Gy JL Sy

OF THE

COUNCIL AND? OF FECERS

OF THE

LOOLOGICAL

SOCIETY

OF LONDON.

Ie,

COUNCIL.

His Grace Tue DuxKe or Beprorp, K.G., F.R.S., President.

Sir JoHN Rosé

IR CIE IN EGID

F.R.S., Vice-President.

Ricuarp H. Burne, Esq., M.A.

Surg.-Genl. Str R. HAvetock |

Cuarugs, G.C.V.O., M.D.

THe Rv. Hon. tree Haru

Cromer, P.C., G.C.B.,

BRADFORD, |

DE Scea

OF |

Srrk WALTER Roper LAWRENCE,

Bt., G.C.L.E.

Sir Epmunp G. Loprr, Bt.,

Vice- President.

Prof. Ernest W. MacBribs,

MeAGy (DSc, oHARsS: /ace-

President.

| Prof. Epwarp A. MuUINcHIN,

GOW, KOR, BIAS, |

Vice-President.

F. G. Dawrrey Drewirr, Esq., |

M.A., M.D.

CHARLES DRUMMOND,

Treasurer.

Sir Epwarp Duranp, Bt., C.B.

FREDERICK GILLerr, Esa.

THE LorpD GLENCONNER.

F. Du Canr Gopman,

D.C.L., F.R.S.

EKsq.,

Ksq.,

M.A., F.R.S., Vice-President.

| P. Cuatmers MircHe.y, Ese.,

M.A., D.Sc., LL.D., F.B.S.,

Secretary.

| W. R. Ocitvir-Grant, Esq.

ApRIAN D. W. Potiock, Ese.

OLpFIELD THomas, Esgq., F.R.S.

AntHOoNY H. WINGFIELD,

Ksaq.

Henry WoopwArp, Ksq., LL.D.,

F.R.S., Vice-President.

PRINCIPAL OFFICERS.

P, Cuatmers MircHetn, M.A., D.Se., LL.D., F.RS.,

Secretary.

Frank E, Bepparp, M.A., D.Sc., F.R.S., Prosector.

R. I. Pococn, F.R.S., F.L.8., Curator of Mammals and

Resident Superintendent of the Gardens.

D. Sera-Surru, Curator of Birds and Inspector of Works.

Epwarp G. BouLencer, Curator of Reptiles.

Prof. H. Maxweiu Lerroy, Honorary Curator of Insects.

Henry G,. Purmmer, F.R.S., M.R.C.S., Pathologist.

Henry G. J. Peavor, Librarian and Clerk of Publications.

JoHn Barrow. Accountant.

W. 4H. Cots, Chief Clerk.

LIST OF CONTENTS.

1913, pp. 339-1104.

EXHIBITIONS AND NOTICES.

The Secrerary. Report on Additions to the Society’s

Menagerie during the month of March 1913 .........

Mr. H. J. Etwes, F.R.S., F.Z.S. Exhibition of the Head

of an Asiatic Wapiti. (Text-figs. 83 & 84.) ............

Mr. R. H. Burne, M.A.,F.Z.S. Exhibition of malformed

Antlers of an Axis Deer (Cervus aitis) ............-.-:::

Mr. E. G. Bousenerr, F.Z.8., Curator of Reptiles. Exhi-

bition of living specimens of the Leaf-Insect

(CMO GPO OUCOHIO)) Acco dab ccaees Bobs sscouedgankucéodeLee

Mr. E. G. Boutencer. Exhibition of a living melanistic

specimen of the Green Lizard (Lacerta viridis) ......

Dr. R. Broom, C.M.Z.S. A new Species of Golden Mole. .

Dr. R. Broom. Exhibition of living specimens of the

female and young of the South African Lizard,

EUUEROUS Chua EmOUCOIGE (Vl IUD, OSIIN))) Gaadoncedoossosasrose

The Secrerary. Report on Additions to the Society’s

Menagerie during the month of April 1913 ............

The Rev. T. R. R. Sraesine, M.A., F.R.S., F.Z.S. Notice

of Prof. F. KE. Schulze’s ‘ Nomenclator Animalium’...

The LrprariaAn. List of dates of publication of the early

parts of the Society’s ‘ Transactions’ ...............0.068.

546

“Page

Mr. R. E. Hotpine. Exhibition of Antlers, Skulls, and

Photographs illustrating Variations in the growth of

the Antlers of Deer. (‘Text-fig. SAS) tector cohen eee 815

Ir. D. Sera-Smira, F.Z.S., Curator of Birds. , Exhibition

of the Egg and Young of the Mikado Pheasant

(COMO DOCSIS TOUBEUD). 95 3ccdacd00e gaa506 do00 00 0bce scene oonconaae 818

The Secrerary. Report on Additions to the Society’s

Menagerie during the months of May, June, July,

August, and September OMe tear ake cto ean eee 1090

The Secretary. Exhibition of the Alimentary Tract of a

young Elliot's Pheasant (Phasianus elliott) ............ 1094

Mi. R. I. Pocock, F.R.S., F.L.S., F.Z.8., Curator of Mam-

mals. Exhibition of a female and three young speci-

mens of the Woolly Opossum (Philander laniger) ... 1094

Mr. R. I. Pocock. Exhibition of two youne Degus

(Cctodonidegus) ern raa. eee ceneee sat Secure dasaysecete ek 1095

Mr. D. Seru-Suira. Exhibition of a spirit specimen of a

young Toucanet (Selenidera maculirostris) showing

the well-developed serrated Heel-pads. (Text-fig.191.). 1095

Mr. C. Tare Reaan, M.A., F.Z.8. Exhibition of a speci-

men of a Deep-sea Angler-Fish, Melanocetus johnsonii

Giinnbly,. 39255 ats sie coe oe ee ere tT ee 1096

My. E.G. Boutencer, Exhibition of a young specimen of

the Matamata Terrapin (Chelys fimbriata) ......4..2.. 1097

Mr. D. Sera-Smirn, Exhibition of a hybrid Black=winged

Peafowl (Text-fig. 192), a hybrid Pheasant, and @

hybrid Jungle- Bowlin eer ne tie tte eee 1098

Mr. R. I. Pocock, Exhibition of Armadillos to show the

Dorsal Glands. (Ci sseotai NEB HOG)). coeocsousbscocoode 1099

The Secrerary. Report on Additions to the Society’s

Menagerie during the month of October 1913 ......... 1108

Prof. H. Maxwent Lerroy, M.A., F.Z.S, Remarks open

the New Insect Efonee Sonu spbudacuna dacHAosEsaossedeoaes 1104

Mr. R. H. Burne. Exhibition of specimens of Foetal

Skeletons prepared by the Beale-Schultze method ... 1104

28.

29.

30.

31.

34,

35,

PAPERS,

. New Species of Rhopalocera from Costa Rica. By

WAMLTAM SCHAUS. EIZ.S5) (Rist WeiiVi.)y-2-5-.25-2.

. A Collection of Fishes made by Professor Francisco

Fuentes at Easter Island. By C. Tare Reean, M.A.,

Le Acer jee {CLEA EBS UD: RRR RB iene alsin Ah Tiber Heap

A Revision of the Fishes of the Genus Auhlia. By C.

Tate Reean, M.A., F.Z.S. (Text-figs. 68 & 69.)

The Affinities of the Antarctic Wolf (Canis antarcticus).

By R. I, Pococn, F.R,S., F.L,S,, F.Z.8,, Curator

OH Wilerairmelis, (UNeexhies) (Oi) nonpAbpasubeooecoanbee se

On the Patella in the Phalacrocoracidee. By Dr. R. W.

EU PEED IE CoV eZ anes (dca [UIXGIs) Wa sa vdaseee sake seeecits-

Experiments on the Metamorphosis of the Mexican

Axolotl (Amblystoma tagrinwm), conducted in the

Society’s Gardens. By E. G. Bourencer, F.Z.S.,

Curator of Reptiles, (Text-figs. 75 & 76,) ...,........

. Contributions to the Anatomy of the Ophidia. By

Surgeon JosepH C. THompson, U.S.N, (Text-figs.

TES Ths) eecene anh 2 voller oie sah oes POV OP a a aa

The Polyzoa of Waterworks, By Srpney F. HARMeEr,

SGD), Une dd4isy | (dallsaMUDCUL, we LpQUE Le) eee nade

The Marine Fauna of British Kast Africa and Zanzibar,

from Collections made by Cyril Crossland, M,A,,

B,Sc., F.Z.8., in the Years 1901-1902. Bryozoa—

Cheilostomata. By ArtHur Wm, Waters, F,[.5.,

F.G.S, (Pls. LXTV.-LXXIII. and Text-figs. 79-82.)

Notes on Albinism in the Common Reedbuek (Cervicapya

arundinum) and on the Habits and Geographical Dis-

tribution of Sharpe’s Steenbuck (Raphiceros sharpei),

By Major J, SrevEnson-Haminton, C.M.ZS, ....0010

Page

399

368

374

382

393

403

414

426

d37

36.

37.

39.

40.

4].

43.

44,

45.

A6.

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—X. On Two Species of Tape-

worms from Genetta dongolana. By Frank KE.

Bepparp, M.A., D.Sc., F.R.S., F.Z.S., Prosector to

themsociety.sa(ext—tlesn8o—04,)\erne re eeseeeeE ee eeraee

Pacific Salmon: An Attempt to evolve something of

their History from an examination of their Scales.

By Jonn ApAmM Mitne. (Text-figs. 95-118.) .........

. Notes on Peripatoides woodwardi Bouvier. By Karu-

TARE NAD DOIN pcos eae elas eee teeta

Field-Observations on the Enemies of Butterflies in

(Cleydlarats 1837 di, Chie IMirwaoin, WING, INIIUSE S54 s¢ocnenae

On the South-African Pseudosuchian Huparkeria and

Allied Genera. By R. Broom, M.D., D.Sc., C.M.Z.S.

(TIE) DDO. Viral BD.O.D. Gaia oe Shean peabaiaanas po5cocgace codon

On some Cases of Blindness in Marine Fishes. By

SG yea Deel Bo C1 ba ON Nseries oe Precine ebh A era ec

2. Notes on Turacin and the Turacin-Bearers. By

Sir ARpHpR JE CHrurcH KUCAVCOL SROs) aaeeeereeee

Observations on the Anatomy of the Shoe-bill (Bale-

niceps rex) and allied Birds. By P. CHALMERS

Mircueni, M.A., D.Sc., LL.D., F.R.S., Secretary to

the Society. (Pls. LXXX.-LXXXIII. and Text-

LS Sel O13.) Cee crctets wetioceeceo ah et see eee EERE ae

The Classification and Phylogeny of the Calcareous

Sponges, with a Reference List of all the described

Species, systematically arranged. By Arruur Denby,

D.Se., F.R.S., F.Z.8., and R. W. Harotp Row, B.Sc.,

HES. -(@ext=fig. 133.) 02s a.en foi ngaie Wo teenie ees

The Transvaal Race of the Cape, or Khama, Hartebeest.

By R. Lyprxxker, F.R.S., F.Z.8. (Text-fig. 135.) ...

On a Collection of Mammals from the Inner Hebrides.

By G. E. H. Barrerr-Hamitton, F.Z.S., and Marrin

A. C. Hinton. (Pl. LXXXIV. and Text-figs. 136-

BGP) sec ca We gndsce oi dokets scot BEE eee eee

Page

D49

572

611

613

619

634

639

644

704

818

47.

48.

49,

5].

vil

Some Miocene Cirripedes of the Genera Hevelasma

and Scalpellum from New Zealand. By Tuomas H.

Wiruers, F.G.S. (Pls, LXXXV. & LXXXVI. and

Memes a OAL) Clea saecijcmacscieen vide wesctoed valnceees

Observations on Osteomalacia in the Zoological Col-

lections of Manchester and Cleveland. By T. Wrneate

Topp, M.B., F.R.C.S. (Els. LXXXVIT.-LXXXIX.)

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—XI. On a new Tapeworm

from Wdienemus. By FRANK EK, Brpparp, M.A.,

D.Se., #.R.S., F.Z.8., Prosector to the Society. (Text-

Heese oe MEA ME ae ac esas Sarai As tse else anaes Sela

. The Brain and Brain-Case of a Fossil Ungulate of the

Genus Anoplotherium. By R. W. Paumer, M Sc.

(Manchester), Research Fellow in Zoology, University

College, Reading. (Text-figs. 150-157.) ..,..,.........

The Fossil Crinoils referred to Hypocrinus Beyrich.

bya 105 Avo letemenaic. | WIS, IDES @ye Sita Nasi N/A iS

(PIX and Mext=fiess VOSNG602) 5 eet eee se cnc cces-

- On Freshwater Decapod Crustacea (Families Potamo-

nide and Palemonide) collected in Madagascar by

the Hon. Paul A. Methuen. By W. T. Catman,

D.Sc., F.Z.8. (Pls. XCI. & XCII. and Text-fig. 161.)

A New Trematode of the Genus Lechriorchis from the

Dark Green Snake (Zamenis gemonensis). By Marre

Y. Lesour, M.Sce., Assistant Lecturer and Demon-

strator in Zoology, Leeds University. (Pl, XCIII.) ,

. Cirripedes from the Cenomanign Chalk Mar] of Cam-

bridge, By Tsomas H, Wiruers, F.G.S. (Pls,

. Batrachiderpeton lineatum Hancock & Atthey, a Coal-

Measure Stegocephalian. By D, M, 8. Warzson,

M.Sce., Lecturer on Vertebrate Paleontology in Uni-

versity College, London, (Pls. XCVI, & XCVII,

aide Dextre saat OL MOMs) caccecnutsasesecse meer niseieriens ;

. On two New Actinians from the Coast of British

Columbia. By J, Puayrarr McMourricn, C.M,Z8,

as ce ONY LETS) “oopane oooenbupvonunseonenee on popapne pevevegvenes

Paga

840

895

878

894

914

933

937

949

963

Vill

Page

57. Sponges in Waterworks. By W.N. Parker, Ph.D.,

F.Z.S., Professor of Zoology, University College,

Wand iity A fugetcith..d Jaded Riehoretnareacecnee sean eae 973

58, A Revision of the Cyprinodont Fishes of the Subfamily

Peciliine. By C. Tare Recan, M.A., F.Z.8. (Pls.

NC X= Cian Mext-figsilGS= ids) eeeceneeer sesso: Sa

D9. On a Collection of Batrachians and Reptiles made by

Dr. H. G. F. Spurrell, F.Z.8.,in the Choco, Colombia.

By G. A. Boutencer, F.R.S., F.Z.8. (Pls. CII.=

CWWaltisicvand Wext- tose (4 lis) i cepecee sees cere eck 1019

60. The Peroneal Muscles in Birds. By P. CHALMERS

Wilewornsine, = Wl oNGy IDES, IbIELIDL, GIRS, | Arsen

Secretary to the Society. (Text-figs. 179-190.) ...... 1039

61. On the External Characters and Biology of Bryde’s

Whale (Balenoptera brydei), a new Rorqual from

the Coast of South Africa. By Oran OLsEN,

Zoological Laboratory, Christiania University. (Pls.

CTE FOTN) so cis Sebi pa es Werder gael wae dene) ok eee 1073

Alphabetical Mist) of Contributors) 2. «ssa. sseeseseeteneees 1X

iNew. Generic WBC Siem. o.jshesr scrarccken tle cstroc Seeieee score ops xvi

TroGlebe Git SONGTTUAUINS INEWINGS — dacooonpoonscodaonodonodoodobodeodoedse X1X

indexvotqlilustrabvonssicats ccecee tes coe ie eeioe ees xlii

ALPHABETICAL LIST

OF THE

CONTRIBUTORS,

With References to the several Articles contributed by each.

(1913, pp. 339-1104.)

Page

Barrett-Hamizton, G, E. H., F.Z.S., and Hryton, Martin

A.C.

On a Collection of Mammals from the Inner Hebrides.

(Pb TOO ein) eesti USJONGISh) bcoeeseoncoseceonacas 821

Batuer, Francis A., M.A., D.Sce., F.R.S., F.Z.8.

The Fossil Crinoids referred to Hypocrinus Beyrich.

CeEEXCr andi Dextre OS — 100s) esacacncaealaeiecdseete ace 894

BEDDARD, FRANK E., M.A., D.Sc., F.R.S., F.Z.8., Prosector

to the Society.

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.

X. On Two Species of Tape-worms from Genetta dongo-

aiipoiig. (QUEERS, (35 8))) pee copqopgpabbecsoo se odenconconnogd: 549

XI. On a New Tapeworm from @dicnemus. (Text-

fos yA eA On ea Sind Sate inher Bann len ccce Salada cra dcte <a 86]

Page

Bovutencer, Epwarp G., F.Z.8., Curator of Reptiles.

Experiments on the Metamorphosis of the Mexican

Axolotl (Amblystoma tigrinum), conducted in the Society’s

Gardens: an (Next-fisss 1 160s) menses neh n tees tanner 403

Exhibition of living specimens of the Leaf-Insect

(CECH OUTD, CELIO OINTEDN). "achboebennaoagoosennscensebaebocesadee cc: 546

Exhibition of a living melanistic specimen of the Green

JINR (OL RIGIG, CHPUCHIS)) aoa oabepocpbadaedeadoooosoonbsacdoon oso: 546

Exhibition of a young specimen of the Matamata

UVeroenyjarin (Claas jUMULPOGTD) ccocascooossps0q0n0cacadbaen00900350 1097

Bounrencer, Grorce A., F.R.S., F.Z.S.

On a Collection of Batrachians and Reptiles made

by Dr. H. G. F. Spurrell, F.Z.8., in the Choco, Colombia.

(Pls. CII.-CVIII. and Text-figs. 174-178.) .,....,..,... 1019

Broom, Rospert, M.D., D.Sc., C.M.Z.8.

A new Species of Golden Mole..............-..:cscssseceoes 546

Exhibition of living specimens of the female and young

of the South African Lizard, Zonurus giganteus. (PI.

NBD. BES) anne ere Ferien thn Vals AAMT ae Aad ond ohbonasbaa une 548

On the South-African Pseudosuchian Huparkeria and

Allied Genera. (Pls. LX X VX XO) eee ee 619

BULLEN, G. E.

On some Cases of Blindness in Marine Fishes _......... 634

Burne, Rrenarp H., M.A., F.Z.S.

Exhibition of malformed Antlers of an Axis Deer

(CCE CD TIS) Danna eHn AN PENEERB HORM E an aco chin Gia jnbquanantandoc 545

Page

Exhibition of specimens of Foetal Skeletons prepared

by the Beale-Schultze method ...............sseeeeeeeeeseeees 1104

CaLMAN, Witii1aM T., D.Sc., F.Z.8.

On Freshwater Decapod Crustacea (Families Pota-

monidz and Paleemonidz) collected in Madagascar by the

Hon. Paul A. Methuen. (Pls. XCI. & XCII. and Text-

in, UG pesbdaddudeschsnedescodabosh aad satpocooceoesr suceecnecseatnc 914

Cuurcsa, Sir Artuur H., K.C.V.O., F.R.S.

Notes on Turacin and the Turacin-Bearers............... 639

Drenpy, ArtHuR, D.Sc., F.R.S., F.Z.8., and Row, R. W.

Haro, D.Sc., F.L.S. :

The Classification and Phylogeny of the Calcareous

Sponges, with a Reference List of all the described Species,

systematically arranged. (Text-fig. 133.) .................. 704

Ewes, Henry J., F.R.S., F.Z.8.

Exhibition of the Head of an Asiatic Wapiti. (Text-

HUGS HSA I0L) OAn) erence dnecgec madsen acecneede ce stewigwcieae aetna 542

Fryer, J. C. F., M.A., F.E.S.

Field-Observations on the Enemies of Butterflies in

(CES TINOT ONS Sommer bred dee TOsOe acc Ine BOGE erick Ctra Tir LOCC Cees ORES 613

Happon, Miss KATHLEEN,

Notes on Peripatoides woodwardi Bouvier ............. 611

Hamitron, G. EK. H. Barrerr-. See Barrerr-HAmItroy,

Gals EH.

Hamiuron, J. SrevENsSON-. See SrrveNsoN-HAmILtoN, J.

X1L

Harmer, Sipney, F., Se.D., F.R.S., F.Z.S.

The Polyzoa of Waterworks. (Pls. LXII. & LXITI.). 426

Hinton, Martin A. C., and Barrerr-Hamitron, G. E. H.,

E.Z.S.

On a Collection of Mammals from the Inner Hebrides.

CEI IDXOXGXeVE and Mext=neseek3 Ota. a) ee eEee ee nae etary 821

Ho.prine, R. EK.

Exhibition of Antlers, Skulls, and Photographs illus-

trating Variations in the growth of the Antlers of Deer.

(Dext=fig. Moa) cise. dates oan, seinen oes cig aber enlace 815

Lesour, Miss Marte V., M.Sc,

A New Trematode of the Genus Lechriorchis from the

Dark Green Snake (Zamenis gemonensis), (Pl, XCIII.). 933

Lerroy, Prof. H. Maxwett, M.A., F.Z,8., Hon, Curator of Insects.

Remarks upon the New Insect House..,....,,,........... 1104

LYDEKKER, RicHarp, F,R$,, F.Z.8,

The Transvaal Race of the Cape, or Khama, Harte-

beest; Bext-fig, 135.) :cscsd.. dads scutes tae eee eee eee 818

McMoraicu, J. Prayrarr, C,M.Z.S,

On two New Actinians from the Coast of British

Columbia, «(PE XC VILL icc eee Cee eee ens 963

Mine, Joun ADAM.

Pacific Salmon: An Attempt to evolve something of

their History from an examination of their Scales. (Text-

eS eo) a Speen SME MRP ago Hrs Sudgaaconatcadon 572

xiil

’ Page

Mireneuy, P. Caatmers, M.A., D.Sc., LL.D., F.R.S., F.Z.S.,

Secretary to the Society.

Report on Additions to the Society’s Menagerie during

Taeemomulmotevlanche MOMS ssrausetases ecisstccwetcrecse seas 542

Observations on the Anatomy of the Shoe - bill

(Baleniceps rex) and allied Birds. (Pls. LXXX.-

LXXXIII. and Text-figs. 119-132.) ..................-.e 644

Report on Additions to the Society’s Menagerie during

UME MAL OLIN ON PADIEM MON ed therserasye s stisicistensints otras clerieletate ajtaietasier 813

The Peroneal Muscles in Birds. (Text-figs. 179-190.) 1039

Report on Additions made to the Society's Menagerie

during the months of May, June, July, August, and

Be pbemaly err USM AIG ae ee. ee SUI athe de Rlaateened aa torars 1090

Exhibition of the Alimentary Tract of a young Elliot’s

Bheasaintg (hasvaneuscelliote)i-saeses anes esse eee eee ee 1094

Report on Additions to the Society’s Menagerie during

iln® sonore Or Crewe orere OMS? ceccacogssacnpaco soedaeaccsesessecac 1103

OLSEN, ORJAN.

On the External Characters and Biology of Bryde’s

Whale (Balenoptera brydei), a new Rorqual from the

coast of South Africa. (Pls. CLX.-CXIIT.)

Pater, R. W., M.Se.(Manch.),

The Brain and Brain-Case of a Fossil Ungulate of the

Genus Anoplotherium. (Text-figs, 150-157.) ve... eee. 878

ParkKER, Prof. W. Newton, Ph.D., I'.Z.S.

Sponges in Waterworks

Pravor, Henry G. J., Librarian to the Society.

List of dates of publication of the early parts of the

Society’s ‘ Transactions’

X1V

Page

Pocock, Reeinatp I., F.B.S., F.L.S., F.Z.S., Curator of

Mammals.

The Affinities of the Antarctic Wolf (Canis antarc-

dics). (Rext=figs MOS ( ANE. ca ja. sch epee eee eee neer te. eecr ance 382

Exhibition of a female and three young specimens of

the Woolly Opossum (Philander laniger) — ..........ece cece 1094

Exhibition of two young Degus (Octodon degus) ...... 1095

Exhibition of Armadillos to show the Dorsal Glands.

(Mextetigs NOS S195) is iiaieeeencortuuatine scone cceren eee cre: 1099

Reean, C. Tats, M.A., F.Z.S8.

A Collection of Fishes made by Professor Francisco

Fuentes at Easter Island. (Pls. LV.-LX.) ............... 368

A Revision of the Fishes of the Genus. Kuhlia.

(Aes ces tata lela (0) )) e Rada cing MogeBaonaaaesdscoddonadessaeasGoon 374

A Revision of the Cyprinodont Fishes of the Sub-

family Peeciliine. (Pls. XCIX.—CI. and Text-figs.

Ios Nfe)8) Waser ota nardndnodeadanccesac SPE SID oe Og

Exhibition of a specimen of the Deep-sea Angler-Fish,

MUNG POCA MS NOMISO MID EMD tin phsq0bbs0a0006000s60a0e0be0G0D00. 1096

Row, R. W. Harotp, B.Sc., F.L.S., and Drenpy, ARTHUR,

ID S55 ISIS) NYAS

The Classification and Phylogeny of the Calcareous

Sponges, with a Reference List of all the described

Species, systematically arranged. (Text-fig. 133.)......... 704

ScHaus, WiLuiAM, F.Z.S8,

New Species of Rhopalocera from Costa Rica. (Plates

1,1 I gs ae Mrs 8 Ae A 339

Sura-Suiru, Davip, F.Z.S., Curator of Birds.

Exhibition of the Egg and Young of the Mikado

Pheasant (Calophasis mikado) ........1sesereeeeete nsec een ees

Exhibition of a spirit specimen of a young Toucanet

(Selenidera maculirostris) to show the well-developed

Semrated Heel pads.) (Mext-ton NON) oor. secaenentectmecee

Exhibition of a hybrid Black-winged Peafowl] (Text-

fig. 192), a hybrid Pheasant, and a hybrid Jungle Fowl .

Suuretpr, Roperr W., M.D., C.M.Z.S.

On the Patella in the Phalacrocoracide. (PL. Xe

Sairn, D, Seru-. See Sera-Smiru, D.

STEBBING, The Rev. T. R. R., M.A., F.RB.S., F.Z.S.

Notice of Prof. F. E. Schulze’s ‘ Nomenclator Ant-

malium ’

Pee eee ewe ee ea eee eee e eres ress sees eseesseseessssaessestsssose

SLEVENSON-Hamiuron, Major J., C.M.Z.S.

Notes on Albinism in the Common Reedbuek (Cervi-

capra arundinum) and on the Habits and Geographical

Distribution of Sharpe’s Steenbuck (Raphiceros sharpei) .

THompson, Surgeon Josepu C., U.S.N.

Contributions to the Anatomy of the Ophidia. (Text-

LIS SHRM OD Sa) BAI. Menthe toeenemade sah aNen saute uctauncuedaa

Topp, T. Wrineate, M.B., F.R.C.S.

Observations on Osteomalacia in the Zoological Col-

lections of Manchester and Cleveland. (Pls. LX XXVIT-—

TE XSNOXSIENG nae

ee ee i ee oe ry

Page

818

1095

1098

393

814

537

Al4

XV1

Waters, Artuur W., F.L.S., F.G.S.

The Marine Fauna of British East Africa and Zanzibar,

from Collections made by Cyril Crossland, M.A., B.Sc.,

F.Z.S., in the Years 1901-1902. Bryozoa—Cheilostomata.

(Pls. LXTV.-LX XIII. and Text-figs. 79-82.) ............ 458

Watson, D. M.8., M.Sc.

Batrachiderpeton lineatum Hancock & Atthey, a Coal-

Measure Stegocephalian. (Pls. XCVI. & XCVIT. and

Pext-figs. [62-16 0.) cease rae eer ctr ree etedste screenees 949

WITHERS, THOMAS H., E:G:S:

Some Miocene Cirripedes of the Genera Hexelasma

and Scalpellum from New Zealand. (Pls. LXXXV. &

IDXOOXOVS andi Vext-figss 139d 140.) ree eset eeeeeeeee 840

Cirripedes from the Cenomanian Chalk Marl of Cam-

lover Fegen, (Ue CONAN 05 ACORVG)) So gecn ondoadssocsocsonacosntoc 937

XVil

NEW GENERIC TERMS

PROPOSED IN THE PRESENT VOLUME (pp. 339-1104).

Page | Page

Ascute (Spongiz) .............:. 729 Pamphoria (Pisces) ............ 1003

Brachyrhaphis (Pisces) ...... 997 Pamphorichthys (Pisces) 10038

Browniella (Reptilia)............ 627 Peciliopsis (Pisces) ............ 966

Corrachia (Insecta) ............ 351 Priapella (Pisces) ............... 992

Diplopylidium (Vermes) ...... 569 Priapichthys (Pisces) ......... 991

Epinosis (Insecta) ..............- 366 Pseudopecilia (Pisces) ...... 995

Kugonodzum (Vermes)..... ... we Styriodes (Insecta) ............ 361

Girellops (Pisces) ............... 369 Syeute (Spongia) ............... 763

Hebius (Reptilia) ............... 424 Teichonopsis (Spongiz) ...... 761

Kuarrhaphis (Spongiz) .. ...... 780 Titanolepas (Crustacea) ....... 943

Leptorhaphis (Pisces) ......... 998 Uteopsis (Spongiz) ............ 766

Leucopsila (Spongiz) ......... 7795 Zeugmatolepas (Crustacea)... 938

Proc. Zoo. Soc.—1913.

tainty

Jy)

TT tne CAM:

: ahh! i a 4 ac ‘ i b Lets

INDEX

SCIENTIFIC NAMES,

Acanthistius

cinctus, 868, 369.

Suscus, 868, 373.

Acanthopacelus

bifurcus, 1006.

melanzonus, 1007,

1008.

reticulatus, 1008.

Acauthorbynchus,

1065.

Achradoecrinus, 896-

899.

Achramorpha, 758, 766,

798.

glactalis, 765.

grandinis, 765.

nivalis, 76d.

schulzei, 765.

truncata, 765.

Acineta, 428.

Acleropogon

piceus, 618.

Acmepteron

cinerascens, 356, 357.

poasind, 896, 367.

Acridotheres

melanosternus, 1091.

Acropeecilia, 1014.

tridens, 1018.

Actias

selene, 618.

Actinia

clavus, 969.

cylindrica, 966, 972.

Actitis

hypoleucus, 1054.

Adeona

Joliacea fascialis, 532.

Adeonella, 492, 519,

528.

contorta, 504, 529.

platalea, 463, 501, 514,

529-532, 537.

polystomella, 530, 531.

| Adeonella *

(Laminopora)

contorta, 529.

Adeonellopsis, 528.

crosslandi, 463, 531,

532, 537.

distoma, 531, 532.

imperforata, O31.

subsulcata, 531.

/Kchmophorus

major, 6O4.

| Agialitis

nivosa, 861.

AKigotheles, 1065.

nove-hollandie, 1062.

/Eluredus

melanocephalus, 1065.

Aiyuorea, 970.

forskalii, 967.

Actea

anguina, 462-464, 532.

recta, 463, 464.

truncata, 462, 465,

Aithopyga

seherie@, 1091.

Aétosaurus, 629, 630.

ferratus, 624.

galericulata, 1047.

sponsa, 1047, 1048.

Afaro, 979.

acutiventralis, 981.

amazonum, 981, 990.

cultratus, 981.

Agalychnis

calcarifer, 1023,

1038.

spurrelli, 1024, 1025,

1038.

Alea

torda, 1056.

Alcedo

astaticus, 1060.

Alcyoncellum, 787.

Aleedo

bengalensis, 1060,

ispeda, 1060.

gelatinosum, 746,

787.

Aleyonella

articulata, 441.

fungosa, 450.

Aleyonidium

duplex, 488.

Alopoglossus

coptt, 1038.

fuliginosa, 1034.

Alticus

striatus, 372.

variolosus, 373.

Alytes

obstetricans, 1025,

Amabilia, 571.

Amazilia

felicie, 1063.

Amblystoma

tigrinum, 403-413.

Ameiva

bridgesti, 1032.

festiva, 1033.

| Ampheristocrinus, 911.

| Amphisbeena

fuliginosa, 1094.

Aimphiute, 754, 798.

paulini, 755.

Amphoridium, 787.

viridis, 787.

Amphoriscus, 781, 798,

804.

buceichii, 782, 790.

chrysalis, 782.

cyathiscus, 782, 799.

cylindrus, 782.

elongatus, 782.

flamma, 752.

gregorti, 782.

kryptoraphis, 782.

Amphoriscus

murmanensis, 733.

oblatus, 711, 782.

ovipurus, 782.

poculum, 752.

semoni, 782.

testiparus, 782.

thompson, 753.

urna, 782.

Amphorula, 787.

solida, 787.

Anadia

oceliata, 1033.

vittata, 1033, 1038.

Anamixilla, 758, 786,

798.

torresi, 766.

Anampses

ceruleopunctatus, 372.

pulcher, 371-373.

Anaschisma, 957.

Anastomus, 688, 690.

Anchisaurus, 629, 630.

Ancylus, 427.

Anodonta, 428.

Anolis

breviceps, 1031, 1038.

Jasciatus, 1030.

godmani, 1031.

macrolepis, 1032.

maculiventris, 1030.

notopholis, 1032.

princeps, 1031.

Anoplotherium, 879-

882, 886-888, 890-

893.

Antedon, 903, 904.

Anthropoides

virgo, 1053.

Anthropopithecus

troglodytes, 1090,

Aphaneramma, 956.

Aphelocoma

sordida, 1091.

Aphroceras, 768, 776,

798.

aleicornis, 777, 789.

asconoides, 729.

cespitosa, 777.

cataphracta,

777.

cliarensis, 777.

corticata, 777.

elongata, 777.

ramosa, TO4,

sericatum, 'T56.

syconoides, 764.

Apodemus

hybridensis, 835-837.

sylvaticus sylvaticus,

835, 837.

alu;

Appias, 616.

sp-, 615, 618.

albina, 614, 618.

paulina, 614, 617, 618.

Aprosmictus

cyanopygius, 1008.

Aptenodytes, 1044.

pennantii, 394.

Apteryx, 1043.

Aquila

verreauxt, 1O49,

Arachnechthra

zeylonica, 1066.

Aramides

chiricote, 1052, 1104.

ypecaha, 1052.

Aramus

scolopaceus, 1053.

Archegosaurus, 956.

Archispirostreptus

nitidus, 1093.

Archonias

dismorphites, 356.

intermedia, 3595,

367.

nigrescens, 396.

Arcoscalpellum, 850,

853.

Ardea, 656, 690, 691,

695, 694, 696-699,

703.

egretta, 1045.

herodias, 1045.

— lessoni, 1045,

1091.

purpurea, 1045,

Ardetta, 674.

minuta, 1045.

Arginia

sp., 614, 618.

Argus

giganteus, 1051.

Artamus

Suscus, 613, 615.

leucogaster, 1066.

Arthropoma

cecillii, 462, 508.

Artynas, 787.

compressus, 787.

villosum, 746.

Artynella, 787.

compressa, 787.

Artynes, 787.

compressa, 787.

Artynium, 787.

compressum, 787.

Artynophyllum, 787.

compressum, 787.

Asarcia

variabilis, 1055.

Ascaltis, 717, 787.

INDEX OF SCIENTIFIC NAMES.

Asealtis

botryoides solanderii,

720.

canariensis,

787.

cerebrum, G24.

— decipiens, 725,

compacta, "724.

darwinii, 725.

— caroli, (24.

gegenbauri, 725.

— charybdea, 724.

goethet, 725.

lamarckii, 726.

— agassizti, 724.

solanderti, "725.

Aseandra, 717,

787.

angulata, 721.

hotrys, T21.

complicata ameboides,

720.

contorta, 721.

conulata, 726.

corduta, 721, 787.

densa, 721.

echinoides, 722.

falcata, "17, 720, 722,

788, 790.

hermesti, "722.

licherkiihnit. 723.

nirabilis, 761.

nitida, 723.

panis, 723.

pinus, 723.

reticulatum, 728.

— reticulata, 723.

sertularia, 723.

tenuis, 723.

variabilis, 723.

— arachnoides, 721.

— cervicornis, 721.

— confervicola, 721.

— hispidissima, 722.

Ascetta, 717, 788.

blanca, 724.

— phillipina, 726.

celathrus, 725.

coriacea himantia,

726.

flewilis, 725.

macleayt, 726.

primordialis, 726, 730,

788.

— dictyoides, 725.

— loculosa, 726.

— poterium, 726.

— protogenes, 726.

procumbens, 726.

sagittaria, 727.

sceptrum, 727.

724,

Ascetta

spongiosa, 727.

vesicula, 727.

Ascilla, 7388.

coriacea, 725.

gracilis, 725, 788.

— convallaria, 725.

— japonica, 726.

Ascometra, 788.

primordiale, 788.

Ascortis, 788.

clarkit, 721.

corallorhiza, 721.

Sragilis, 722.

horrida, 722, 788.

lacunosa, 72%.

Asculmis, 758.

armata, 721, 788.

Ascuris, 788.

arrecifé, 788.

Ascute, gen. nov., 719,

29% 798; 799!

asconoides, 729.

uteoides, 710, 718,

G29.

Ascyssa, 717, 719, 767,

798, 799.

acufera, 729.

troglodytes, 729.

Asellus, 426, 451, 4384,

437.

aquaticus, 427.

Asio

otus, 1061.

Aspatha

gularis, 1060.

Aspidelectra, 461.

Aspidostoma

giganteum, 487.

Asterias

rubens, 1104.

Astrosclera, 788.

willeyana, 788.

Ate

canace, 359, 367.

Athene, 1062.

noctua, 1061.

Atheris, 1034.

Atractus

iridescens, 1035.

multicinctus, 1035.

Aulacorhamphus

prasinus, 1063.

Auloplegma, 7&8.

haeckeli, 721.

loculosum, 738.

Aulorhiza

intestinalis, 726, 788.

Baeria, 768, 780, 798, 803.

INDEX OF SCIENTIFIC NAMES.

Baeria

ochotensis, 779.

Baleniceps, 645-703.

rex, 644, 650, 701-708,

1039, 1045, 1092.

Balxnoptera

acutorostrata, 1076,

1077, 1085, 1086.

borealis, 1073, 1074,

1076-1030, 1083,

1085-1087, 1089,

1090.

brydei, 1074-1977,

1079, 1080, 1082-

1087, 1089, 1090.

musculus, 1076, LO89.

physalus, 1076, 1077,

L085.

Balanus, 842, 846.

callistoderma, 847.

corolliforneis, 847.

evermanni, S47.

hirsutus, 847.

hockianus, 847, 848.

psittacus, 847.

Balearica

chrysepelargus, 1053.

Basiliscus

americanus, 1032.

galeritus, 1032,

Bathus

hoitentotta, 1093.

Bathystethus

cultratum, 373, 374.

orientale, 379.

Batrachiderpeton, 949,

955, 956, 958-961.

lineatum, 950, 951,

953, 954, 962.

Batrachosuchus, 956.

Beania

hirtissima, 467.

hyadesi, 487.

intermedia, 467.

magellunica,

530.

mirehilis, 462, 467.

quadricornuta, 467.

spinigera, 462, 467.

Belodon, 623, 629.

Belonesox, 979.

467,

belizanus, 977, 990,

991.

Bematiscus

transvaalensis, 547,

548.

trevelyani, 547.

villosus, 546-548.

Bicellaria, 476.

chuakensis, 462, 467,

5384.

XX1

Bicellaria

ciliata, 468, 470.

Bicidium, 970.

equoree, 967-969, 972.

clavus, 969.

parasiticum, 969, 972.

Bifaxaria

abyssicola, 528.

corrugata, 527.

denticulata, 527.

levis, 528.

minuta, 528.

papillata, 528.

reticulata, 527, 528.

submucronata, 527.

vagans, 463, 527, 528,

582, 533.

Biflustra

armata, 486.

Bithynia, 437.

tentaculata, 427.

Bithynis

gaudichaudii, 930.

hildebrandti, 926, 928,

930.

Biziura, 393.

lobata, 396.

Boa

occidentalis, 1094.

Bolla

cylindus, 359.

machuca, 359, 367.

sodalis, 359, 367.

Boobon

fuliginosus, 1092.

Botaurus, 652, 699.

stellarts, 1045.

Bothriceps, 956, 957.

Bothrodytes, 424.

Bothrops

acrochordus, 1037,

Boulengerina, 374.

Brachypternus

auranticus, 1063.

Brachyrhaphis, gen. nov.,

997, 980.

rhabdophora, 990, 997,

998.

Bradypus

tridactylus, 1094.

Brettia, 484.

australis, 465.

longa, 465,

tropica, 462, 465, 532.

Browniella, gen. noy.,

627.

africana, 627, 628, 633.

Bubalis

caama, 818, 819.

— selhornet, 820, 821.

— typica, 821.

Xxll

Bubalis

cokei, 819.

Bubo, 151.

lacteus, 1061.

maculosus, 1061.

maximus, 668, 1061.

Bucorvus, 670, LO61.

Buffonella, 505.

Bufo

hematiticus, 1022.

hypomelas, 1022, 1038.

marinus, 1022.

typhonius, 1022,

Bugula, 459, 470, 522.

avicularis, 479.

bicornis, 476.

caliculata, 469, 479.

capensis, 471, 472.

dentata, 472.

gracilis, 472.

mirabilis, 472.

murrayanda, 479.

neritina, 472.

— minima, 462, 471.

— ramosa, 472.

— rubra, 472.

— tenuata 472.

reticulata, 472.

— unicornis, 472.

robusta, 462, 471, 534.

versicolor, 472.

Bugulopsis

peachit, 473, 474, 584.

Buthus

gibbosus, 542.

Butleria

faula, 360.

lalage, 360.

lethea, 360, 367.

lysis, 360, 367.

Butorides

cyanurus, 650.

Caberea, 474, 475, 481.

crassimarginata, 480.

darwinit, 477,

ellisit, 534.

retiformis, 479, 480.

Caberiella, 475.

Cacatua

sulphurea, 1058,

Cecilia

intermedia, 1020, 1034.

isthmica, 1020.

nigricans, 1022,

pachynema, 1020, 1022,

palmeri, 1021.

tentaculata, 1020.

Caica, 1059.

melanocephala, 1058,

Caiman

selerops, 1030.

Calantica, 937, 938, 942,

947.

(Titanolepas) — ¢uber-

culata, 943, 948,

Calcispongia, 789.

compressa, 789.

Calodromas, 1051.

elegans, 1050.

Calophasis

elliott, 1052, 1098.

mikado, 818. 1094.

Caloporella, 484.

Calopsittacus

nove-hollandie,

1059.

Calpidium, 483.

Calwellia, 465.

Campylorhynehus

unicolor, LOG.

Cancroma, 645, 649, 652,

674, 689-691, 693-

696, 698-700, 703.

cochlearia, 699.

zeledoni. 1045, 1091.

Canda, 473, 474.

arachnoides, 480.

Jfossilis, 480.

retiformas,

481, 534.

tenuis, 480.

Canis

antarcticus, 382-893.

anthus, 384, 386, 393.

aureus, 384.

azare, 392.

CUNCTLLOTUS,

390, 333.

familiaris, 384,

Fulvipes, 390,

gracilis, 390.

Jubatus, 383, 384.

latrans, 382-393.

462, 479,

lupaster, 386, 9890,

392.

lupus, 384, 390, 393.

mesomelas. B84.

mnicrotis. 390.

occidentalis, 884, 390.

1058, |

382-384, |

pallipes, 386, 390, 392,

393.

parvidens, 390.

rudis, 390.

sclatert, 390, 391, 1108.

urostictus, 390.

vetulus, 383, 384, 393.

(Nyctereutes) pro-

cyonoides, 391.

Capitosaurus, 956.

Caprimulgus, 1065, 1072. |

INDEX OF SCIENTIFIC NAMES. |

Caprimulgus

europeus, 1062.

Caranx.

cheilio, 370,

guard, 3710.

Cariacus

leucurus, 817.

Cariama

burmeistert, 1094.

cristata, 1054.

Carphibis

spinicollis, 654, 1047.

Carpophaga

rufiguia, 1056.

Carystus

commodus, 366.

subrufescens, 865, 367.

Castor

canadensis, LOL.

Casuarius

keysseri, 542.

scluteri, 1091.

uniappendiculatus,

1043.

Catantica

(Scillelepas), 947, 948.

Catargynnis

dryadina, 342, 367.

Catenaria, 465.

diaphana, 462,

485, 532.

lafontii, 462, 481, 482,

489.

parasitica, 489.

Catenicella

contei, 485.

continua, 485.

cornuta, 483, 484.

delicatula, 488.

elegans, 484.

fusca, 483.

internodia, 484, 485.

perforata, 453,

seplentrionalis, 48d.

taurina, 483.

umbonata, 433.

Jathartes, 661, 1069.

aura, 655, 1048.

Cathrina, 717.

Catopsilia, 616.

crocale, 615.

pyranthe; 614, 615,

Cellaria, 461, 493.

australis, 495, 496.

bicornis, 495.

contiqua, 497.

dennanti, 496. 497.

divaricata, 495.

dubia, 42h.

Jistulosa, 490, 496.

gracilis, 490, 495-497,

482,

Cellaria

gracilis tessellata, 492, |

495, 497, 535.

hirsuta, 496.

Jullieni, 496.

magnifica, 497.

malvinensis, 495-497.

oculata, 490.

rigida, 495, 496.

salicornia, 494.

salicornioides, 497.

tenella, 490.

tenuirostris, 495, 497.

variabilis, 495, 496.

wandeli, 496.

wasinensts, 462, 495—

497, 533.

Cellarinella, 528.

Cellepora, 461, 529.

albirostris, 522.

— typica, 522.

avicularis, 5V8.

cidaris, 521.

ciliata, 509.

columnaris, 521.

conica, 508.

coronopus, 508-510.

eatonensis, DO8.

evexd, 508.

hyalina, 509.

incrassata, 509.

megasoma, 508.

pumicosa, 509, 510.

ramulosa, 509, 510.

redoutii, 508.

rota, 510,

sardonica, 509.

simonensis, O08.

spongites, 509.

Cellularia, 461. 493.

diplodidymoides, 492.

spatulata, 478.

tubucellaria, 494.

Centropomus

rupestris, 375.

Ceraterpeton, 960, 962.

galvani, 958, 959, 962. |

reticulatum, 98d, 959,

961.

Cercopithecus

randti, 542.

Cereopsis

nove-hollandie, 1047.

Cerianthus

membranaceus, 966.

Cervicapra

arundinum, 537.

Cervulus

muntjac, 817.

Cervus

axis, 040, 817.

INDEX OF SCIENTIFIC NAMES.

Cervus

canadensis, 816.

canadensis siberica, 545.

dama, 817.

elaphus, 543, 545, 816.

eustephanus, D4).

unicolor, 817.

Ceryle

alcyon, 1060.

americana, 1060.

anda, 1060.

maxima, 1060.

Ceyx

rufidoris, 1060.

Charadrius

pluvialis, 1059.

Charis

chrysus, 352.

subressellata, 351, 367.

turrialbensis, BOK

367.

Chasmorbynchus

nudicollis, \065.

Chauna, 1042, 1043.

chavaria, 1040, 1041, |

1047.

cristata, 1092.

Cheilio

inermis, 371.

Cheilopora, 515,

Chelys

Sinbriata, 1097.

Chilopora, 515.

Chionis

alba, 1055.

Chironomus,

459.

Chizerhis, 641.

Chlidonia

433, 45C-

cordiert, 462, 489, 490, |

492, 583.

Chloéphaga

tornata, 1047.

Choanoteenia, 564.

coronata, 861.

Chorizopora

brongniarti. 507.

Chrysaora, 969.

Chrysochloris, 546,

villosa, 547.

Chrysopelea

ornata, 420.

Chrysotis, 1059.

Cicada, 618.

Ciconia, 645, 679, 688, |

699.

nigra, 1046.

Cinnamopteryx

rubiginosa, 1094.

_ Cinosternum

spurrelli, 1030, 1058.

xxi

Circus

qouldi, 1049.

hudsonius, 1104,

Citharoscelus

koekti, 1098.

Cittotzenia, 569.

Cittura

cyanotis, 1060.

sanghirensis, 1060.

Clathrina, 730, 789.

cavata, 724.

clathrus, 789.

daminoclathrata, 727.

latitubulata, 787,

738.

osculum, 726.

reticulum, 718.

sulphurea, 725, 727.

tripodifera, 706, 717,

728.

— gravida, 724.

ventricosa, 724.

Clava, 505, 522.

Claviporella

pulchra, 483.

Clystolynthus, 789.

vesicula, 789.

Cnesterodon, 980,

carnegie, 1000.

decemmaculatius, 1000,

1001.

scalpridens, 1008.

Cobalopsis

latonia, 363, 367.

Cobalus

gabina, 362.

lateranus, 362, 867,

laureolus, 362, 367.

nigrans, 862, 367.

pindar, 362, 567.

Codiacrinus, 896.

Coendu

mexicanus, 1108.

Coenocystis

richardsoni, 897.

Ceenostomella, 789.

caminus, 789.

Ccenostomium, 789.

crambessa, 789.

Ccenostomus, 789.

primigenius, 789.

Coereba

cyanea, 1065.

Colaptes

meaxicanoides, 1068.

Colius

capensis, 1063.

Coluber

helene, 424.

Columba

livia, 1056.

XX1V

Columbula

picut, 1056.

Colymbus

septentrionalis, 394.

Compsognathus, 629.

Conger

vulgaris, 635.

Connochates

albojubatus, 1090.

Conurus

jendayt, 1058,

Copsychus

saularis, 614.

Coracias

garrula, 1060.

indica, 1060.

Cordylophora, 427,

431,

Corrachia,

gen. nov.,

leucoplaga, 351,

367.

Corynoporella, 465.

Corythzola, 641.

Corythaix

albicristata, 629.

persd, 1057.

Corythophanes

cristatus, 1032.

Costaticella

lineata, 483.

Costicelia, 483.

Cotugnia, 570, 571.

Coturnix

coromandelica, 1052.

Cracticus

destructor, 1065.

Craspedozoum, 475.

Creadion

carunculatus, 1065.

Crenothrix, 431.

hiihniana, 430.

polyspora, 430.

Crex, 670.

Cribricella, 488.

Cribrilina, 529.

figularis, 474, 476.

radiata, 462, 501.

Crisia, 473, 493.

pilosa, 477.

Cristatella, 431, 976.

Crossoptilon

manichuricum, 1052.

Crotalus

confluentus, 422.

Cryptocrinus, 895-897.

Cuculus

canorus, 1057.

Cumulipora

angulata, 502.

INDEX OF SCIENTIFIC NAMES.

Cupularia, 481.

owent, 508,

Cursorius, 651.

Cyanea

arctica, 969.

Cyanocorax

luxuosus, L065.

Cyanops

flavifrons, 1068.

Cyanorhamphus

alpinus, 1058.

Cyathiscus, 789.

actinia, 777, 789.

Cyclocorus

lineatus, 419, 425.

Cycloszemia

subcerulea, 358,

367.

Cyclotosaurus, 956.

Cydonocrinus, 913.

parvulus, 894, 911.

Cygnus

nigricollis, 1047.

olor, 1047.

Cylindrecium, 464.

Cymbirhynebus

macrorhynchus, 1065,

Cyneelurus

Jubatus, 1093.

Cynictis

penicillata, 559.

Cynocephalus

anubis, 856, 859.

Cyphosus

cinerascens, 378.

Cypselus

apus, 1062.

Dacelo

gigantea, 1060.

Danais, 619.

aglea, 617.

limniace, 616, 617.

septentrionis, 615-

618.

Daption

capensis, 1044,

Dasylophus

superciliosus, 643.

Dasyprocta

colombiana, 814.

variegatus, 1094.

Dasypus see Euphractus.

Davainea, 570, 876,

877.

Decapterus

sancte-helene, 378.

Dendrobenia, 479.

Dendrobates

aurotenia, 1029, 1038.

paraensis, 1028, 1029.

tinctorius, 1026-1029,

1038.

— chocoensis,

1028, 1038.

—coctei, 1026, 1027,

10388.

— cocteawi, 1027.

— daudini, 1027.

trivittatus, 1029.

1026,

Dendrocopus, 1065,

major, 1063.

Dendrocygna

Julva, 1047.

Dendrograptus

serpens, 471.

Dendrohyrax

dorsalis, 1104.

Dendya, 706, 712, 716,

717, 719, 730, 736,

737, 742, 798-802.

prolifera, 728.

tripodifera, 710, 728.

Dermatreton, 789.

chartaceum, 789.

hodgsont, 760.

Diachoris

spinigera, 467.

Diastopora, 461.

Diceum

hirundinacewm, 1092.

Diceratosaurus, 962.

punctolineatus, 959.

Dicrurus, 1066.

cerulescens, 614.

leucopygialis, 614.

Didelphys

azare, 1094.

Dinetopia, 465.

Diplocaulus, 960-962.

Diplodidymia, 492.

conplicata, 459, 462,

489-491, 529, 534.

Diploglossus

monotropis, 1032.

Diplophallus, 569, 570.

Diploposthe, 569, 570.

sui-generis, 570.

Diplopylidium, gen. nov.,

559, 567-571, 872-

874.

genett@, 559-561, 563-

566,

Diplostomum, 635.

Dipsadoboa

unicolor, 10938.

Dipsadomorphus

blandingii, 1093.

INDEX OF SCIENTIFIC NAMES.

Dipsas

JSorstenti, 1092.

Dipylidium, 564, 57], |

872.

caninum, 564-

599, 870.

551,

dongolense, 549, 550. |

553-559, 561, 567.

echinorhynchoides, 558,

559.

genette, 558, 559.

gervaisit, Ho8.

monoophorum, 5A9.

monticellit, 5d9.

pasqualii, 554, 558,

BIS),

trtnchesti, 554, 558,

559, Siow

triseriale, 559.

eschokkei, 552, 559.

Discopora, 461.

Dissura, 681, 688, 693, |

703.

episcopus, 1046.

Ditaxipora. 485.

Djedda, 789.

vtolacea, 753, 789.

Dolichotis

magellanicus, 1091.

Dreissensia

polymorpha, 427,

439.

Dromeus, 692.

nove-hollandie, 1043.

Dromas, 651.

Drymobius

dendrophis, 1034.

Dryonastes

cerulatus, 814.

ruficollis, 814.

Dules

caudovittatus, 379.

Suscus, 375, 377.

leuciscus, 378.

maculatus, 378.

malo, 378.

marginatus, 378.

— boninensis, 378.

mato, 378. j

teniurus, 381.

Dunstervillia, 789.

corcyrensis, 746.

elegans, 745, 789.

Jormosa, 746.

lanzerote, TA5.

schmidtii, 746, 748,

Dyssycarium, 790.

egedii, 790.

Dyssyconella, 790.

caminus, 769,

pumila, 790.

Dyssyeum, 790.

Jistulosum, 790.

periminum, 773.

Dyssycus, 790.

primigenius, 790.

Ebnerella, 790.

buccichti, 790.

gregortt, 782.

kitkenthali, 753.

lanceolata, 752.

nitida, 752.

schulzei, 765.

Echelatus

lucina, 358, 367.

Echidna, 1072.

Echinoencrinites, 895,

896.

Echinorhynehus, 427,

Eclectus, 685.

pectoralis, \Q58.

roratus, 1058.

Edwardsia

kerquelensis, 969.

Egernia

depressa, 1092.

Eilhardia, 768, 798.

schulzet, 780, 781.

Elaps

corallinus, 1021, 1036.

microps, 1036, 1038.

mipartitus, 1035, 1036.

rosenbergri, 1021, 1034,

1036.

Hnantia

lua, 307.

— costaricensis, 307,

367.

Enodes

erythrophrys, 1065.

Entomyza

cyanotus, 1065.

Enyalioides

heterolepis, 1082.

Ephydatia

fluviatilis, 427, 436.

Epimachus, 651.

Epinosis, gen. nov., 366. |

angularis, 367.

EKresia

cwla, 344, 567.

pacilina, 344.

sticta, 344, 367.

Krithizon

dorsatus, 542.

Eryops, 956, 957, 961.

Erythrosuchus, 623, 6380.

Eschara

hexagonalis, 529,

platalea, 529.

Escharina, 505.

Kscharinella

cecilleana, 529.

Escharoides, 461.

oceiusa, 463, 519-

521.

Estrelda

phenicotis, 1065.

Eubagis

hecubu, 347, 367.

vicaria, 247.

HKueratea

chelata, 462, 466.

cordieri, 492.

Tudocimus

longtrostris, 650,

Eudyptes

chrysocome, 394.

Eugonodeum, gen. nov.,

872, 874-877.

edicnemi, 862-866,

- 868, 870, 871, 873,

877.

Eunectes

murinus, 814.

Euparkeria, 620, 624,625,

629, 630, 632.

capensis, 619, 627, 628,

632, 633.

Hupetoieta

poasina, 343, 367.

| Euphractus

sexcinctus, 1102, 1103.

villosus, 1099.

villosus, 1099,

Euplectella, 787.

Euplea, 617-619.

core, 615, 616.

coreta, 6195.

kollart, 615, 616.

Eupodotis

kori, 564.

Euptychia

agnata, 842, 367.

antonoé, 342.

confusa, 342.

drymo, 341, 367.

juani, 342,

labe, 342.

Eurygona

cataleuca, 349.

leucon, 3849, 367.

leucophryna, 849,

367.

matuta, 849, 867.

mystica, 348, 367.

EKurylemus

cchromelas, 1065.

Eurypyga, 645, 651,

699.

helias, 1054, 1092.

XXV1

Eurystomus

orientalis, 1060.

Eutimalphes

indicans, Y70.

Evotomys, Nb

alstoni, 522, 827, 829,

&30, 839.

cesarius, 829.

glareolus, 829.

—. britannicus,

830.

norvegicus, 829, 830.

skomerensis, 829.,

827,

Falcinellus, 651.

Falco

pereyrinus, 1049.

Falga

hermione, 366, 367.

Farcimia, 490.

oculata, 462, 489, 535

Felis

caracal, 1098.

concolor, LOO1.

ocreata, 1098.

pardalis, 1091.

tigrina, LUOL.

Flabellaris, 474.

cuspidata, 475.

flabeliata, 475.

ligulatum, 475.

multisertata, 475.

roborata, 475.

triseriata, 475.

(Craspedozoum) ligz-

latum, 475.

(—) rohorata, 475.

(Menipea) cuspidata,

534.

Flustra, 494, 529.

abyssicola, 474, 476.

cecilii, 508.

mamillaris, 497.

Francolinus

infuscatus, 1052.

Fredericella, 429, 431,

444, 447.

sultana, 427, 438, 459,

441, 448, 457.

Fregata, 661, 1045.

Fulica

leucoptera, 1052.

Fulmarus, 670.

Gadus

eglefinus, 578.

pollachius, 637.

Galago

maholi, 814.

JNDEX OF SCIENTIFIC NAMES.

Galbula

albivostris, 1063.

rufiventris, 1063.

Galeopithecus, 631.

Gallinago

celestis, LOD4.

Gallinula

chloropus, 1052.

phenicura, 10d2.

Gallirex, 639.

jJohustoni, 641,

Gallus

gallus, 1051, 1052,

1098.

varius, L099.

Gambusia, 979,

1000.

affinis, 982, 984.

annectens, 992.

bimaculata, 993.

bonita, 993.

caudovittata, 982, 986.

caymanensis, 982, 990.

dominicensis, 982, 989,

990, 1018.

dovit, ‘982, 986.

episcopi, 992.

Jasciata, 995.

991,

gracitior, 982, 983, 989,

1018.

gracilis, 984, 999.

holbrookti, 982, 988,

990.

humilis, 984.

infans, 998.

Jonesti, 993.

melanopleura, 982, 988.

melanosticta, 982, 987.

nicaraguensis, 982, 983,

985.

nigropunctata, 982,983,

987.

nigroventralis, 992.

nobilis, 98d.

oligosticta,

990, 1018.

parismina, 992.

patruelis, 982, 984.

picturata, 981.

punctata, 982,

986.

982,

983,

puneticulata, 982, 987.

rhabdophora, 997.

senilis, 982, 983, 98d.

speciosa, 984.

terrahensis, 993.

tridentiger, 992.

turrubarensis, 992.

umbratilis, 998.

wrayt,

1018.

988,

982, 983, 988,

Gammarus, 426, 431,434,

437.

pulex, 427.

Gasterocoma, 897,

Gasterosteus

aculeatus, 428.

Gavia

stellata, 394.

Gazella

bennetti,

Gecinus

vittatus, 1063.

Gemellaria

loricata, 465, 466.

Gemellipora

glabra striatula, 507.

898.

1093.

protusa, 462, 506,

53).

striatula, 507.

Genetta

dongolana, 549.

Gennzeus

horsfieldi, 542.

Geocichla

citrina, 1066.

Geopelia

cuneata, 1056.

tranguilla, 1056.

Geophaps

plumitera, 1056.

Geothelphusa, 924.

Gephyrophora

polymorpha, 521.

Giraffa

camelopardalis

quorum, 542.

Girardinus, 978, 980.

caucanus, 1013.

caudomaculatus, 999.

creolus, 1002.

denticulatus, 1002.

Jormosus, 995.

garmant, 1002.

guppyt, 1008, 1010.

theringit, 999.

januarius, 999.

lutzi, 996.

metallicus, 1001, 1002.

pleurospilus, 997.

presidionis, 996.

reticulatus, 1008.

uninotatus, 1000.

versicolor, 1018.

zonatus, 999.

Girella

nebulosa, 369.

Girellops, gen. nov., 369,

373.

nehulosus, 369, 373.

Glareola, 651.

pratincola, 1055.

antt-

INDEX OF SCIENTIFIC NAMES,

Glaridichthys, 978, 980.

falcatus, 1000, 1002.

latidens, 1002.

torralbasi, 1000, 1002.

uninotatus, LOOU, LO02.

Glaridodon

Januarius, 999.

latidens, 1002.

wuninotatus, 1000.

Glossiphonia, 427.

Glossostoma

aterrimum, 1025.

Glyptolepis

paucidens, 957.

Grantessa, 704,

798.

compressa, TA2.

erecta, T52.

erinaceus,

flamma, 750, lige

glabra, 751, 752

glacialis, 753.

hastifera, 711, 752.

hirsuta, 7H0-752.

hispida, 711, 750-752.

intusarticulaza, J11,

Toll, Tass

kitkenthali, 753.

lanceolata, 752.

murmanensis, 7)3.

nitida, 72.

elanitoy7 WO

“all,

pluriosculifera, 752.

poculum, T\1, 752.

polyperistomia, 711,

(52, 753.

preiwischt, 753.

sacca, 711,

703.

simplex, 753.

spissa, 759.

stauridea, 753, 789.

sycilloides, 753.

thonpson, 753.

zanzibarensis, 703.

(Ainphoriseus) pocu-

lum, TAO.

Grantia, 704, 738,

749, 750, 751,

766, 768, 769,

798, 805.

aculeata, 761.

arabica, 773.

asconoides, 760.

atlantica, 199.

botryoides, 721.

— nen 725,

brevipilis, TH9.

canadensis, 759.

capillosa, TH9.

lod

750, 751

723.

=t)

744,

7TdA,

781,

Grantia

chartacea, 759, 789.

ciliata, 745.

clathrus, 725.

clavigera, T59.

cliftont, 724.

comoxensis, 759.

compressa. T11, 757,

759, 760, 761, 787,

789, 795.

cupula, 761.

ensata, 763.

extusarticulata, 761.

fistulosa, 770.

Joliacea, 760.

genuina, 711, 760.

gracilis, 760, 796.

hodgsoni, T6V.

indica, 711, 713, 761.

intermedia, 759, 760,

804.

invenusta, TAL.

labyrinthica, 762.

lacunosa, 722.

levigata, 761, 793,

796.

lieberkithniti, 723.

lobata, 760, 795.

longipilis, T60.

mirabilis, 761.

monstruosa, 760.

multicavata, 725.

nivea, 773.

pennigera, 760.

phillipsii, 761.

pulchra, 726, 727.

scotti, 760.

singularis, 761, 794.

solida, "734.

striatula, 786.

strohbilus, 761.

subhispida, 748.

tenuis, 760.

tessellata, 749.

truncata, 765.

tuberosa, 761.

urceolus, T61.

virgultosa, 746.

vosmaeri. 711, 760.

Grantilla, 751, 798, 804.

hastifera, 752, 756.

quadriradiata, 76,

797.

Grantiopsis, 766, 798.

cylindrica, 762, 763.

infrequens, 11, 769,

Tol.

Grus, 698.

australasiana, 1053.

Gryponyx

africanus, 638.

XXVil

Guancha, 790.

blanca, 724, 790.

Guire

piririgua, LOST.

Gulapinnus, 1000.

Guttera

cristata, 1051.

edowardi, 1051.

pucherant, 1051.

Gymnoschizorhis, 641.

Gymnothorax

dovti, 373.

meleagris, 373.

Gypaétus, 645.

barbatus, 1049.

Heematopus

ostralegus, 1055.

Halcampa, 968.

chrysantheilum,

ale

clavus, 969.

fultoni, 970, 972

medusophila, 970.

septentrionalis, 969.

Halcampoides

abyssorum, 969.

Haleyon

pileata, 1060.

rufa, 1060.

Haliaétus

leucegaster, 1049.

Halysis

diaphana, 482.

Haplochilus

melanopleura, 988,

melanops, 984.

Haswellia

auriculata, 512.

australiensis, 462, 511,

OUZA520}

coronata, 512.

gracilis, 512.

grandipora, 512.

Hebius, gen. nov., 424.

Heliconius

galanthus, 343.

— subrufescens,

367.

Helicops

angulatus, 424,

leopardinus. 425,

modestus, 425.

Heliornis, 678.

Fulica, 1054.

Hemeschara

gigantea, 518,

Hermathena

owen, 300,

Herpetodryas

carinatus, 1034.

970,

343,

367.

XXVIII

Herpetosuchus, 629.

granti, 626. ~

Hesperia

parvipuncta, 366.

Hesperornis, 394, 396.

regalis, 395.

Hetrandria, 980.

affinis, 984.

colombianus, 996.

cubensis, 1002.

Jasciata, 995.

formosa, 977, 994, 995. |

holbrookii, 983.

lutzi, 996.

minor, 1003.

nobilis, 985.

occidentalis, 1018.

petruelis, 984.

Heteropegma, 728, 737,

790.

nodus-gordit, 706, 736-

738, 790.

Heteropia, 798.

compressa, 752.

erecta, 72.

glomerosa, 711, 754.

macera, TOD.

patulosculifera, 752.

pluriosculifera, 752.

polyperistomia, 758.

ramosa, TO4.

rodgeri, 754.

simplex, 754.

spissa, 753.

Heterusia

cingala, 614.

Hexelasma

sp., 847.

arafurae, 847.

aucklandicum,

841, 843, 847,

854.

corolliforme, 847.

velutinum, 847.

Hierococcyx

varius. 1057.

Hierofaleco

candicans, 542.

Himantodes

cenchoa, 1035.

Himantopus

nigricollis, 1055.

Hincksiella, 483.

Hippopodina, 515.

feegensis, 514.

Hippoporina, 515.

Hippothoa

distans, 462, 501.

divaricata, 462, 501.

Hirundo, i069.

rustica, 1066, 1067.

840,

848,

Holocentrum

punctatissimum, 3873.

Holocentrus

caudovittatus, 379.

Holoporella, 461,

509, 529.

albirostris, 463,

522, 523, 537.

aperta, 463, 522.

bispinata, 522, 523.

celosia, 521.

cidaris, 521.

columnaris, 463, 505,

621.

pigmentaria, 509.

Holoptychius, 957.

Holoscalpellum, 850.

Homalocranius

alticola, 1036.

coralliventre,

1038.

melanocephalum, 1030.

Homandra, 717, 720,

788.

Jalcata, 790.

Hometta, 791.

Homoderma, 744,

802

sycandra,

716,

748.

Howesia, 628, 632.

Hydra

viridis, 427.

Hydrocherus

hydrocherus, 1092.

Hydrophasianus

chirurgus, 1055.

Hydrophis

fasciatus, 414.

Hydrornia

alleni, 1052.

Hydrothelphusa,

923, 924.

agilis, 915, 922.

Hyla

‘ arborea, 1028, 1024.

baudintt, 1023.

maxima, 1022,

rubra, 1026.

Hylobates

lar, 813.

Hylodes

conspicillatus, 1025.

palmatus, 1025.

Hymenolepis, 564.

Hyperodapedon, 628.

Hypocrinus, 894, 897,

899, 903.

miilleri, 896, 900, 913. |

piriformis, 896, 900,

905, 907, 910-913.

508,

516,

1035,

791, |

742,

918,

1028.

INDEX OF SCIENTIFIC NAMES.

Hypocrinus

pyriformis, 896, 905.

schneideri, 895, 896,

900, 901, 904, 910,

913,

Hypodictyon

longstaffi, 44, TAT,

TOM

Hypograntia, 791.

extusarticulata, 761.

hirsuta, 752.

infrequens, 762,

TM.

intusarticulata, 753.

medioarticulata, 753.

763,

Hypotnidia

philippensis, 1052.

Hypoxanthus

rivolt, L063.

Hyracotenia

hyracis, 552.

Tanthia

rufilata, 814.

Tbis, 690.

Ichthyoteenia, 562.

Icterus

jamaicat, 1065.

mesomelas, 1104.

mississipprensis, 1104.

Tetinia, 1050.

mississiptensis, 1049.

Inermicapsifer

capensis, 568.

Ischnognathus

lineatus, 424.

Ithomia

bolivart, 341, 367.

Jacana

jacana, 1055.

Julis

umbrostigma, 371.

Kebira, 798, 800.

uteoides, 78d.

Kellogella

oligolepis, 373.

Ketupa, 651.

Kuarrhaphis, gen. nov.,

767, 780, 798, 803.

cretacea, 775, 780.

Kuhlia

boninensis, 374, 3875,

378.

cerulescens, 875-377.

caudovittata, 375, 378,

379.

Kuhlia

humilis, 369, 374, 375,

379, 380.

maculata, 378.

malo, 378-381.

marginata, 874, 875,

317, 378.

mutabunda, 3869, 373-

375, 381.

prozima, 380.

rupestris, 314-377.

— hedleyi, 379.

sandvicensis, 369, 374,

379, 381.

sauvagii, 375, 377.

splendens. 374, 379,

379, 380.

sterneckit, 381.

teniura, 375, 381.

urvillit, 375, 377.

venura, 381.

Labrichthys, 368.

fuentesi, 371, 375..

luculenta, 371.

Labyrinthodon

leptognathus, 997.

Lacerta

viridis, 546.

Lachesis

atrox, 1037.

brachystoma, 1037.

lanceolatus, 1037.

monticellii, 1037.

mutus, 814, 1037.

punctatus, 1037.

Lagenipora, 508.

borywt, 511,

lucida, 511.

nitens, 511.

rota, 462, 510, 511.

socialis, O10, 511.

Lagothrix

poppigt, 1091.

Lalocitta

lidthit, 1104.

Lamontia, 768, 778, 798.

zona, 179.

Lampanyctus

crocodilus, 1097.

Lanius

excubitor, 1066.

Larus

argentatus, 1055,

ridihundus, 105d.

Leander, 428.

(Palemon) squitla,

427.

Lebistes, 981, 1010.

pecilioides, 1008.

INDEX OF SCIENTIFIC NAMES, b--db:4

Lebistes

reticulatus, 978, 1007,

1008.

Lechriorchis

inermis, 933, 936.

primus, 934.

validus, 933, 935, 936.

Leevthiocrinus, 896.

adamsi, 899, 900, 913.

Lelapia, 739, 740, 784,

798, 800, 804.

australis, 711, 785.

Lepralia, 461.

adpressa, 515.

circumcineta, 515,

cleidostoma, 517, 518.

— mermis, 463, 517.

clivosa, 503.

crassa, 519.

cucculata, 528.

dorsiporosa, 518, 519.

Jeegensis, 463, 514,

Silisy, lays}oy,

Foliacea, 503.

hippopus, 515, 524,

inornata, 507.

japonica, 518.

longipora, 510.

margaritifera, 503.

mont ferrandi, 506.

occlusa, 489, 519.

pallasiana, 518, 515.

pertusa, 502.

porcellana, 517.

prelonga, 515.

prelucida, 5198.

rectilineata, 515.

striatula, 463, 507.

syringopora, 504, 529,

turgescens, 507.

turrita, 463, 516, 537.

venusta, 506.

wasinensis, 463, 516,

ole

(Mucronella). pre-

lucida, 517.

Lepreus

| tricolor, 1091.

Leptodactylus

pentadactylus, 1025.

Leptodira

albo fusca, 1035.

Leptognathus

annulata, 1037.

spurrelli, 1036, 1038.

temporalis, 1037.

Leptophilus

crumeniferus, 1046.

| Leptophis

bilineatus, 1034.

occidentalis, 1035.

Leptoptilus, 668, 672,

678, 677-679, 681,

685, 685, 686, 699.

argala, 654.

crumeniferus, 654.

Leptorhaphis, gen. noy.,

980, 998.

infans, 998.

Leptosia

xiphia, 616,

Lerodea

phyllas, 361.

rupilius, 360, 367.

Leucaltis, 712, 736, 738,

757, 798, 800, 804.

bathybia, 773.

— mascarenica, ‘737,

738.

clathria, 706, 710, 728,

736-738, 790.

crustacea, 770.

Horidana, 734.

— australiensis, 734.

helena, 774.

impressa, 774.

nausicae, TTA.

pumila, T74.

solida, 734.

(Heteropegma), 804.

Leucandra, 709, 731, 733,

736, 738, 758, 766,

168, 779, 781, 783,

798, 804.

alcicornis, 777.

— cespitosa, 777.

amorpha, 772.

ananas, 769.

anfracta, 772.

anguinea, 769,

anomaa, 769,

apicalis, 72.

armata, 769,

aspera, 730, 769, 774,

775, 795.

_ australiensis, 711, 769,

803.

balearica, 772.

bathybia, 773.

bomba, 732.

brumalis, 774.

bublosa, 772.

callea, 772.

caminus, 769, 789.

capillata, 770.

cartert, 734.

cataphracta, 777.

cirrata, 770.

cirrhosa, 770.

claviformis, 770.

coimbre, 772.

compacta, 770.

XXK

Leucandra

conica, 772.

crambessa, 770, 776,

789.

— callea. 772.

erosslandi, 772.

crustacea, 77).

cucumts, 735, 778.

cumberlandensis, 770.

curva, 774.

cyathus, 736.

cylindrica, 772.

donnant, 770.

echinata, 711, 770.

egedii, 770, 790.

elongata, 777.

falcigera, 770.

fernandrica, 772.

fistulosa, 770, 790.

Frigida, 774.

gelatinosa, 774.

gemmipara, 770.

gladiator, nae

gossei, 710, 791.

heathii, 772.

helena, 774.

hiherna, 770.

hirsuta, 770.

hispida, 711, 771.

impressa, 774.

infesta, 769, 771, 803.

inflate, 774.

innominata, T74.

intermedia, (74.

johnstonii, 773.

joubini, 772.

kerquelensis, 771.

lendenfeldi, T7\.

levis, TTA.

lohata, 773.

loricata, 772.

lunulata, 771.

masatierre, 771.

meandrina. T1\. 771.

minima, T11, 771.

minor, 772.

multifida, 773.

multiformis. 771.

nausicae, T7T4.

nived, 773.

ochotensis,

ovata, 773.

pailida, 771.

pandora, 774.

panicea, 77

775.

75.

phillipensis, T11, 771.

plate, 772.

prava, Hbitexs

pulvinar, 771, 792.

pumila, 774, 790.

pyriformis, 773.

Leucandra

rodrigueztt, 773.

rudifera, 773.

saccharata, 734.

sagittata, 774.

schauinslandi, 774.

solida, 787.

spissa, 173.

stylifera, 776.

taylori, 773.

teluim, 774.

thulakomorpha, 711,

771.

typica, 773.

vaginata, 771.

valida, 771.

verdensis, 774.

villosa, 771.

vitrea, 773.

wasinensis, 772.

(Leucetta) pandora,

785.

(Leucortis) pulvinar,

78).

Leucaseus, 716, 717, 732,

736, 786, 798, 800.

clavatus, 10, 730,

Veil

insignis, 710, 731.

simplex, 710, 730,

Tile

Leucetta, 709, 731, 735,

738, 757, 758, 768,

783, 798, 800, 804.

carteri. TH.

chagosensis, 710, 753.

clathrata, 724.

corticata, 739.

expansa, 710, 734.

floridana, 734.

haeckeliana, 739.

homoraphis, 734.

imperfecta, 739.

infrequens, T34.

microraphis, 710, 734.

pandora, 774.

— anomala, 769.

— intermedia, 774.

-— loculifera, 777.

primigenia, 730, 732,

734, 789, 790-792,

796.

— microraphis, 734.

prolifera, 710, 734,

796

pyriformis, T10, 734.

sagittata, T74.

sambucus, 739.

schauinslandi, 774.

solida, 734.

trigona, 734.

INDEX OF SCIENTIFIC NAMES.

—

Leucetta

vera, 739.

(Teichonella) prolifera,

762.

Leucettaga, 768, 798.

loculifera, V77.

loculosa, 749.

Leucettusa, 732, 728,

757, 768, 798, 800,

804.

corticata, 738, 739.

dictyogaster, 710, '736,

738, 739.

haeckeliana. 733, 739.

imperfecta, 739.

sambucus, 739.

vera, 138, 739.

(Leucetta) haechkeliana,

738.

(—) vera, 738.

Leucilla, 733, 736, 768,

778, 728, 798.

amphora, 730, 783.

australiensis, 711,

783.

capsula, 784.

connexiva, TAR,

crosslandt, 774.

echinus, 784, 792.

intermedia, 769,

803.

leuconides, 784.

nuttinyt, 784, 793.

oxeodraginifera, T34.

princeps, 711, 783, 734.

prolifera, 734.

proteus, 784.

saccharata, 784.

uter, 784, 793.

wasinensis, 772.

Leuckartea, 791.

natalensis 748, 791.

Leucogvpsa, 778, 791

795.

algozensis, 786.

gosset, 770, 791.

Leucomalthe, 731, 768,

798, 800.

bomba, 732.

Leucometra, 791.

primigenia, 791.

Leuconia

anfracta, 772.

armata, 769.

balearica, 77

cirrhosa, 770.

cotmbre, 772.

compacta, 770.

crucifera, 768.

dura, 734.

echinata, 770.

—

Leuconia

erinaceus, 752.

Sernandensis, 772.

Jistulosa australiensis,

769.

Sruticosa, 734.

glomerosa, 754.

hispida, 771.

johnstonii, 773.

— australiensis, 783.

levis, 774.

lendenfeldi, 771.

lobata, 773.

loricata, 772.

masatierreé, 771.

minor, 772.

multifida, 773.

multiformis, 771.

— amorpha, 172.

— capillata, 770.

nivea, T34.

ovata, 773.

platei, 772.

prava, 773.

pumila, 774.

pyriformis, 773.

rodrigueztt, 773.

rudifera, 773.

somesi, 723.

spissa, 775.

stylifera, 776.

typica, 773.

vitrea, 773.

Leucopsila, gen. noy.,

775, 768, 798, 803.

stylifera, 776.

Leucopsis, 792.

pedunculata, 726, 731,

792.

Leucortis, 792.

anguinea, 769.

elegans, 771.

pulvinar, 771, 792.

Leucosarcia

picata, 1056.

Leucosolenia, 705, 708,

TG, 7, ZN), "eat,

798, 812.

agassizti, 724.

ameboides, 720.

angulata, 721.

arachnoides, 721.

armata, 721, 788.

atlantica, 72).

bella, 709, 721, 742.

ilanca, 724, 790.

botryoides, 721,

723.

botrys, 721.

canariensis, 720, 723,

72+, 787, 788, 796.

INDEX OF SCIENTIFIC NAMES.

Leucosolenia

cancellata, 724.

caroli, 724.

cavata, 710, 720, 724.

cerebrum, 724.

cervicornis, 721.

challengeri, T24.

charybdea, 724.

clarkit, 721.

clathrata, 724, 728,

789.

clathrus, 725,

complicata, 709, 721,

799.

confervicola, 721.

contorta, 721.

convallaria, 7295.

corallorhiza, 721.

cordata, 721, 787.

coriacea, 710, 725, 793-

796, 799.

darwinii, 725.

decipiens, 725.

densa, 721, 796.

depressa, 710, 725.

dictyoides, 729.

discoveryt, 722.

dubia, 722.

echinata, 722.

echinoides, 722.

eleanor, 722.

fabricii, 722.

falcata, 709, 720, 722,

790.

Jalklandica, 725.

flexilis, 725.

fragilis, 722.

W217.

goethet, 725.

gracilis, 725,

793.

grantit, 725.

hermesi, 722.

himantia, 726.

hispidissima, 722.

horrida, 722, 788.

incerta, 722.

intermedia, 724.

irregularis, 722.

japonica, 726.

lacunosa, 722.

lamarckhit, 726, 788.

laminoclathrata, 727.

laxa, 722.

lieberkithnii, 723.

loculosa, 726, 788, 794,

796.

lucasi, 709, 718, 720,

723, 742.

Proc. Zoou. Soc.—1913, No. LXXIV.

OM!

Leucosolenia

macleayt, 726.

minchint, 723.

ninoricensis, 726.

multiformis, 726.

nansent, 724.

nitida, 723, 793, 794.

osculum, 726.

panis, 723.

pedunculata, 726, 792.

pelliculata, 710, 726.

philiipina, 726.

pinus, 728.

poterium, 726.

primordialis, 726, 727,

788, 793.

protogenes, 726.

proxima, 710, 727.

psammophila, ‘727.

pulcherrima, 710, 727.

pulchra, 727.

reticulata, 723.

reticulum, 723, 792.

robusta, 723.

rosea, 727.

sagittaria, 727.

sceptrum, 727.

sertularia, 723.

spinosa, T27,

spongiosa, 727.

stipitata, 727.

stolonifer, 709, 718,

720, 723:

sulphurea, 727.

sycandra, T48.

tenutpilosa, 723.

tenuis, 723.

thamnotdes, 72°2.

trépodifera, 728.

uteoides, 729.

variahilis, 710, 7238.

ventricosa, 709, 720

724.

vesicula, 727, 739.

vitrea, 710, 727. -

wilsont, 727, 777.

(Ascandra) falcata,

720.

Leuculmis, 792.

echinus, 784, 792.

Leucyssa, 767, 798,

803.

eretacea, 780.

tncrustans, 759.

-— lichenoides, 779.

— villosa, 780.

spongilla, 779.

Lichenocystis, 897.

Lichenopora, 461.

Limenitis

cealidasa, 614.

XXX]1 INDEX OF SCIENTIFIC NAMES.

Limia, 981, 1010. | Mamosa | Menipea

arnoldi, 1014, 1016, elegans, 1092. | flagellifera, 473, 474,

1018. Mastigina, 799. 476.

caudofasciata, 1014, Medon Suegensis, 474.

1017, 1018. barbata, 786, 792. gracilis, 473.

couchiana, 1008. wmberbis, 786. occidentalis, 473.

cubensis, 1014. »Megaleema patagonica, 473, 534.

dominicensis, 1014, virens, 1063, 1064. | portert, 473.

LOTS; LOM Megalichthys, 957. smithii, 4738.

formosa, 1012. Megalotis ternata, 473, 475.

heterandria, 1014, cerda, 558. Menura

1017, 1018. Megapogon, 755, 769, superba, 1065, 1067.

matamorensis, 1011. 798, 803. Mergus

nigrofasciata, 1014, crispatus, 767, 768. serrator, 1047.

1015, 1018. cruciferus, 767, 768. | Merlia, 792.

ornata, 1014, 1016, poilicaris, 767, 768. normant, 792.

1018. raripilus, 767, 768. | Merops

pecilioides, 1011. | _villosus, 767, 768. | sp., 615.

versicclor, 1009, 1014, | Megistias | apiaster, 1060.

1017. cantho, 363, 367, | -philippensis, 1060.

vittata, 1014. Melanerpes | Merula

Limnea, 427, 437. flavifrons, 1091. | tamaulipensis, 1.066.

peregra, 434, 439. | Melanitis | Mesites, 645, 702.

Limosa | ismene, 614. Mesocestoides

lapponica, 1054. | Melanocetus, 1097. ambiguus, 549.

Liophis | johnsonii, 1096. | Mesosemia

albiventris, 1035. | Meleagris albipuncta, 350, 367.

Liothrix gallopavo, 1051. Mesosuchus, 628, 632.

luteus, 1066. Meliceritites | brownt, 619, 627, 632,

Lipostomella, 792. | magnifica, 505. | 633.

capsuia, T84. | royana, 505. Metopiana

elausa, 734, 792. | _vielbanci, 505. peposaca, 1047.

Lithonina, 739. | Melopsittacus Micropholis, 956.

Lophius, 960. | wndulatus, 1058. Micropora

piscatorius, 1097. Membranipora, 529. articulata, 490, 492.

Lophoceros | armata, 459, 462, 486— | ratoniensis, 490,

erythrorhynchus, 106). | 488, 534, 535. _ Microporella

Lophortyx | catenularia, 462,488. | ciliata, 463, 523.

douglasi, 1052. | eraticula, 475. | Microtus

Lophura lineata, 475. agrestis, 834, 838,

nobilis, 10938, | magnilabris, 498. 839.

Lorius mamillaris, 497. — exsul, 830-834,

donvicella, 1058. marginella, 488. | 839.

Lota | melontha, 461. | — levernedii, 834.

vulgaris, 428. | monostachys, 488. | —macgillivrati, 822.

TLoxomma, 958-958, 961. | nigrans, 488. 831, 832, 834, 839.

Lutra panhoplites, 487. | — neglectus, 833, 834.

leptonyx, 1092. savartii, 462, 486, | arvalis, 838, 839.

Lycaon | 535. | corneri, 838.

pictus, 538. unicornis, 475. hirtus, 834.

Menipea, 472. — bailloni, 834.

aculeata, 474. orcadensis, 838.

Macacus benemunita, 475. sandayensis, 838.

silenus, 1103. | buskii, 473, 534. sarnius, 838.

Machetes cervicornis, 473, 474. ~ | Mimo

pugnax, L054, | cirrata, 473-475. dumontt, 1065.

Macropora clausa, 475. Mimus

cribrilifera, 496. crystallina, 478, orpheus, 1066.

Macropus, 592. | ATA. | Minchinella, 739, 798,

Malapterurus cyathus, 474, 554. 800.

electricus, 1093. flabellum, 475, | damellosa, 710, 740.

INDEX OF SCIENTIFIC NAMES.

Mitua

tuberosa, 1051.

Mlea, 792.

dohrnii, 771, 792.

Mobiusispongia, 792.

parasitica, 792.

Mollia

tubereulata, 507.

Mollienisia, 978, 981.

elongata, 1010, 1013.

formosa, 1010, 1012. |

gracilis, LO10, 1012.

Jonesit, 993.

latipinna, 1010, 1011.

occidentalis, 1010,

1013

petenensis, 1010, 1012.

sphenops, 1009, 1010,

1012. |

spilurus, 1010, 10138.

Molo

apella, 865, 367.

herea, 365.

humeralis, 365.

nebrophone, 364, 367.

Molothrus

fringillarius, 1094.

Molpastes

hemorrhous, 614.

Momotus

lessoni, 1060.

subrufescens, 1060.

Monacanthus

cirrifer, 373.

Monieza, 569.

Monopylidium,

872.

macracanthum, 571. |

rostellatum, 568, 876, |

877.

unicoronata, 877.

Moronopsis

argenteus sandvicensis,

567,

fuscus, 377.

sandvicensis, 881,

Mueronella, 461.

vultur, 518, 519.

Murrayona, 789, 740,

798, 800.

phanolepis, 710, 741.

Mus

musculus, 838.

Musophaga, 639.

violacea, 641.

Mustela

erminea ricine, 829.

Mycteria, 699.

americana, 1046.

Myiarchus |

tyrannulus, 10G5.

Myopsittacus, 1059.

monachus, LO58.

Myriozoella, 520.

Myriozoum

australiense, 511.

coarctum, 520.

marionensis, 520, 521.

simplex, 520.

subgracile, 520.

truncatum, 520.

Myripristis

pralinius, 373.

Myrtillocrinus, 898.

americanus, 899,

élongatus, 899.

Mysis

chameleon, 427.

Naia

bungarus, 1093.

nigricollis, 1093.

Nannopterum

harrisi, 393, 899, 401, |

A02.

| Nanocrinus, 897, 898.

Nanodes

(Lathamus) discolor,

1058.

Napeogenes

hemisticta, 340, 367.

Nardoa

arabica, 726.

canariensis, 724.

labyrinthus, 725.

pelagica, 752.

reticulum. 728, 792.

rubra, 724.

spongia, 725.

spongiosa, 727.

sulphurea, 724.

Nardoma, 793.

nitida, 793.

Nardopsis

gracilis, 721, 798,

horrida, 722.

Nardorus, 793.

primordialis, 793.

Nascus

phintias. 358.

phocus, 358.

Necturus. 403,

Nellia, 49v.

tenella, 489.

_ Neophron, 661,

Nephelis, 427.

| Nicoria

nasuta, 1030,

Niltava

grandis, 814,

Notamia

bursaria, 465, |

XXXi1ll

Nothura, 1051.

maculosa, 1050.

Notodela

leucura, 814,

Nototragus

melanotus, 1092.

Nycticorax, 649, 656, 679,

686, 699.

gardeni, 654, 1045,

1046.

violaceus, 1045.

Nyetidromus, 1065,

1072.

albicollis, 1062.

Nymphicus

uveensis, 1058.

Oceanodroma, 1044.

Octacnemus, 440.

Octodon

deqgus, 1095.

Octorchis, 970.

Ocydromus

australis, 10D2.

Gidemia

negra, 1O4T.

Ciidicnemus, 861.

bistriatus, 861, 863.

edicnemus, 861.

scolopax, 1055.

CEnus

nausiphanes, 363, 367.

Oligodon, 424.

Olindias, 969.

Olynthella, 793.

coriucea, 799.

Olynthium, 798.

nitidum, 723, 793.

splendens, 723.

Olynthus, 719, 793,

798.

cathus, 722.

guancha, 724.

hispidus, 72).

pocillum, 721,

simplex, 726, 793.

Oncorhynchus, 576.

gorbuscha, 572,

608, 610.

keta, 572, 577, 604,

605, 610.

kisutch, 572, 599-602,

610.

masu, 572,

nerka, 572, 586, h88—

590, 592, 593, 610.

guinnat, 572.

tschawytscha, 572,

B75, 585, 594,

597, 598, 610.

74"

602,

574,

o95,

XXXIV

Oochoristica, 557, 567,

568, 571, 876.

wagenert, 874, 875.

Opisthocomus

cristatus, 1052.

Oreicola

Serre, 814.

Oriolus

galbula, 1066.

Orizomys

longivaudata, 1092.

Ornithorhynchus, 1072.

Ornithosuchus, 629, 631.

taylori, 625, 626, 630,

633.

woodwardi,, 620, 624-

626, 6380, 633.

Ortalis

araucuan, 1104.

ruficauda, 1051.

Ortygospiza

atricollis, 1093.

Ortyx

virginianus, 1052,

Orycteropus, 886, 888,

889, 892, 893.

Osearella, 738.

Osinotreron

bicincta, 1056.

Osteolepis, 957.

Osthimosia, 510.

zanzibariensis,

508, 537.

Ostinops

decumanus, 1065.

Otiditzenia, 564.

Otis

tarda, 1054.

Ovis

musimon, 1094.

Oxynaspis, 938, 942,

948.

Oxyrhopus

clelia, 1035.

462,

Pachynus

brachyurus, 1058.

Palemon, 930.

dolichodactylus,

926.

hildebrandti, 925, 928,

931, 932. :

hilgendorfi, 926.

ide, 927, 928.

lepidactylus, 925, 926.

mooret, 929.

925,

ritseme, 925, 927,

928.

sintangensis, 928.

(Kupalemon) 7itseme,

927.

Palzemon

(Macrobrachium) Jepi-

dactylus, 926.

(Parapalemon) do-

lichodactylus, 926.

Palxornis

fasciata, 1058.

Palxosyops, 886, 893.

Palamedea

cornuta, 1047.

Palmicellaria, 461.

Paludicella, 429, 431,

444, 446, 448,

449.

articulata, 427, 489,

441, 443, 447, 456.

ehrenbergii, 441.

Pamphila

angularis, 366,

Pamphoria, gen. nov.,

980, 1003.

scalpridens, 1008.

Pamphorichthys, gen.

noy., 980, 1003.

minor, LOO.

Pancerina, 571.

| Pandion, 1069, 1072.

haliaétus, 1050.

Papilio

agamemnon, 616.

aristolochie, 614, 617,

618.

demoleus, 615.

hector, 614, 615.

jason, 615, 618.

nomius, 615.

parinda, 614.

polytes, 614-616, 618,

619

— cyrus, 615, 616.

— polytes, 614-618.

— romulus, 614-616.

Papio

anubis, 1090.

Paragambusia, 982.

nicaraguensis, 985.

Paraleucilla, 735, 798.

cucumis, 778.

Paramecium, 428.

Paraphysa

manicata, 1093.

| Parathelphusa, 925.

tridentata, 923.

Parotia

lawestt, 1065, 1066.

Passerculus

savanna, 1091.

Pauxis

galeata, 1051.

Pavo

nigripennis, 1098.

INDEX OF SCIENTIFIC NAMES.

Peachia

boeckii, 966, 967.

carnea, 966, 967.

fultoni, 970.

hastata, 963, 964, 966,

967, 970, 971.

koreni, 966, 967.

quinquecapitata, 963,

966, 967-969, 972.

tricapitata, 964, 967.

triphylla, 966, 967,

972.

undulata, 966.

Pectinatella, 431.

Pelargopsis, 1065.

Pelecanus, 661, 688, 698,

703, 1045.

Penelope, 1052.

pileata, 1104.

purpurascens, L051.

Pennella, 1078, 1089,

1090.

Pericharax, 731, 758,

768, 798, 800.

cartert, 735, 778.

— heteroraphis, 735.

— homoraphis, 734,

735.

heteroraphis, 710, 735.

peziza, T10, 735,

780.

polejaevi, 739.

Pericrocotus

brevirostris, 814.

Peripatoides

orientalis, 612, 613.

suteri, 611.

woodwardi, 611, 612.

Perisama, 348.

barnest, 347, 867.

Peristomella, 521.

Petalognathus

nebulata, 1085.

Petalosoma

amazonum, 981.

cultratum, 981.

Petalurichthys, 981.

Petaurus, 631.

Petralia, 488.

castanea, 519.

chuakensis, 463, 518,

519, 535.

dorsiporosa, 518.

ellerii, 519.

Japonica, 463, 518,

519

thenardii, 519.

undata, 519,

vultur, 519.

— armata, 463, 518,

519, 535.

INDEX OF SCIENTIFIC NAMES.

Petralia

(Escharella) bistnwata,

519. .

(Mucronella) castanea,

519.

(—) magnifica, 519.

(—) porosa, 519.

Petrostroma, 798.

schulzei, 740.

Phaéthon, 661, 1045.

Phalacrocorax, 395,

1045.

albiventris, 401, 402.

auritus, 401, 402.

bicristatus, 596.

carbo, 400.

dilophus, 402.

magellanicus, 400, 402.

pelagicus, 400-402.

penicillatus, 398-400,

402.

punctatus, 402.

urile, 400, 402.

vigua, 401, 402.

Phalloceros, 978-980.

caudomaculatus, 999,

1001, 1018.

Phalloptychus, 978, 980,

999

januarius, 998, 999,

1018.

Phaps

chalcoptera, 1056.

elegans, 1056.

Philander

laniger, 1094.

Philomedusa

vogtii, 969, 972.

Phlogeenas

cruentata, 1056.

luzonica, 1056.

Pheenicopterus, 670, 671,

681, 687, 703.

antiquorum, 1047.

Phonicosia, 508.

Phrynonax

JSasciatus, 1034.

Phyciodes

dora, 345, 367.

niveonotis, 847, 367.

phlegias, 346, 367.

Phyllium

erurifolium, 546.

Phyllobates

chocoensis, 1028.

pratti, 1026.

Phyllomedusa

hypochondrialis, 1025.

Phyllopezus

africanus, 1055.

Physa, 427.

| Plotus, 668,

Picolaptes

affinis, 1065.

Pieris

diana, 356.

limona, 356, 367.

Pitangus

sulfuratus, 1065.

Pitta

strepitans, 1065.

Plagiosternum, 956.

Planorbis, 427, 437.

Platalea

leucorodza, 1047.

Platemys

platycepha'a, 1104.

Platycercus

eximius, 1058, 1059.

Platypcecilus, 980.

couchianus, 1003,

1004.

maculatus, 1001, 1004, |

1005.

perugi@, 1014.

tropicus, 1013.

Platystethus

cultratum, 379.

Plectroninia, 740, 798.

deansti, 741.

halli, TAL.

hindet, 741.

| Plegadis

Jalcinellus, 651, 1047.

691,

695, 703.

anhinga, 396, 397, 654,

655, 702.

levaillanti, 655.

Plumatella, 429, 4381,

436, 488, 448.

coralloides, 449, 450.

emarginata, 452-454.

— muscosa, 435, 441, |

452-455, 457.

— spongiosa, 453, 455.

fruticosa, 449, 452-

454.

fungosa, 427, 449-452, |

454

== coralloides, 437,

441, 449-451, 454,

457.

polymorpha, 450, 452,

454,

punctata, 452.

repens, 450, 452-454,

457.

spongiosa, 455.

Podargus, 1065, 1069,

1072.

cuviert, 1062.

nacunda, 650.

693, |

XXXV

Podica

| senegalensis, 1054.

| Podiceps

cristata, 1044,

| minor, 1044.

Podoa

surinamensis, 1054.

Peecilia, 981, 999, 1002.

amates, 1013.

amazonica, 1006.

| branneri, 978, 1005,

1007.

couchiana, 1003.

cuneata, 1010, 1013.

decemmaculata. 1000.

dominicensts, 1015,

1017.

| elongata, 1018.

| _feste, 996,

gracilis, 1000.

heteristia, 1007, 1008.

lineolata, 1011.

| maculata, 1004.

melanogaster, 1015.

|. melanopleura, 988.

multilineata, 1011.

occidentalis, 1013.

pare, 978, 1005, 1006,

1009.

petenensis, 1012.

picta, 1005, 1007-1009,

1018.

pittieri, 997.

presidionis, 996.

reticulata, 1008.

retropinna, 997.

salvatoris, 1013.

schneideri, 1005.

sphenops, 1012.

spilonota, 1013.

spilurus, 10138.

surinamensis, 1006.

tenuis, 1013.

unimaculata, 1006.

vittata, 1014.

vivipara, 978,

107-1009.

— pare, 1006.

| Peciliopsis, gen.

996. 980, 1010.

isthmensis, 996-998,

1018.

| dutzi, 996.

| pittiert, 996, 997.

pleurospilus, 996,

997

presidionis, 994, 996.

retropinna, 996, 997.

Pceocephalus

Fuscicapillus, 1094.

meyert, 1058.

1008,

noy.,

XXXV1

Polejna, 793.

telum, 774.

uter, 793.

Pollicipes, 840, 842, 852

957, 958, 941, 947.

aucklandicus, 841, 847, |

848.

rigidus, 946.

Polyborus, 661.

Polychrus

gutturosus, 1032.

Polyodontophis

bivittatus, 428.

Pomacentrus

inornatus, 3870, 873:

genkinsi, 370, 371.

ntomatus, 370.

Porella

acutirostris, 503.

plana, 603.

Poricellaria, 492.

Porina

coronata labrosa, 511.

Porphyrio

porphyrio, 1052, 1053.

Porzana

carolina, 1052.

Potamon

bombetokense, 916, 919.

edule, 924.

fluviatile, 923-925.

goudoti, 919-921, $28,

924.

grandidieri, 916, 919.

humbloti, 916, 919.

madagascariense, I15— |

917, 921, 923-925.

methuent, 920, 923

924, 982.

(Parathelphusa)

tongilense, 925.

(Potamon) goudoti,

915, 920.

(--) madagascariense,

915, 916.

(—) methueni, 915,

920:

(—) pittarellii, 925.

(Potamonautes) de-

pressum,s 925.

Precis

iphita, 614.

Presbytis

entellus, 1108.

Priapella, gen. nov.,

992, 979.

bonita, 993, 994.

Priapichthys, gen. nov.

991, 979, 995, 995.

annectens, Y91, 992

S94,

INDEX OF SCIENTIFIC NAMES.

an= |

Priapichthys

episcopt, J92.,

nigroventralis, 992.

parismind, 991, 992.

tridentiger, 992.

turrubarensis, 992.

Procellaria, 1044.

Prosycum, 793.

primordiale, 726,

simplicissimum, 793.

Proteus, 403,

Protopterus

ethiopicus, 542.

Pseudocordylus

microlepidotus, 548,

1091.

Pseudoflustra

solida, 518.

Pseudolabrus

inscriptus, 371.

Pseudomonacan thus

ayraudi, 372.

paschalis, 372, 373.

Pseudopeecilia, gen. nov.,

995, 980.

Teste, 994, 996.

Pseudotantalus, 661.

ibis, 654, 702, 1046.

Pseudoxiphophorus,

ioe

bimaculatus, 993, 994.

Jonesii, 993.

pauciradiatus, 993.

terrabensis, 993.

Pseudupeneus

multifasciatus, S73.

Psophia

erepitans, 1058.

leucoptera, 1053.

obscura, 1053.

Pterocella, 483.

Pterocles

alchata, 1056.

Pteroglossus

inseriptus, 1063.

Pteronymia

fumida, 341, 367.

godmani, 341.

simplex, 341.

Pteroplax, 955, 956.

Ptilorhis, 651.

Ptychobothrium

belones, 552.

Ptyonius

estocephalus, 959.

Pudu

pudu, O91.

Pygosceles

papua, 1091.

Pyrrhula

nepalensis, 1091.

Querquedula

castunea, 1047.

Rallus

celebensis, 1052.

longirostris, 1052.

maculatus, 1052.

Rana

capito, 1092.

holsti, 1028.

palmipes, 1026.

Raphiceros

melanotis, 539.

sharpet, 539.

Raya, 960.

Recurvirostra

avocetta, 1050.

Retepora, 523, 529.

axillaris, 527.

cellulosa, 489, 503,

524,

contortuplicata, 524.

denticulata, 463, 526,

586.

elongata, 524.

Jormosa, O24.

gelida, 524, .

hippocrepis, 524,

hirsuta, 463, 528, 524,

527.

imperatt, I24.

ancrustata, O24.

inordinata, 524.

Jjermanensis, 463, 526.

lapralicides, 524.

monilifera, 524, 525.

— wmbonata, 524,

536.

novezealandie, 524.

philippensis, 527.

procellana, 525.

— laxa, 525.

producta, 463, 525,

536.

stnwosa, 524.

solanderia, 524.

tessellata, 524.

tubulata, 468, 526.

wmnbonata, d24.

Reteporella

dendroides, 526.

| Rhabdodermella, 793.

nuttingt, 784, 793.

Rhamphastos

discolor, 1068, 1064.

Rhamphoceelus

brasilius, 1065.

Rhawphostomella, 520.

Rhea, 663.

americana, 1043,

INDEX OF SCIENTIFIC NAMES.

Rhinochetus, 645, 651,

Jubatus, 1054. 1092.

Rhinthon

luctatius, 361, 367.

Rhizodopsis, 957, 958,

962.

Rhizodus, 957, 958, 962.

Rhyacornis

fuliginosa, 814,

Rhynchea

capensis, 1059,

Rbynchosaurus, 628,

Rhynchotus, LO5L.

rufescens, L050.

Rbynchozoon

bispinoswin, 523.

longirostre, 523.

profundum, 525.

— laminatum, 463,

523, 536.

Rhytidoceros

undulatus, 1061.

Rollulus

roulroul, 1052.

Safole, 374.

Salarias

arenatus, 372.

Salicornaria, 404.

Salicornia, 494.

Salmo

fario, 594.

gairdneri, 572, 576,

577, 605, 606, 610.

salar, 574, 579, 581,

582,607. - ‘

Sareorhainphus

gryphus, 1091,

Sauropatis

chloris, 1060.

sancta, 1060.

sordida, 1060.

vagans, 1060.

Savignyella, 482,

Sceorhynchus

ruficeps, 814.

Scalpellum, 840, 851,

946, 947.

sp., 82.

aucklandicum,

842.

cret@, 937, 941, 942.

magni, 842, 945.

michelottianum, 853.

— gassinensis, 853.

quadratum, 853.

subplanum, 842, 848,

854.

tuberculatum, 937, 938,

9403, D44.

841,

Sealpellum

| wulgare, 945.

| zancleanum, 8d0.

(Arcosealpellum) wngu- |

| latwm, 850, 853, 854. |

| Sceloporus

spinosus, 1092.

| Schismopora, 509, 510,

| Schizoblastus, 900,

Schizoporella, 459.

| auriculata, 505, 515,

biaperta, 509, 508.

| biturrita, 520.

| cecil, 508.

| cerustacea, 520.

galeata, 505, 506.

harmsworthit, 50d.

levigata, 506.

levata, 506.

| linearis, 504, 505, 515.

| longirostris, 505.

| marsupifera, 506.

montferrandt, 462,

506.

nivea, 462, 502-506,

1 a8, 687.

— wasinensis, 504, 506.

pachnoides, 506.

pertusa, 462, 502.

polymorpha, 520.

ridleyi, 505.

rimosa, 506.

sanguinea, DUD.

stmplex, 50d.

spongites, SO.

striatula, 507.

subimmersa, 515,

triangula, 615.

tuberosa, 520.

tumida, 506.

unicornis, 462, 501,

504, 505.

— errata, d0d.

venusta, DUG.

viridis, dV.

vulgaris, 50d.

Schizotenia

haymanni, 867.

Schultzierinus, 898.

Scillwlepas, 937, 938.

carinata, 947, 948.

ornata, 948.

parone, 948,

Scincosaurus

crassus, 959.

Sciurus

| saltuensis bonde, 814.

Sclerodomus, 522,

| Scleromochlus, 651.

| taylori, 629, 632.

Scoliocrinus, 897, 898.

XXXVI1

Scolopax

rusticula, 1Od4.

Scopus, 645, 646, 648-

652, 661, 671-674,

676, 682-709, 703.

umbretia, 650, 699,

701, 1045.

Seruparia

chelata, 466.

diaphana, 482.

Serupocellaria, 459, 481.

annectans, 477.

bertholettit, 474.

cervicornis, 462,

477, 554.

ciliata, 477.

clypeata, 477, 480.

delilii, 474. 477.

diadema, 477.

jerox, 462, 474, 476,

534.

incurvata, 476, 477.

inermis, 473, 475.

jolloisit, 473, 474, d34.

macandret, 462, 474,

476-478, 534.

mansueta, 474, 476,

479.

marsupiata, 475.

minuta, 77.

obtecta, 474, 477, 479.

occidentalis, 474.

ornithorhynchus,

477.

pilosa, 462, 478, 534.

portert, 474,

pusilla, 478.

reptans, 474.

scabra, 473-475.

scruped, 477, 478.

seruposa, 474.

— dongolensis, 474.

serrata, 475, 479.

smitti2, 473.

varians, 479.

WasiNensis,

534.

Seuticella, 483.

Seypha, 704.

coronata, 793.

ovata, 769.

Scyphognathus, 629, 631.

Selenidera

maculirostris, 1063,

1093, 1095, 1096.

Serpentarius, 661, 1048,

1069.

Sertularia, 494.

Seymouria, 955.

Simia

satyrus, 1092,

474,

462, 479,

XXXVIll

Siniopelta, 510.

Siphonactinia

boeckii, 966.

tricapitata, 966.

Siren, 403.

Sistrurus

miliarius, 10938.

Sitta

cinnamomeiventris,

814.

Smittina, 461, 515.

sp., 463, 514.

longtrostris, 514.

nitida, 513.

oculata, 514.

trispinosa, 508.

— bimucronata, 513.

— nitida, 518.

— protecta, 468, 513.

— spathulata, 463,

513.

tropica, 468, 514.

Solea, 960.

Solenidium, 794.

mitidum, T94.

Soleniscus, 794.

loculosus, 794.

Solenula, 794.

coriacea, 794.

Sorex

araneus, 824, 825, 838, |

839.

— araneus, 825.

— castaneus, 823-826,

828, 839.

— tetragonurus, 825,

828, 838, 839.

grantti, 822, 824-826, |

Spongia

inflata, 748, 769, 774.

nivea, T7193.

panicea, 769, 775.

pulverulenta, 769.

urceolus, T61.

Spongilla

lacustris, 427, 4389,

973.

Stagonolepis, 629.

Starncenas

cyanocephala, 1056.

Steatornis

caripensis, 1062.

| Steganoporella

minutus, 826-828, 838. |

Speotyto

cunicularia, 1062.

Spherium

(Cyelas), 427.

Spheniscus

demersus, 1044, 1085.

Sphenodon, 632.

Sphenophorina

singularis, 761, 794.

Sphenophorus

singularis, 794.

Spilotes

pullatus, 425,

Spongia, 794.

ananas, 769.

botryoides, 721.

ciliata, 704, '745.

complicata, 721.

compressa, 704, 760.

coriacea, 725.

coronata, 7495.

foliacea, 760.

| Styriodes, gen. nov., 361.

alveolata, 499.

buskit, 498, 501.

connexa, 499.

lateralis, 499.

magnilabris, 462, 468, |

489, 498-500, 536.

simplex, 499.

sulcata, 199.

truncata, 499.

tubulosa, 499.

Stenorhina

degenhardti, 1036.

| Stentor, 428.

| Stilesia, 552.

Stirparia, 458, 465.

annulata, 468, 469.

dendrograpta, 462,468—

471, 533.

exilis, 462, 468, 471,

DSoe

glabra, 468, 471.

zanzibariensis, 462,

469, 470, 534.

Streptoconus, 791, 794.

australis, 744, 745, 794.

Streptophorus

atratus, 1021, 1034.

Stringops, 1059.

habroptilus, 1058.

Strix

castanops, 1091.

flammea, 651,

1062.

Strongylopora, 483.

Struthio, 679, 1048.

lyco, 361, 367.

Sula

bassana, 395, 1045.

capensis, 1085.

Sycaltis

conifera, 743.

glacialis, 753.

leuconides, 784.

ovipara, 782.

perforata, 786.

1061,

INDEX OF SCIENTIFIC NAMES.

Sycaltis

testipara, 782.

Sycandra, 777, 798, 802.

alcyoncellum, 746.

— virgultosa, 749.

ampulla, 'TA4.

— alopecurus, 744.

— petiolata, 747.

arborea, 746.

arctica, T44.

— maxima, TAT.

— polaris, 747.

aspera, 744.

barbadensis, 745.

borealis, 745.

capillosa brevipilis,

759.

— lengipilis, 760.

ciliata, 745.

— lanceolata, 746.

— ovata, 747.

clavigera, 759.

coacta, 745,

compressa, 760.

— clavigera, 759.

— foliacea, 760.

— lobata, 760.

— pennigera, 760.

coronata, 745.

— commutata, 745.

— tubulosa, 749.

elegans, 745.

— formosa, 746.

— tessellata, 749.

glabra, 764.

— ensata, 763.

— rigida, 764.

helleri, 746.

humboldtii, 746.

hystrix, 764, 765.

inconspicua, 746.

lobata, 760.

parvula, 747.

pennigera, 760.

quadrangulata, 747.

— tesseraria, 749.

quadrata, 748.

ramosa, 748, 791.

ramsayt, 748.

raphanus, 748.

raphanus proboscidea,

747.

— procumbens, 747.

— tergestina, 748.

schinidtii, 748.

setosa, 748.

staurifera, 748.

tabulata, 748.

tuba, 749.

utriculus, 749.

villosa, 749.

INDEX OF SCIENTIFIC NAMES.

Sycandra

villosa impletum, 746.

Syeantha

tenella, 744, 745, 748, |

Use

Sycarium

ampulla, 744, 795.

rhopalodes, 760.

vesica, 748.

villosum, T49.

Sycetta, 744, 758, 798,

801, 802.

asconoides, 743, 760.

conifera, 748.

cupula, 761.

primitiva, 743, 796.

sagittifera, 748.

stauridea, 753, 789.

strobilus, 761.

Sycidium

gelatinosum, 795.

Sycilla

chrysalis, 782.

cyathiscus, 782, 798.

cylindrus, 782.

urna, 782.

Sycinula, 790.

aspera, 795.

egedii, 770.

penicillata, 769.

Sycocystis

oviformis, 745, 795.

Sycodendrum

ramosum, 745, 795.

Syecodorus, 798.

hystrix, 764, 765, 796.

Sycolepis, 779.

incrustans, TSO, 79).

palvinar, 77).

Sycometra, 795.

compressum, 795.

Sycon, 704, 716, 720, 733,

TAZ, (43, 790, 757, |

758, 798, 802, 803.

alopecurus, 744.

ampulla, 744, 798,

antarcticum, 744, 796.

arcticum, 744.

asperum, 744,746, 769.

australe, '745, 794.

barbadense, 745.

boomerang, 710, 745,

803.

boreale, 745.

caminatum, 745.

carteri, 710, 745.

ciliatum, 745, 795,

796.

coactum, T45.

commutatum, 745,

compactum, 745,

Sycon

compressum, 760.

coronatum, 745, 798.

dendyt, 763.

eglintonensis, 745.

elegans, 745, 748, 789.

ensiferum, 713, 746,

ioe

formosuna, TA6.

gelatinosum, 710, 746,

78%, 795.

giganteum, 710, 746.

heileri, 746.

humboldtii, 746, 748.

impletum, 746.

inconspicuum, T46.

tncrustans, 746.

karajahense, 746.

kergquelensis, 740.

lambei, 746.

lanceolatum, 746.

lendenfeldi, 710, 747,

748.

lingua, 747.

longstaffi, T47, 791.

maximum, TAT. |

minutum, 747.

mundulum, 747.

munitum, 747. |

ornatum, 747.

ovatum, 747.

parvulum, 747. |

pedicellatum, T47.

petiolatum, 744, 747.

polare, TAT. |

proboscideum, 747.

procumbens, 747.

protectum, T47.

guadrangulatum, “47, |

748, 795.

ramosum, G48, 791,

795. |

ramsayt, 710, 748.

raphanus, 710, 744, |

748.

schmidti, 748.

schuffneri, 748 |

setosum, 710, 748. |

stauriferum, 748.

subhispidum, 748.

sycandra, 748, 791.

tabulatum, 748.

tenellum, 748, 795.

tergestinum, 748.

tessellatum, 749.

tesserarium, 749. |

tuba, 749. |

tubulosum, 749.

verum, 710, 718, 749.

villosum, 749.

virgultosum, 749.

XXX1X

Sycon

(Acyoncelllum) gelati-

nosum, 787.

Syconella

proboscidea, 747.

quadrangulata, 747,

795.

Syeophyllum

lohatum, 799.

Sycorrhiza

corallorhiza, 721.

coriacea, 799.

Sycortis

levigata, 761, 79d.

lingua, TAT.

sycilloides, 753.

Sycortusa

levigata, 761, 796.

Sycothamnus

Sruticosus, 734, 796.

Syculmis, 798.

synapta, 783.

Sycum

alopecurus, T44.

areticum, TA4.

ciliatum, 796.

clavatum, 749.

giganteum, 745.

lanceolatum, 746.

lingua, 760.

ovatum, 747.

petiolatum, 744, 747.

primitivus, 196.

procumbens, 747.

tergestinum, 748.

Sycurus, 796.

primitivus, 796.

Sycute, gen. nov., 763,

798.

dendyi, 757, 763.

Sycyssa, 798.

huxleyi, 767.

Sylvia

nisorta, 1093.

Sylvicapra

coronata, 1090.

Synnotum

aviculare,

466.

contorta,

532.

pembaensis, 462, 465,

532.

Synute, 798.

pulchella, 711, 764.

462, 465,

462, 466,

Tenia

bifaria, 570.

cucumerina, Ho.

genette, d49.

Tenia

marchali, 568.

platydera, 549.

Tainsia, 1099,

Talegalla

lathami, 1051.

Tanagra

sayaca, 1065.

Tantalus, 688-691, 693- |

698, 7U3.

loculator, 1046.

Tarroma, 796.

canariense, 796.

Tarropsis, 796.

coriacea, 796.

Tarrus

densus, 721, 796.

labyrinthus, 725.

reticulatus, 723.

Tatusia, 1099.

Taxocrinus, 911.

Teichonella

labyrinthica, 762.

prolifera, 734, 762, 796.

Teichonopsis, gen. nov.,

761, 796, 798.

labyrinthica, 711, 762.

Telemiades

mnemon, 857, 367.

Tenthrenodes, 796.

antarcticus, 796.

scotti, 760.

Tephreops, 368, 369.

richardsontt, 370.

Terpsephone

paradisi, 614.

Tethrenodes

artarcticus, 744.

scottt, 744.

Tetrabothrium, 562.

Netrarhynchus

tetrabothrius, 551.

Tetrastemma, 428.

Teuthis

umbra, 373.

Thairopora

manillaris, 462, 497.

Thalamoporella, 501.

mamillaris, 497.

Thalassoma

purpurewm, 373.

Thanatophis

sutus, 1037.

Thaumantias

(Phialidium), 970.

Thecla

amphrade, 355, 367.

dolylas, 353.

guapila, 354, 367.

hypocrita, 302, 367.

Boo 367,

leos, 358,

INDEX OF SCIENTIFIC NAMES,

Thecla

melma, 894, 367.

politus, 305.

subflorens, 353, 367.

Thecotetra, 7388.

loculosa, 796.

Thinocorys

sp., 1055.

—rumicivorus, 1O92.

Thylacinus, 892.

Tima, 970.

Tinnuneulus

alaudarius, 1049.

Titanolepas, subgen. noy.,

943.

tuberculata, 938, 943,

944, 946-948.

Tomeurus, 979.

gracilis, 98.

Toxus, 980.

riddlei, O02.

Trachinus

draco, 636.

Trachyboa

houlenger?, 1034, 108

gularis, 1034.

Trachypoma

macracanthus, 373.

Trematosaurus, 956,

Tribonyx .

mortiert, L052.

Trichogaster

Jasciatus, 542.

Trichoglossus, 1059.

nove-hollandie, 1058.

Trichogypsia, 767, 798,

805.

incrustans, 779, 780,

795.

villosa, 779, 780, 795.

Triclaria

cyanogastra, 1104.

Tripleurocrinus, 898.

Trogon

atricollis, 1063.

puella, 1068.

Tropidonotus

vibakari, 423, 424,

vittatus, 424.

Trypostega

venusta, 462, 506, 507.

Tubucellaria

cereoides chuakensis,

462, 512.

Susiformis, 462, 512.

opuntioides, 494.

zanzibariensis,

512.

Tubulipora, 461.

Turacus, 639.

corythaia, 1057,

462,

Turacus

donaldsont, 641.

Jischeri, 641.

merian, 641.

Turdus

tristris, 1066.

Turnix

dussemiert, 10d).

Tyrannus

melancholicus, 10698.

Unio, 428.

Upupa

epops, 1061.

Urceolipora 528.

Urocordylus

reticulatus, 959.

wandesfordii, 959.

Urogalba

puradisea, 1063.

Uromys

ae she

Ute, 754, 798

oe ae.

capillosa, 759, 764.

chrysalis, 782.

ensata, 763.

glabra, 764.

papillosum, 786.

rigida, 764.

spencert, 764.

spiculosa, 711, 764.

syconoides, 711, 764.

utr ees ee 764.

viridis, 786, 787.

Utella, 764.

hystriz, 796.

eee gen. noy., 766,

58, 798.

gente 766.

Vanellus

vanellus, 1050.

Vespa

cincta, 618.

Victorina

steneles stygiana, 348,

Vittaticella,

505.

buskei, 485.

contet, 484.

elegans, 462, 484, 485,

533.

elegans zanzibariensis,

462, 484, 485, 533.

(Catenaria), 484,

Viverra

genetta, 549,

465, 483,

Vivipara

(Paludina), 428.

Vorates

cotiso, 364, 367.

Vorticella, 428.

Vosmaeria

corticata, 776, 777.

gracilis, 760, 76.

Vosmaeropsis, 798, 803.

connexiva, 756.

cyathus, 756.

dendyi, 711, 756.

depressa, 711, 759.

macera, 711, 75d.

primitiva, 711, 756.

sericatum, 756.

wilsoni, 711, 735.

Vulpes

chama, 392.

(Alopex) lagopus, 391.

Wagnerella, 797.

INDEX OF

SCIENTIFIC

Wagnerella

borealis, 797.

Xanthichthys

lineopunctatus, 373.

Xenopeltis

wnicolor, 415, 417.

Xenopus

miillert, 1093.

Xenorhynehus, 688, 692-

694, 696, 703.

Xiphophorus, 978, 980.

bimaculatus, 993.

brevis, 1005.

gracilis, 998.

guenthert, 1004, 1005,

helleri, 1004, 1008.

Jjalape, 1004.

montezume, 1004,

Xiphophorus

rachovti, 1005.

strigatus, 1004.

NAMES.

Yphtbima

ceylonica, 614.

Zamenis

gemonensis, 933.

grahani, 1092.

Zaudidura

carolinensis, 1056.

Zeugmatolepas, gen. nov.,

9338.

eret@, 942.

macklert, 937,

941, 942, 948.

Zonurus

giganteus, 548.

Zy gonectes

atrilatus, 984.

brachypterus, 985.

inurus, 985.

938,

INDEX

ILLUSTRATIONS.

Acanthistius fuscus, Pl. LV. p. 368.

Acmepteron poasina, Pl. LIT. p. 839

Adeonella platalea, Pl. LXXIIT. p. 458 ;

Fig. 82, p. 530. ;

Adeonellopsis crosslandi, Pl. LXXIII.

p. 458.

Aetea unguina, Pl, LXIV. p. 458.

truncata, Pl. LXIV. p. 488.

fix sponsa, Fig. 181, p. 1048.

Agalychnis calcarifer, Pl. CII. p. 1019.

spurrelli, Pl. CIII. p. 1019;

Fig. 177, p. 1025.

Alfaro amazonum, Fig. 169, p. 993.

Amblystoma tigrinum, Figs. 75, 76,

pp. 406, 407.

Anadia vittata, Pl. OVIT. p. 1019.

Anampses pulcher, Pls. LVILII., LIX.

. 68.

Das breviceps, Pl. CVIT. p. 1019.

Anoplotherium, Figs, 150-157, pp. 879—

882, 886, 887, 890.

Archonias intermedia, Pl. LII. p. 539.

Ardea, P|. UXXXIII. p. 644.

Ate canace, Pl. LIV. p. 339.

Baleniceps rex, Pls. UXXX., LXXXI.

p. 644; Figs. 119-132, pp. 646-

695.

Balenoptera borealis, Pl. CXII.

p. L073.

brydei, Pls. C1X.-CXIII. p. 1073.

Batrachiderpeton lineatum, Pls. XCV1.,

XCVII. p. 949; Figs. 162-166,

pp- 950-954.

Belonesox belizanus, Fig. 169, p. 990.

Bicellaria chuakensis, Pl. LXVIII.

p- 458.

Bicidium equoree, Pl. XCVIIL. p. 963.

—— parasiticum, Pl. XCVIII. p. 963.

Bifaxaria vagans, Pl. LXV. p. 458.

Bolla machuca, Pl. LY. p. 339.

sodalis, Pl, LIV. p. 339.

Brachyrhaphis rhubdophorus, Fig. 169,

990.

Brettia tropica, Pl. LXLV. p. 458.

Browniella africana, Pl. LXXIX.

p- 619.

Bubalis caama_ selbornei, Fig. 135,

p- 820.

Bufo hypomelas, Pl. CII. p. 1019.

Bugula robusta, Pl. LX1X. p. 468.

Bugulopsis peachii, Pl. LUX1X. p. 458.

Butleria lethea, Pl. LIV. p. 339.

lysis, Pl. LIV. p. 339.

Butorides cyanurus, Fig. 120, p. 650.

Caberea ellisit, Pl. LXIX. p. 458.

Cecilia intermedia, Fig. 174, p. 1020.

palmeri, Fig. 175, p. 1021.

Calantica (Titanolepas) tuberculata,

EAROXC VER peor

Cancroma, Pl. LXXXII. p. 644.

Canda retiformis, Pl. LXIX. p. 458.

Canis antarcticus, Figs. 70, 71, 73, 74,

pp. 889, 387, 389, 390.

latrans, Figs. 70-72, 74, pp. 885,

387, 388, 390.

Carystus subrufescens, Pl. LIV. p. 359.

Catargynnis dryadina, Pl. L. p. 339.

Catenaria diaphana, Pl. LXILYV. p. 458.

Cellaria gracilis, var. _ tessellata,

Pl. LXVII. p. 458.

wasinensis, Pl. LX VII. p. 458.

Ceraterpeton vreticulatus, Hig. 167,

p. 958.

Cervus axis, Fig. 134, p. 816.

canadensis, Fig. 184, p. 816.

dama, Fig. 134, p. 816.

— claphus, Fig. 134, p. 816.

xliv

Cervus eustephanus ?

pp. 548, 544.

leucurus, Fig. 134, p. 816.

muntjac, Wig. 134, p. 816.

unicolor, Fig. 134, p. 816.

Charis subtessellata, Pl. LILI. p. 339,

turrialbensis, Pl. LILI. p. 339.

Chauna chavaria, Big. 179, p. 1041.

Chlidonia cordieri, Pl. LXY. p. 458.

Cinosternum spurrelli, Pls. CV., CVI.

p. LOL9.

Cnesterodon decemmaculatus, Fig. 172,

p. 1001.

Cobalopsis latonia, Pl. LIV. p. 339.

Cobalus lateranus, Pl. LITT. p. 339.

laureolus, P\. LIV. p. 339.

— nigrans, Pl. LIV. p. 339.

pindar, Pl. LIV. p. 339.

Corrachia leucoplaga, Pl. LILI. p. 3839.

Cyclosemia subcerulea, Pl, LIV.

. 039.

Cynocephalus anubis, Pls. LXXXVILI.-

LXXXIX. p. 809.

Figs.

Dasypus. See Ewphractus,

Dendrobates aurotenia, Fig. 178,

p- 1029,

— paraensis, Fig. 178, p. 1029.

—— tinctorius, Fig. 178, p. 1029.

— yar. chocoensis, Pl, CLY.

p. LO19.

war, caer, I OIL

p- 1019.

Diagram of Peroneal Muscles in Birds,

Vig. 190, p. 1071.

Diplodidymia complicata, TAL, JLRS WIE,

p- 458; Fig. 79, p, 491.

Diplopylidium genette, Figs.

pp. 560, 561, 563, 565, 566.

Dipylidium caninum, Fig, 88, p. 999.

dongolense, Figs. 85-87,

pp. 550, 553, 594, 556.

Dissura, Pl, LXXXI. p. 644,

90-94,

309.

1OL9,

IPM

Echelatus lucina, Pl. LLY. p.

Hlaps microps, Pl, CVIILI. p.

Enantia lua costaricensis,

p. 399,

Epinosis angularis, Pl. LMI.

Eresia cela, Pl. L. p. 339,

sticta, Pl, L. p. 339,

Hubagis hecuba, Pi. LI. p. 339.

Eudocimus longirostris, Fig.

p. 650.

Eugonodewm edicnemi, Figs. 141-148,

pp. 862, 864-866, 868, 870-873.

Euparkeria capensis, Pls. UXXV.,

LXXVL., LXXVIII., LXXIX. p. 619.

Eupetoieta poasina, Pl. LI. p. 389.

LI.

p. 039,

1120)

INDEX OF ILLUSTRATIONS.

Huphractus — seacinctus, 195,

pawlO2

villosus, Figs. 193, 194, pp. 1109,

1101.

Euptychia agnata, Pl. L. p. 339,

drymo, Pl. L. p. 339.

Eurygona leucon, Pl. LI. p. 339.

leucophryna, Pl. LIL. v. 339.

matuta, Pl. LI. p. 339.

mystica, Pl. LI. p. 339.

Evotomys alstoni, Fig. 137, p. 829.

Fig.

Falco peregrinus, Fig. 182, p. 1049.

alga hermione, P|. LIV. p. 389.

Farcimia oculata, Pl. LXVITI. p. 458.

Hlabellaria (Menipea) cuspidata,

Pl. LXIX. p. 458.

Fredericella sultana, Pl, UXIII.

p- 426.

Gadus eglefinus, Fig. 98, p. 578.

Gambusia dominicensis, Pl.

p. 977 ;_Fig. 169, p. 990,

gracilior, Pl. XOIX. p.

Fig, 168, p. 983.

holbrookit, Fig. 169, p, 990.

nicaraguensis, Fig. 168, p. 983.

nigropunctata, Fig. 168, p, 983.

oligosticta, Pl. XCIX. p. 977;

Fig. 169, p. 990.

punctata, Hig, 168, p. 983.

senilis, Big. 168, p. 983.

wrayi, Pl. XCIX. p.

Fig. 168, p. 983.

Gemellipora protusa, Pl. LXX. p. 458,

Girardinus metallicus, Fig. 172,

p. 1001.

Girellops nebulosus, Pl. LVIT. p, 368.

Gryponyx africanus, Pl, UXXIX,

p. 619.

XCIX.

O77 ;

ime

Heliconius galanthus, ab. subrifescens,

Pl. LIL: p. 339.

Hermathena oweni, P\. LILI. p. 339.

Heternadria formosa, Fig. 170, p. 994.

Hexelasma aucklandicum, Pl, uXXXV.

p. 840; Figs. 139, 140, p. 848.

Holoporelia albirostris, Pl, LXXIIT,

p. 458.

Homalocranium coralliventre, Pl. CV ILL,

p. 1019.

Hyla maxima, Fig, 176, p. 1028.

Hypocrinus piriformis, Pl, XC, p. 894;

Fig. 160, p. 907.

schnetderi, Pl, XC. p.

Figs. 158, 159, pp. 901, 904,

894 ;

Intromittent Organ in the Peeciiiines,

Figs. 168-173, p. 983, ete.

Tthomia bolivari, Pl. L. p. 339.

INDEX OF ILLUSTRATIONS,

Kuhlia cerulescens, Big. 68, p. 376.

—— humilis, Fig. 69, p. 380.

—— mutabunda, Pl. LVI. p. 368.

splendens, Fig. 69, p. 380.

Labrichthys fuentesi, Pl. LVIII. p. 368.

Lebistes reticulatus, Fig. 1738, p. 1009. |

Lechriorchis inermis, Pl. XOILT. p. 933.

Lepralia feegensis, Pl. UXX. p. 458.

turrita, Pl. LX XIII. p. 458.

wasinensis, Fig. 81, p. 517.

Leptognathus spurretli, Pi,

p- L019.

Leptoptilus, Fig, 131, p. 686.

Lerodea rupilius, Pl. L1V. p. 389.

Limia arnoldi, Pl. CI. p. 977.

caudofasciata, Pl. OL. p. 977.

heterandria, Pl. Cl. p. 977.

—— nigrofasciata, Pl. CL. p. 977.

ornata, Pl. Cl. p. 977.

versicolor, Fig. 173, p. 1009.

CVIII.

Megalema virens, Fig. 187, p. 1064. |

Megistias xantho, Pl. LIV. p. 339, |

Membranipora armata, Pls. LXVIL., |

LXXI. p. 458.

savartit, Pl. LXXI. p. 458.

Menipea buskii, Pl. LXIX. p. 458.

cyathus, Pl. UXIX. p. 458.

patagonica, Pl. LXIX. p. 458.

Mesosenia albipuncta, Pl. LILI. p. 359. |

Mesosuchus browni, Pls. LUXXVIIL.,

LXXIX. p. 619.

Microtus agrestis exsul, Fig, 138, p. 832.

-macgilivrati, Fig. 1838, |

p. 892.

Mollienisia sphenops, Fig. 173, p. 1009.

Molo apelia, Pl. LILI. p. 839.

nebrophone, Pl. LLY. p. 339.

Nannopterum harrisi, Pl. UXT. p. 393.

Napeogenes hemisticta, Pl. L. p. 339.

Nyeticorax, Fig, 131, p. 686.

gardeni, Fig. 180, p. 1046.

Enus nausiphanes, Pl. LIV. p. 339.

Oncorhynchus gorbuscha, Fig. 116,

p. 603.

keta, Fig. 117, p. 605. |

—— hkisutch, Figs. 112-115, pp. 599- |

602.

nerka, Figs. 103-108, pp. 586,

590, 592, 593. |

tschawytscha, Figs. 95, 96, 102, |

109-111, pp. 574, 575, 585, 595,

597, 598.

Oochoristica, Fig. 149, p. 875.

Ornithosuchus taylori, Pl. LXXVII. |

p- 619. |

—— woodwardi, Pls. DLXXVIL.- |

LXXIX. p. 619.

xlv

Osteomalacia in a young Anubis

Baboon, Pls. LXXXVII.-LXXXIX.

p- 855.

Osthimosia zanzibariensis, Pl. LX XIII.

p- 408.

Palemon hildebrandti, Pl. XCII. p. 914.

Paludicella articulata, Pl, LXII. p. 426.

Parathelphusa tridentata, Fig. 161,

HO23:

Parotia lawesti, Fig. 189, p. 1066.

Pavo nigripennis, Fig. 192, p. 1098.

Peachia quinguecapitata, Pl. XCVIII.

963.

Pelecanus, Pl. UXXXIII, p. 644.

Perisauma barnesi, Pl. LI. p. 389.

Peroneal muscles in Birds, Figs. 179-

190, p. 1041 ete.

Petralia chuakensis, Pl. UXX. p. 458.

vultur, var. armata, Pl. LXX.

p. 458.

Phalacrocorax albiventris, Pl. UX1.

p. 993

auritus, Pl. LXT. p. 3938.

—— dilophus, P\. LXI, p. 393.

magellanicus, Pl, LXI. p. 395.

pelagicus, Pl. LXI. p. 393.

pemcilatus, Pl. LXI. p. 393,

—— punctatus, Pl. LXT. p. 393.

urile, Pl. LXTI. p. 3898.

vigua, Pl. LXT. p. 393.

Phatloceros caudomaculatus,

p. 977; Fig. 172, p. 1001.

Phalloptychus ganuarius, Pl. C. p. 977;

Fig. 171, p. 998.

Phyciodes dora, P\. LI. p. 339.

niveonotis, Pl. LILI. p. 839.

phlegias, P|. L. p. 339.

Pieris limona, Pl. LIL. p. 339.

Platycercus eximius, Fig. 185, p. 1059.

JPL. GL

Platypecilus maculatus, Fig. 172,

p. LOOL.

Plotus, Pl. LXXXIII. p. 644.

Plumatella emarginata, var. juscosa,

Pl. LXITI. p. 426.

Sungosa, var. coralloides, Pl. UX111.

p. 426.

repens, Pl. LXIIT. p. 426.

Podargus nacunda, Fig. 120, p. 650.

Pecilia pare, Big. 173, p. 1009.

-pieta, Pl. ©. p. 977; Fig. 173,

p- L009.

—— vivipara, Fig. 173, p. 1009.

Peciliopsis isthmensis, Pl, C. p. 977;

Fig. 171, p. 998.

presidionis, Fig. 170, p. 994

Pomacentrus inornatus, Pl. LVIII.

p. 368.

Porphyrio porphyrio, Fig. 183, p. 1053.

Potamon fluviatilis, Fig. 161, p. 928.

madagascariense, Fig. 161, p. 923.

xlvi

Potamon methuent, Pl. XCI. p. 914.

Priapella bonita, Fig. 170, p. 994.

Priapichthys annectens, Fig, 170, p. 994.

Pseudomonacanthus paschalis, Pl. LX.

p: 368.

Pseudopecilia feste, Fig. 170, p. 994.

Pseudoxiphophorus bimaculatus, Fig.

170, p. 994.

Pteronymia fumida, Pl. L. p. 839.

Tel,

Retepora LXXII.

p. 458.

monilifera, var.

LXXIT. p. 458.

producta, P). LXXII. p. 458.

Rhanphastos discolor, Fig. 188, p. 1064.

Rhinthon luctatius, Pl. LIV. p. 889.

Rhyncozoon profundum, var. laminatum,

Pl. LXXITI. p. 458.

denticulata,

umbonata, Pl.

Salmo gairdneri, Figs. 97, 118, pp. 577,

606.

salar, Figs. 99-101, pp. 579, 581, |

582.

Scales of Salmon, ete., Figs. 95-118,

pp- 574-606. :

Scalpellum subplanum, Pl. UXXXVI.

p- 840.

(Arcoscalpellum) ungulatum, PI.

LXXXVI. p. 840.

Schizoporelia nivea, Pls. LXX., LX XIII.

p. 458; Fig. 80, p. 505.

Scopus wmbretta, Pl. UXXXT. p. 644;

Figs. 120, 132, pp. 650, 695.

Scrupocellaria cervicornis, Pl. LXIX.

p. 458.

—=—= ORE:

p. 458.

jolloisii, Pl. LXIX. p. 458.

macaudret, Pl. LXVILI. p. 458.

pilosa, Pl. LXVIII. p. 458.

wasinensis, Pls. LXVIII., LXIX.

p. 458.

Selenidera

p. 1096.

Pls, LXVINI, LXIX. |

Tales, ht,

maculirostris,

INDEX OF ILLUSTRATIONS.

Skulls of Storks, ete., Pls. LXXXI.-

LXXXITII. p. 644.

Sorex araneus castaneus, Pl. LXXXIV.

p. 821.

grantit, Pl. LXXXIV. p. 821;

Fig. 136, p. 825.

Sponges, Phylogeny of

Fig. 183, p. 798.

Steganoporella magnilabris, Pl. LXXII.

p. 458.

Stirparia

p. 408.

-—— evilis, Pl. LXVI. p. 458.

zanzibariensis, Pls.

LXIX. p. 458.

Stringops habroptilus, Fig. 184, p. 1058.

Strix flammea, Fig. 186, p. 1061.

Styriodes lyco, Pl. LIV. p. 339.

Synnotum contorta, Pl. LXIV., p. 458.

penbaensis, Pl. LXLV. p. 458.

Calcareous,

dendrograpta, Pl. LXVI.

LXVIIL,

Tantalus, Pl. UXXXITI. p. 644.

Telemiades mnemon, Pl. LIV. p. 339.

Thecla amphrade, P|. LIL. p 359.

guapila, Pl. LIT. 3389.

hypocrita, P|. LI. p. 339.

leos, Pl. LIL. p. 339.

mela, Pl. LIT. p. 539.

—-— subflorens, P\. LAI. p. 339.

Victorina steneles, ab. stygiana, Pl. LI.

p. 339.

Vittaticella elegans, Pl. LXV. p. 458.

-——., var. zanzibariensis, Pl. LXV.

p. 458.

Vorates cotiso, Pl. LIM. p. 339.

Xenopeltis unicolor, Figs. 77, 78, pp. 418,

417.

Xenorhynchus, Pl. UXXXT. p. 644.

NXiphophorus hellert, Fig. 172, p. 100i.

Zeugmatolepas mockleri, Pl. XCIV.

p. 937. -

Zonurus giganteus. Pl. LXXIV. p. 548.

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STRUET.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.

1913.

PART III.

CONTAINING Paces 339 ro 860, witH 40 PuLaTEs

AND 73 TEXT-FIGURES.

SEPTEMBER 1913... =~ -. “©

PRINTED FOR THE SOCIETY,

SOLD AT ITS HOUSE IN REGENT’S PARK.

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Olas [Price Twelve Shillings.]

LIST OF CONTENTS.

1913, Part III. (pp. 339-860).

EXHIBITIONS AND NOTICES.

Page

The Sxcrrrary. Report on the Additions to the Society's Menagerie during the month of

March 1913 2... .... sass Bsa ie aime waavercietceiere Ere de anlastoddntoeiadac sos sees oe O42

Mr. H. J. Eiwss, F.B.S., F.Z.S. Exhibition of, and remarks upon, the head of an Asiatic

Wapiti. (Text-figs. 83 & 84.) ......... Boe Bis ike dio Shc lols ae Me hate onctalo rele ts eG erin 542

Mr. R. H. Burnu, M.A., F.Z.S. Exhibition of malformed antlers of an Axis Deer (Cervus

LIS) stares onus Rew seaes saicteiniois eioiete dosed ded caaiced so 36-06 elstnlae\olnis love loteenaena 545

Mr. E. G. Bouumnarr, F.Z.8. Exhibition of living specimens of the Leaf-Insect (Phylliwm

Crurifolium) ..20+...0.03- aia treorelexe iio setae: nai sie Wiolneleeeen anole Gis iejnefecsie cv eieteae -.. 546

Mr. E. G. Bovutrzncer, F.Z.8. Exhibition of a melanistic specimen of the Green Lizard _

(Lacerta viridis) ..c.secsecsse “oN opibooaconD OGus aya TelswaR anetlalatetenerepctonetererr eje basse OAG

Dr. R. Broom, C.M.Z.8. Exhibition and Description of a new Species of Golden Mole .. 546

Dr. R. Broom, C.M.Z8. etiaisad of the female and young of the §. African Lizard,

Zonurus giganteus. (Pl. LXXIV.).......... wie clobatei quate se tobetareteisiere 18 eiaislcletel ates 548

The SzcreTary. Report on the Additions to the Society's Menagerie during the month of

PNT OUD ccna rere eae ra Sod nce ne EEIOAD oth So's seueneiaeks aterete Shoo 5 oe esses. 813

The Rev. T. R. R. Steszine, M.A., F.RS., F.Z.8., notice of Prof. F. E. Schulze’s

‘Nomenclator Animalium’ .......... OARS Aero ECC aT CREDO Cocke Beer oS 814

The Liprartan. Dates of publication of early parts of the Society’s “Transactions” .... 814

Mr. R. E. Hotvine. Exhibition of a number of antlers, skulls, and phoiographs, illus- _

trating variations in the growth of the Antlers of Deer. (Text-fig.134.)............ 815

Mr. D. Seru-Surru, F.Z.8. Exhibition of the Eggs and Young of tne Mikado Pheasant

(Calophasis mthado) os-ceecccrevcccrsvereere cs eettennees aa venga gin ates sue ee 818

PAPERS.

26. New Species of Rhopalocera from Costa Rica. By Wiuttam Scuavs, F.ZS.

(Pls. L.-LIY.) eeerteeoreeeneenn ® @eceovce tC eChCeeeeee et ee oveseeeeeeeosevese e@oeeeecsece 339

97. A Collection of Fishes made by Professor Francisco Fuentes at Haster Island.

By C. Tarz Ruaan, M.A., F.Z.8S. (Pls. LV.-LX.) ccccee seca ce ee ceneeeenes Abaoos clas

98. A Revision of the Fishes of the Genus Kuhklia. By C. Tats Ruaan, M.A., F.Z.8.

(Text-figs. 68 & 69.) ecVeepeeecetreeet®Oce P®soooep ee ee oe Fe eee @Cs.oececrecece tee e@ @¢sece 374

29, The Affinities of the Antarctic Wolf (Canis antarcticus). als R. I. Pococs, F.R.S.,

E.L.S., F.Z.8., Curator of Mammals. (Text-figs. 70-74.) . ernaharereieneverere -. 882

Contents continued on page 3 2 Wetine

Sh tastes teat

DY Aso is vSi2 ell, Ik,

Horace Knight del.et lith. West, Newman chr.

(6p)

COSTA RICAN BUTTERFLII

PAS ASS Jel, Jbl,

Horace Knight del.et lith. West, Newman chr.

COSA IRUNCYAIN] © 12) 1O/ DIE IS. Ly aS).

IWASS), wue)ikSy, IM. IL.

e’

Ie 7

West, Newman chr.

Horace Knight del.et lith.

GOSIWA WRIGAIN IBUPaa RUE Is S.

Horace Knight del.et lith.

COSTA

TSU) AM Tes Wale) Ie

PIA. Sr lhe ele ee

West, Newman chr.

IBNAyS) US) hee}, IPM IL IDW/

West,Newman chr.

Horace Knight del.et lith.

COSTA RICAN BUTE RE iE SS:

PROCEEDINGS

OF TIE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

PAPERS.

26. New Species of Rhopalocera from Costa Rica.

By Wicu1am Scuavs, F.Z.8.

| Received February 17, 1913: Read March 4, 1913.]

(Plates L.-LIV.*)

SYSTEMATIC :—

Subfam. DanaInz.

Napeogenes hemisticta, sp. n.......

Pteronymia fumida, sp. n.

godmeni, nom. nov.

Ithomia Bolsa, Sp. We...-.. aseeeeee

Subfam. SaTyRiInz.

Huptychia drymo, sp. n. ............

5 agnata, Sp. N..........-..

Catargynnis dryadina, sp. Ni. ......

Subfam. HELIconInz.

Heliconius galanthus, ab. sub-

FPO AS OCIS,, B)D> TAG)\ Ye eeappaenareees5o5c

Subfam. NyMPHALIN”.

Hupetoieta poasina, sp. n. .........

WER ESUCISUZELE,| SPs IS vs eesseasss se eese

~, wether When 65 it

Phyciodes dora, sp. n. ......

Ay phlegias G.&S., Qin ;

Fa niveonotis Btl.

Pym.

fake

INDEX,

Page

346

Eubagis hecuba, sp. n.

Perisama barnesi, sp. n.

Victorina steneles, ab. stygiana,

ab. nov.

Fam. ERYcINID2.

Eurygona mystica, sp. 0. .....0...++

oD UGHEOIOS JVs We sce aoc oopo0s

op leucophryna, sp. N.......

op MAtULA, SP. Nl. ........+-s-

Mesosemia albipuncta, POE Te. cos soc

Hermathena oweni, sp. 0.

Corrachia, gen. nov. soe

leucoplaga, sp. Ibe.» de

Chari is subtessellata, sp. N..........

5 turrialbensis, SPSMeg. swans

Fam. Lycx#NID2.

Thecla hypocrita, sp. n. eats

5 GOTO PIDIHEOD, Ss. oosens ssp see

Sian gUCOSH SDs buccuemereeere aw Aaas

6p COATES SNo Do ono cab ess as cdbonc

cg) GROOT S Ds Wo. oon codossoseo>:

50 COM OURHOE SH, Do soon secace

* For explanation of the Plates see p. 367.

Proc, Zoou, Soc.—1913, No. XXIV, 24

340 MR. WILLIAM SCAAUS ON

INDEX (continued).

Page | Page

Subfam. Prerin x. | Lerodea? rupilius, sp.n. ......... 360

Archonias intermedia, sp. 0. ...... 355 | Simio des: eee TON g Goins plas oe

JEG PIS UOROME, D6 oasseaccncoeccsna BD | Rh nth oe ae athe Ae Ste ie 361

Acmepteron poasina, sp. n.......... 356 | 6 ae LOY ed OtUS; SD ee aia 362

Enantia lua costaricensis, subsp.n. 357 | ~?7“"S Se Sy SPs eens ;

| 5 aureolus, sp. n............. 862

Kam. ElEspeRriip =. | ap lateranus, SDs Weacoxsonodecy By

my FOOREIBIP, Fe Wo covecsscacpad a GhO2)

Telemiades mnemon, sp.n. .......... 357 | Cobalopsis latonia, sp. n. ......... 363

Nascus phintias, sp. n................ 3858 | Ginus nausiphanes, sp. un. ......... 363

Cyclosemia subcerulea, sp... ... 3808 | Megistias xantho, sp. n. ..-... 0.0... 363

Echelatus lucina, sp.n. ............ 858 | Vorates cotiso, sp.n. .............. 864

ANA CONDGICO, Fide Ds socscocsoconcenbenccn GOS) || Lalo nebrophone, SO Ua ancnak cad 364

Bolla sodalis, sp. n. .................. 859 | Molo? apella, sp. n................... 365

» machuca, sp. nD. ............... 859 | Carystus subrufescens, sp. n....... 365

Butleria lethea, sp.n................. 860 | Hpinosis, gen. nov. .................. 3866

53 UISISS (305 Wood0 006 seo cndananee 360 | Falga? hermione, sp.n. ............ 366

The following new species are the result of a recent visit to

Costa Rica, where nearly three years were spent in collecting.

1081 species of Rhopalocera were taken, 409 more than were

previously known from that country, and of these I am here

describing 54 speciesas new. With the Heterocera I was equally

successful, taking 4000 species, whereas only 560 are recorded in

the ‘ Biologia Centrali-Americana’ from Costa Rica, to which

number 87 were added by Pittier and Biolley in Part 11. of the

‘Invertabrados de Costa Rica.’

Subfam. DANAIN#.

NAPEOGENES HEMISTICTA, sp. n. (PI. L. fig. 1.)

g. Antenne black on basal half, apical half yellow-brown.

Head black; whitish points on frons and neck behind; a white

medial line on vertex. Collar and thorax black; a greyish

medial line on thorax and small cluster of similar scales behind.

Abdomen slate-brown above, yellow-white below. Fore wings

black ; basal third of inner margin and shorter shade above

submedian rich brown ; spots semihyaline yellowish white ; two

spots near end of cell obliquely placed ; a spot beyond cell and an

elongated spot above it on costa ; an outer row of five spots, the

one between veins 3 and 4 elongated, oblique; seven marginal

spots, the one at apex preceded by a small elongated spot on

costa. Hind wings orange-brown, the costa to discocellulars

grey ; apical portion beyond it black, its inner edge inbent to

vein 3 or before it, then outbent to near margin, which becomes

narrower, the black not reaching anal angle; five marginal

spots ; subterminal spots above and below vein 5, sometimes

almest obsolescent. Wings below similar; no brown at base of

fore wing; the costa of hind wing black.

Expanse 54 mm.

Hab. Cavillo, Guapiles.

In British Museum.

COSTA RICAN BUTTERFLIES, 341

PTERONYMIA FUMIDA, sp.n. (PI. L. fig. 3.)

3. Antenne black, apical fourth orange. Head and thorax

black-brown; a white circle around eyes. Abdomen dark slate-

brown; a whitish vertical line below terminally. Fore wings

semihyaline smoky black, the veins, costal margin, inner margin

to median, and termen narrowly brown- black : faint yellow ich

spots, one at end of cell close to median; an outangled post-

medial series of spots, the one between veins 2 and 3 elongated,

suffusing with the lowest of marginal spots, of which ee

are fave. Hind wings reddish ‘bow, semihyaline, the margins

black, more broadly so on outer margin, the inner mar gin very

finely black; traces of a yellowish snibbenaniie! shade at apex.

Wings below duller; two white spots at apex of fore wing ;

mar ginal white spots on hind wings, paired on interspaces.

Expanse 57 mm.

Hab, Carillo.

PTERONYMIA GODMANT, nom. nov.

This form is the Pteronymia simplea of the Biologia, but not

P. simplex Salv.; the discocellular line is narrow and curved ;

apex of hind wings below usually without spot, sometimes ane,

never two as in P. “simplex Salv.

Found generally in Costa Rica above 3000 ft.

ITHOMIA BOLIVARI, sp.n. (PI. L. fig. 2.)

3. Head black spotted with white. Collar dark reddish

brown. Thorax black shaded with grey. Abdomen black above,

ventrally white. Fore wings black; the cell to near end, which

is filled by the broad oblique fascia from costa, the space between

veins 2 and 3 to near termen, a large terminal spot between 2

and 3, and a subapical fascia from costa to vein 3 hyaline, the

fascia mostly shaded with white. Hind wings hyaline, the veins

and outer margin narrowly black; the costa grey-black. Fore

wings below: inner margin fuscous grey; dark spaces brown

edged with black, more broadly in cell and at apex; base of costa

black; 4 apical white spots, the one on costa minute. Hind

wings below: margins brown edged with black; a broad yellow

streak at base of costa ; veins black; small paired whitish spots

on three interspaces below apex.

Expanse 45 mm.

Generally distributed along the Caribbean coast.

In British Museum.

Subfam. SATyRINE.

EUPTYCHIA DRYMO, sp.n. (PI. L. fig. 6.)

¢. Body and wings dark brown, the disc of wings lighter

brown, especially on “fore Wings ; the two lines of randerside

faintly indicated ; a terminal black line, presen by a faintly

greyish shade, aol fuscous marginal tener a subterminal fuscous

24%

342 MR. WILLIAM SCHAUS ON

wavy line on hind wings. Fore wings below yellowish buff; the

two lines bright reddish brown, vertical, the inner line not

reaching costa; costal margin shaded with grey; a subterminal

dull brown shade interrupted by an ocellus below costa, black,

circled with yellow, and containing a bluish-white point; the

termen greyish crossed by two fine lines, faintly wavy, and a

terminal fuscous line. Hind wings below lilacine grey tinged

with pale buff; the two lines parallel, slightly inbent; a large

ocellus near apex, surmounted by a very small ocellus, and with

a yellow spot below it; a large ocellus between veins 2 and 3,

containing two bluish points with a yellow spot above it, out-

wardly broken; the ocelli edged and connected by a dull brownish

shade; termen grey, the two marginal lines more deeply wavy,

shaded with reddish brown at anal angle; a terminal fuscous line.

Expanse 37 mm.

Hab. Guapiles.

Near FZ. confusa Stgr., and #. labe Butl.

In British Museum.

EupTycHIa AGNATA, sp.n. (PI. L. fig. 4.)

3. Wings brown; traces of antemedial and medial darker

lines. Fore wings: an even marginal darker shade, faintly

darker edged; the apical spot of underside faintly indicated.

Hind wings: a large subterminal fuscous spot between veins 2

and 3; the apical spot of underside indicated; subterminal,

marginal, and terminal darker lines. Wings below pale grey ;

the antemedial and medial lines dark reddish brown. Fore

wings: the two lines vertical; a broad darker brownish-grey

outer shade; the apical ocellus black, edged with yellow and

containing a black point; dark subterminal and marginal lines ;

a fine fuscous brown terminal line. Hind wings: the two lines

slightly inbent, the outer one angled near inner margin; ocelli

black, broadly edged with yellow and containing a white point ;

darker grey shading around ocelli; a small ocellus at apex above

vein 6, and a large one below it; a small yellow spot below

vein 5; a larger yellow spot below vein 4 suffusing with the

large ocellus below vein 3; the subterminal and marginal lines

dark brownish grey, becoming reddish brown on inner margin ;

terminal line fuscous brown.

Expanse 48 mm.

Hab. Guapiles.

Near H. antonoé Cramer, found in the same locality and

E. juan Stgr.

CATARGYNNIS DRYADINA, Sp. n. (PI. L. fig. 5.)

3. Body fuscous, clothed with brown hairs; a white lateral

streak on palpi. Fore wings brown; a vertical outer row of

white spots on interspaces, broadly edged with darker brown,

the spots on costa and at submedian fold smaller, followed

throughout by a broad lighter brown shade, and preceded by a

COSTA RICAN BUTTERFLIES. 343

similar triangular shade between veins 5 and 6. Hind wings

darker, especially terminally ; an outer row of small white spots

on interspaces ; cilia partly white on interspaces. Fore wings

below dull olive-brown to beyond cell, its outer edge outcurved

from costa, uneven; outer third pale buff shaded with dark

ervey; the white spots as above broadly edged with dull dark

brown ; a broad subterminal fuscous brown line, lunular between

veins 5 and 7, obsolescent on costa; some fine dark strize on

apex ; a terminal interrupted fuscous brown shade. Hind wings

below dull olive-brown, mottled with white antemedially and

beyond medial fascia, crossed by darker strie; the medial fascia

defined by darker edging, broad, outbent in cell to end, downbent

towards inner margin; the small outer white spots broadly edged

with fuscous brown, preceded and followed by fuscous brown

shadings ; the outer margin deeply crenulate from vein 4 to anal

angle, less so towards apex.

Expanse 72 mm.

Hab. Volcano Turrialba, at 8000 feet.

Subfam. HELICONINA.

HELICONIUS GALANTHUS, ab. SUBRUFESCENS, ab. nov. (PI. LIII.

fig. 7.)

Differs from 17, galanthus Bates on the underside of hind wings;

the medial space below subcostal black, the fascia being reddish

brown, limited by a postmedial broad black line, followed by a

narrow brown shade having its outer edge incurved on interspaces.

In H. galanthus the medial space is black edged above and below

with reddish brown.

A single specimen taken on the slopes of Turrialba. Belongs

to the group with greyish costa on hind wings.

Subfam. NyMPHALiIn#&,

EUPETOIETA POASINA, sp.n. (PI. LI. fig. 2.)

2. Wings brownish buff. Fore wings: the base olive-brown

irrorated with black; an antemedial black line in cell, and one

below it, outset ; a pale spot near end of cell, broadly edged with

black ; postmedial line black, oblique from costa, inbent, lunular

below vein 4, angled on veins, followed by a straighter, narrower,

fuscous line; an outer row of black spots on interspaces, suffusing

on costa with subterminal fuscous line, which is lunular, inwardly

toothed on veins; termen fuscous, with a pale interrupted

lunular marginal line. Hind wings: base darker brown with a

black line on discocellular, limited by the black medial line, which

is incurved opposite cell, angled at vein 4, followed by a fuscous

straighter line; a black shade at base and below cell ; four outer

black spots between veins 2 and 6; subterminal line lunular

from veins 6 to 3, otherwise straighter; the pale marginal line

broader. Fore wings below light brown to postmedial; an

antemedial black line in cell; the cell spot paler edged with

344 MR, WLLLIAM SCHAUS ON

black except on subcostal; space beyond postmedial brownish

buff, the costa and termen shaded with greyish green; outer

spots below veins 2, 3, and 4; on costa a dark shade, inbent and

preceded by some white; subterminal line brown. Hind wings

below mottled brownish buff and dull green, the veins and termen

whitish grey; a curved antemedial whitish line above cell;

medial line fine, fuscous, inwardly edged with whitish grey ;

subterminal ike fine, brown, outwardly edged w ith whitish ;

cilia white with brown points at veins.

Expanse 46 min.

Hab. Poas.

In British Museum.

EReEsIA stricta, sp.n. (PI. L. fig. 7.)

¢. Antenne black tipped with yellow. Head, collar, and

thorax black ; a white streak on palpi; two white spots on frons,

and one on vertex. Abdomen brown above, laterally irrorated

with white; underneath white with a ventral black line. Fore

wings black, the spots yellow-white, partly greyish ; a round spot

at end of cell ; postmedial streaky spots irom costa to vein 4,

outcurved, and two spots between 2 and 4, inset, vertical; outer

and marginal spots, both slightly inset between veins 3 and 4,

the marginal spot obsolescent at apex. Hind wings reddish

brown ; a black shade along subcostal, expanding from middle of

wing, forming a broad margin, narrowing to anal angle; marginal

white spots at apex. Fore wings below: the spots larger, yellow-

white; some brown shading at base of inner margin. Hind

wings below similar, but the costa also black with a short yellow

streak at base; large marginal white spots.

@. Antenne yellow except at base ; a brown streak below cell

on basal half. Hind wings with the margin much more narrowly

black, and no black shading on subcostal, except subterminally ;

marginal white points from below the apical spots. Hind wings

below: the costa broadly black; a short yellow streak at base;

traces of fuscous subterminal spots; the marginal spots tri-

angular.

Expanse, 3 53 mm., 2? 60 mm.

Some males have the costa of hind wings entirely black; the

species is allied to #. pecilina Bates, but the apex is more bluntly

produced, and the termen more oblique.

In British Museum.

Eresta cara Druce. (PI. L. fig. 8.)

@. Hnt. Monthl. Mag. xi. (1874) p. 37, fig.; B. C.-A. Lep.

Rhop. vol. 1. p- 187 (tab. xxi. figs. 2, 3).

6. Fore wings black; a reddish-brown fascia through entire

cell, downbent ‘between veins 2 and 3 to near termen, with a

small black spot on it close to vein 3 postmedially, and shaded

with yellowish terminally; a small subterminal reddish-brown

spot just below vein 2; an oblique paler fascia postmedially from

COSTA RICAN BUTTERFLIES. 345

osta to vein 5, and large subterminal spots between 4 and 6; a

costa t 5, and large subt 1 spots betw 4 and 6;

small round marginal spot, tinged with whitish above vein 4,

and one below it. Hind wings black; a broad reddish-brown

fascia from inner margin to near apex; a similar marginal line,

interrupted by veins, the apical spot paler. Underneath similar

to female, but all the spots yellow imstead of white; the brown

fascia on fore wings brighter brown, separated by a black line

from the large subterminal spot. Hind wings below with the

fascia as above but tinged with yellow, otherwise agreeing in

markings with the female.

Expanse 45 mm.

Found on the Caribbean coast only.

PHYCIODES DORA, sp.n. (PI. LI. fig. 9.)

@. Palpi dark brown, whitish grey below. Head dark brown.

Body fuscous with fine yellowish irrorations. Fore wings: basal

half black, its outer edge projecting between veins 4 and 5, and

less so outcurved between vein 2 and submedian, followed by a

yellow shade from costa to vein 4, and a yellow spot between

veins 2 and 3; the base with some yellowish irrorations, leaving

a geminate antemedial outbent line clear black; two yellowish-

brown spots across end of cell, suffusing ; a reddish-brown line on

discocellular ; a medial small yellow spot above submedian ; outer

half bright brown, irrorated with black subterminally, interrupted

by round yellow spots edged with black, one below vein 2 and

another between 4 and 5; three smaller yellow spots from above

vein 6 to costa; termen fuscous preceded by a sinuous yellow-

brown line. Hind wings black irrorated with yellow-grey,

leaving clearer black lines on basal third; a brown line on

discocellular; a postmedial bright brown fascia from costa to

vein 2; large outer spots formed by yellow edging, inwardly

rounded, outwardly straight; these spots fill the entire inter-

spaces ; a subterminal wavy yellow line; a terminal yellow line

at anal angle. Fore wings below brighter, the base greyish; a

yellow-grey shade between the antemedial black lines; the cell

spots broader forming a band; the yellow shades beyond dark

space more extended ; the termen brown; a grey shade at apex.

Hind wings below: base to medial grey-brown, darkest in cell,

whitish lilacine above cell; a subbasal and an antemedial lunular

brown line, partly followed by the other less distinct lines; the

medial line dark brown, inbent on vein 4, outset between vein 2

and submedian ; postmedial space dull brown followed by a series

of spots, except on costa, the spot above vein 4 round, fuscous

brown, the spot below 4 triangular, the others more lunular,

mostly edged with geminate lilacine lines; a subterminal geminate

lunular dark brown line; termen lilacine white from apex to

below vein 6, then dark brown, shaded with still darker brown.

between spots and lunular line,

Expanse 39 mm.

Hab. Poas.

346 MR. WILLIAM SCHAUS ON

PuycropEs pHLEGIAS G.&S. (PI. L. fig. 9.)

-g. B.C.-A. Lep. Rhop. vol. ii. p. 680 (tab. eviii. figs. 21, 22).

@. Body and wings above fuscous brown, the body and base of

wings irrorated with ochreous. Fore wings: spots white; an

oblique line across cell near end, followed by a large white spot

between veins 2 and 3, and a few white scales above 3 and below

2; a postmedial oblique broad white line from costa to vein 4; a

small outer spot on costa; a quadrate spot between veins 4 and 5,

and a larger spot above submedian, extending just above vein 2 ;

traces of small whitish spots between veins 5 and 7; traces of a

fine outcurved subterminal line, broad and well marked from

vein 4 to near vein 2. Hind wings: a broad postmedial white

fascia from vein 7 to inner margin, followed below vein 4 by a

line of ochreous irrorations; subterminal line slightly lunular,

fine, white on costa, then ochreous. Fore wings below: base and

costa mottled pale brown and grey; a broad antemedial fascia

defined by edging of dark outcurved lines; the white line at end

of cell downbent to inner margin; the space between postmedial

and outer spots fuscous; apex lilacine grey; a broad terminal

dark brown shade between veins 4 and 6; termen more narrowly

brown at tornus; subterminal line fuscous brown. Hind wings

below lilacine grey, broadly shaded with white postmedially

between veins 2 and 7; basal half crossed by fine brown lines ;

the postmedial better defined, incurved on costa, and followed

there by a brownish shade; a fine lunular outer line, with a series

of faint greyish spots beyond it; subterminal line geminate,

lunular, filled in and preceded by a dark grey shade from vein 7

to vein 2; termen dark brown from above vein 5 to below vein 2.

Expanse 39 mm.

Hab. Peralta.

PHYCIODES NIVEONOTIS Btl. & Druce. (PI. LIII. fig. 10.)

6. Cist. Ent. i. (1872) p. 100, fig.; B.C.-A. Lep. Rhop.

vol. i. p. 200 (tab. xxi. fig. 27).

@. Body above fuscous. Wings fuscous brown, Fore wings:

a large white spot just beyond middle from submedian to near

vein 2, edged with yellow-brown, irrorated with black ; the outer

broad white line from costa to vein 5 similar; small subterminal

similar spots between wings 4 and 6, and a yellow-brown spot

below vein 4. Hind wings: a yellow-brown line on discocellular ;

a line of similar irrorations beyond postmedial fascia which has

its edge also finely yellow-brown; subterminal line brownish,

faintly indicated towards inner margin. ore wings below light

brown; an antemedial fascia defined by dark lines; two whitish

spots near end of cell; a brown spot on discocellular edged with

black; a broad black postmedial shade from costa, outecurved to

near tornus; a subterminal lunular dentate darker brown line ;

the spots all whitish. Hind wings below: base grey-brown

crossed by brown lines; the whitish fascia obscured by brown

COSTA RICAN BUTTERFLIES. 347

irrorations and crossed by a fine line; outer margin broadly

brownish, irrorated partly with grey; an outer row of fuscous

grey spots on interspaces ; a marginal lunular dark brown line,

Expanse 30 mm.

Hab, Juan Vinas.

EUBAGIS HECUBA, sp. n. (PI. LI. figs. 4, 5.)

¢. Body above fuscous, whitish underneath. Wings bluish

green. Fore wings: apex and outer margin black, expanding at

vein 4, its inner edge obliquely outbent to vein 2, below which it

is very finely black. Hind wings: a marginal and a terminal

black line, suffusing at apex. Wings below white. Fore wings:

costa grey-brown with a white streak at base ; subcostal medially

red-brown ; a fine blue line and red-brown streak below subcostal

to beyond cell; a red-brown streak on discocellular, edged with

metallic blue and then black; a broad medial black fascia from

end of cell, and an antemedial black fascia inbent across cell,

interrupted by median and narrowest below it; a broad black line

above vein 3 from cell, to a similar oblique line which extends

from costa to brown terminal space; another black line from

vein 3 is obliquely downbent to a black tornal space ; the tornus

itself whitish; the termen from vein 2 to just above vein 6

reddish brown, its inner edge fuscous, upbent to costa, leaving a

white apical space. Hind wings: lines reddish brown partly

edged with fuseous; an inbent basal line; a medial fascia

expanding to close to inner margin, and then downbent, enclosing

a white space from subcostal; a broad subterminal line, its outer

edging downcurved near apex, forming a marginal fuscous line

partly mottled with reddish brown; anal angle reddish brown

where all the lines suffuse ; some metallic blue scaling near anal

angle.

@ similar below. Fore wings above green; a white spot in

cell above vein 2; a black shade below subcostal expanding at end

of cell, followed by a large postmedial white spot oblique from

costa to just below vein 4; a large postmedial vertical spot between

submedian and vein 3; outer margin broadly black ; a subterminal

small white spot between veins 2 and 3, and smaller spots above

and below vein 5, also a small inset spot near costa. Hind wings

green; a postmedial broad white space from costa to below

vein 5; a broad black subterminal line, and a similar marginal

line; the termen brown-black with a fine green line near anal

angle, the lines all suffusing in a brown shade at apex.

Expanse, ¢ and 2 40 mm.

fab. Carillo, Guapiles.

Allied to #. vicaria Bates.

In British Museum.

PERISAMA BARNESI, sp. n. (Pl. LI. fig. 10.)

@. Palpi black broadly streaked with white. Body black

above, some white scaling around the eyes. Wings black. Fore

348 MR. WILLIAM SCHAUS ON

wings: some dark blue irrorations at base, along veins, and in

cell; a postmedial curved greenish-blue fascia from costa to tornus,

its edges uneven. Hind wings: some scattered blue irrorations ;

the outer margin greenish blue; a terminal brown-black line ;

cilia white. Fore wings below black; the cell to near end and a

space below it crimson; a postmedial curved series of metallic

blue spots ; termen white, broadest at apex, crossed by a fine

marginal black line. Hind wings below grey-brown shading to

whitish on termen; costa narrowly crimson; outer small round

black spots on interspaces; a subterminal and a terminal dark

brown line.

Expanse 43 mm.

Hab. Carillo.

This species is named after J. Barnes, who captured the only

specimen met with.

This is the first Perisama recorded from Central America.

‘VICTORINA STENELES, ab. STYGIANA, ab. noy. (Pl. LI. fig. 6.)

d. Fore wings fuscous brown, darkest medially ; a large pale

green spot on inner margin close to middle, almost reaching

vein 2; a very small outset spot above vein 2; a small white spot

medially on costa ; subterminal whitish spots, larger and better

defined near tornus. Hind wings fuscous brown; base darker

followed by a broad pale green fascia and a medial fuscous fascia,

all stopping at submedian; the inner margin white; an outer

row of oval green spots on interspaces preceded by fuscous

shadings ; veins terminally darker edged. Underneath more like

the typical form.

Expanse 82 mm.

Hab. Avangarez.

Fam. ERYCINIDA.

Eurycona Mystica, sp.n. (PI. LI. fig. 8.)

@. Palpi white. Frons white divided by a brownish line.

Vertex black. Collar and thorax brown. Abdomen above

brighter brown, underneath white. Fore wings fuscous brown :

the cell and basal half of wing below it bright brown; a paler

rust-brown shade between veins 2 and 4, not reaching outer

margin. Hind wings rust-brown, the costal margin broadly

fuscous brown. Wings below pale ih ownish grey, ‘Tne postmedial

line vermilion finely “edged with dark grey. Fore wings: the

postmedial line slightly outbent, straight ; a subterminal narrow

vermilion shade ; a terminal yellow-brown shade. Hind wings:

the postmedial line vertical to below vein 2, then curved and

wayily inbent to inner margin; a subterminal lunular dentate

brownish shade from costa to below vein 3, followed by marginal

points between veins 4 and 7, between 3 and 4 bya large black

spot outwardly edged with white; below vein 2, irregular brown

COSTA RICAN BUTTERFLIES, 349

lines terminating in black points followed by white shading ;

termen from above vein 2 to anal angle vermilion,

Expanse 22 mm.

Hab. Rio Grande, San Mateo.

KuRyYGONA LEUCON, sp.n. (PI. LI. figs. 1, 3.)

3. Palpi, lines on frons and tarsi pale buff. Head fuscous.

Thorax and base of abdomen dark brown, abdomen otherwise

bright brownish red above ; underneath greyish buff. Fore wings

faintly angled at vein 4, fuscous ; a brown-red shade at base from

cell to inner margin; a similar streak at base below subcostal.

Hind wings brownish red; the costal and outer margins to

vein 2 black, the inner margin fuscous brown. Wings below pale

greyish brown, the medial line brownish red finely edged with

black. Fore wings: medial line slightly angled at vein 4,

terminating at submedian fold; an outer vertical dark brown

line, not so well marked. Hind wings: the medial line incurved

between veins 4 and 2, deeply downcurved below 2; subterminal

greyish lunules from costa to vein 4, edged with dark brown;

between 4 and 3 the lunule is larger, yellowish, followed by a

black spot, outwardly edged with white ; the inner brown shading

expands between 3 and 2, and is followed by a dark wavy line ;

from vein 2 to inner margin are three yellow-brown streaks

outwardly irrorated with black and terminating in white points

before a fine marginal fuscous line; cilia reddish brown except at

apex.

@. Fore wings fuscous; a large orange space from middle of

cell and well beyond it, not reaching costa, or below vein 2. Hind

wings dark brown ; the outer margin outcurved from veins 4 to 2,

shaded terminally from 4 to anal angle with brownish red.

Wings below hght yellow-brown, the medial line as in male, but

brighter red ; the outer line on fore wings also red; the lunules

on hind wings heavier, geminate, the marginal shadings faintly

marked,

Expanse, ¢ 26 mm., 2 29 mm.

Hab. Turrialba.

KURYGONA LEUCOPHRYNA, sp.n. (PI. LIII. fig. 8.)

3d. Frons white. Vertex and body fuscous; some brown

shading on abdomen dorsally. Wings brown. Fore wings: costal

margin, front of cell, apex, and outer margin more narrowly

tinged with fuscous. Hind wings: costal and inner margins

greyish ; a subcostal fuscous shade. Wings below silvery white.

Expanse 27 mm.

Hab. Cachi.

Near L. cataleuca, but larger aud brighter brown.

EuRYGONA MATUTA, sp. n. (PI. LI. fig. 7.)

g. Frons white. Vertex and body above fuscous brown,

underneath white. Fore wings fuscous, the base in and below cell

350 MR. WILLIAM SCHAUS ON

shaded with brown; an orange wedge-shaped spot close to cell,

between veins 2 and 4. Hind wings brown, the outer margin

shaded with fuscous ; the costa whitish ; the inner margin grey-

brown; cilia on inner margin and apex white. Wings below

white; a very fine pale olive-brown postmedial line, almost

vertical, bluntly curved below vein 2 on hind wings. Hind wings:

a small black marginal spot between veins 3 and 4; three black

points from vein 2 to inner margin, and a few subterminal black

irrorations.

Expanse 21 mm.

Hab. Juan Vinas.

MESOSEMIA ALBIPUNCTA, sp. n. (PI. LIII. fig. 12.)

¢. Thorax and base of abdomen metallic blue, abdomen other-

wise and head fuscous; underneath grey. Wings vivid blue.

Fore wings: a white point near end of cell ; outer margin broadly

black, widest on costa, its inner edge outbent and curved near

tornus. Hind wings: termen black, the veins close to it streaked

with black; a subterminal black spot at apex, suffusing with

black costal margin ; in another specimen the spot continues as a

line to inner margin. Wings below grey-brown, the lines darker

brown. Fore wings: the two lines before ocellus parallel, outbent

from subcostal ; ocellus large, a white central point, and two outer

points, circled with yellow; two lines beyond ocellus, close

together, diverging slightly towards inner margin ; postmedial

space grey irrorated with metallic blue from near costa to vein 3,

limited by a fine greyish yellow line, outbent from costa, at sub-

median approximated by asimilar subterminal line. Hind wings:

the antemedial and postmedial lines straight, geminate; the

medial shade partly linear interrupted by a smaller ocellus, black,

circled with yellow and containing two white and blue points; a

broad outer shade; a subterminal line; termen dark shaded.

Expanse 27-30 mm.

Hab. Guapiles, Esperanza.

HERMATHENA OWENI, sp.n. (PI. LIII. fig. 9.)

g. Head and thorax black clothed with greyish-white hairs.

Abdomen grey, with transverse black segmental shades. Wings

white, the base mottled with black with traces of a subbasal

whitish line; spots black, two beyond cell; a medial spot below

vein 2; subterminal quadrate spots above and below vein 6, also

above and below vein 3; apex black; spots at end of veins ;

terminal interspaces between veins 2 and 3, and 4 and 6 mottled

with black. Hind wings: terminal spots at veins; subterminal

quadrate spots above and below vein 7. Underneath similar ;

the basal mottling showing through from above; the terminal

markings on interspaces entirely black.

Expanse 42 mm.

Hab. Costa Rica.

Named after Prof. Owen of Madison, Wis., who has kindly

given me this unique specimen.

COSTA RICAN BUTTERFLIES. 301

CoRRACHTA, gen. nov.

2. Fore wings: outer margin straight at apex, then rounded ;

vein 3 close to lower angle of cell; 4 from lower angle; disco-

cellular incurved, slightly oblique; 5 from just above middle ;

6-10 stalked from upper angle, 10 before 6; 11 from close to end

of cell anastomosing with costal vein. Hind wings as long as fore

wings; cell elliptical; vein 3 before end; 4 from end; 5 from

middle of discocellular ; 6 and 7 stalked. Abdomen stout.

Type of genus, C. lewcoplaga Schaus.

CoRRACHIA LEUCOPLAGA, sp.n. (PI. LITT. fig. 2.)

@. Body and wings brownish slate-colour. Fore wings: a

broad medial white fascia, slightly outcurved, from subcostal to

submedian vein, a whitish streak on costa above it between veins

10 and 11. Hind wings: marginal fuscous lunules, faintly edged

with greyish. Wings below grey. Fore wings: the white fascia

followed by a broad outcurved fuscous shade to some whitish sub-

terminal points. Hind wings: termen irrorated with whitish ;

small subterminal fuscous spots preceded by whitish shadings.

Expanse 35 mm.

Hab, Palmital.

This peculiar species will probably be found to have a very

dissimilar male.

CHARIS SUBTESSELLATA, sp.n. (PI. LITT. fig. 11.)

Body above fuscous. Fore wings fuscous, the base tinged with

olive-brown ; a white spot in cell; an outer white spot between

veins 2 and 3, and a smaller one between 5 and 6; a small post-

medial spot above vein 6. Hind wings dark olive-brown; cilia

white between veins 5 and 6; a yellow-grey marginal streak at

anal angle. Fore wings below reddish brown, markings bluish

white partly edged with black ; a streak at base of costa; a basal

streak below subcostal ; an antemedial fascia in cell, and one

below it outset ; a medial fascia in cell, with a white streak above

subcostal, and a buff streak above costal vein; an outset medial

fascia below cell, all of them narrower towards median, the two

below cell suffusing on submedian; a fine black line on disco-

cellular followed by small spots from costa to vein 4; two costal

spots beyond, superposed, and small black spots on interspaces

below; an interrupted subterminal white line preceded by a white

point above vein 6,a large one below it, black spots below 5

and 4, and a still larger white spot between 3 and 2; a marginal

silvery blue line; a fine terminal black line; cilia fuscous.

Hind wings below reddish brown except on outer third ; a white

triangular spot on costa near base, and a quadrate medial spot ;

an antemedial white spot in cell, and transverse line across end

of cell; two white spots below cell suffusing with a white streak

below vein 1; a black line on discocellular, followed by a white

fascia from veins 7 to 4, and from vein 2 to inner margin; black

points in some of the reddish-brown spaces, which are also partly

352 MR. WILLIAM SCHAUS ON

black-edged ; a postmedial reddish brown shade from costa to

vein 4, followed by a macular black line, and then by a broad

grey-brown shade crossed by a fine black line; postmedial space

below vein 4 greyish brown to vein 1, then reddish brown on

inner margin, followed by greyish brown shading from vein 4 to

vein 1, and buff shading on inner margin, these shadings out-

wardly edged with black and followed by a silvery opalescent

shade; outer margin dull dark brown outwardly edged with

opalescent scaling, and containing small fuscous spots below

vein 4 and vein 3; the tails and a terminal buff line from vein 4

to anal angle edged on either side with black; from vein 4 to

apex the terminal line is reddish brown, preceded at apex by a

short white line.

Expanse 24 mim.

Hab. Cavrillo.

Near C. chrysus Cr.

CHARIS TURRIALBENSIS, sp. n. (Pl. LIII. fig. 13.)

3g. Body above fuscous. Wings dark brown, with antemedial

and postmedial fuscous shadings on interspaces. Fore wings:

cilia at tornus orange-red. Hind wings: a black line on disco-

cellular ; a subterminal greyish buff shade from vein 6 to anal

angle, crossed by a black macular line, preceded below vein 3 by

a large brown spot, some reddish brown shading on inner margin

at angle. Fore wings below orange-red ; a pale buff triangular

spot on costa near middle with its apex on median at vein 3,

edged with silvery blue; similar metallic scaling on costa near

base, and two inbent lines across cell; below cell and vein 2,

medial, postmedial, and subterminal pale greyish shades, the

intermediate red portions edged and suffused with black ; a black

line on discocellular, followed by metallic spots; below vein 5a

macular black line; outer pale greyish spots, followed above 5

and 6 by silvery streaks, preceded by silvery spots and some black

edging, below vein 5 the greyish spots are inwardly edged with

black, outwardly with metallic scaling and black edging; a

marginal metallic line interrupted by veins. Hind wings below

orange-red, divided by metallic antemedial, medial, and _post-

medial irregular lines, with some short black lines and spots in

reddish portions; outer margin broadly pale greyish irrorated

with brown; a marginal metallic line finely edged outwardly

with black, and preceded by some black below veins 4 and 3; a

terminal black line, also edging tails.

Expanse 21 mm.

Hab. Tarrialba, 6000 feet.

Fam. Lyc £#NIDS&.

THECLA HypocritA, sp.n. (PI. LIT. fig. 6.)

$. Body dark brown mottled with bluish grey hairs and

irrorations. Fore wings fuscous brown, except a pale blue space

COSTA RICAN BUTTERFLIES, 353

occupying basal half of cell, space below to inner margin, and

just above vein 2 to near subterminal space; a pale brown patch

of coarse scales at end and beyond cell. Hind wings pale blue;

outer margin fuscous brown; inner margin whitish, becoming

fuscous towards anal angle; cilia brown tipped with grey.

Fore wings below fuscous; costal margin broadly whitish, post-

medially to vein 4; some brown mottling at base of costa; a

brown medial fascia to median and end of cell enclosing a white

line on discocellular ; an outer and a subterminal oblique broad

brown line to vein 5; outer margin narrowly white crossed by a

fine marginal brown line. Hind wings below yellowish white,

crossed by broad brown lines; a fuscous basal space, outangled

on median ; the inner margin broadly irrorated with fuscous and

pale brown; medial and postmedial lines vertical, angled at

vein 2 and inbent to inner margin; a subterminal brown shade

suffusing with postmedial from veins 5 to 2, then continuing to

anal angle; a broken marginal brown shade; anal angle slightly

lobed, no tail.

Expanse 19 mm.

Hab. Tuis, Juan Vinas.

A specimen from Cordoba, Mexico, has the lines below partly

greyish, but undoubtedly belongs to this species.

Near 7’. dolylas Cr.

In British Museum.

‘THECLA SUBFLORENS, sp. n. (PI. LIT. fig. 9.)

gd. Antenne black ringed with white. Body above fuscous,

underneath white; some green scales on frons. Wings dark

purple-slate colour. Fore wings: the costa, apex broadly,

and outer margin fuscous brown. Hind wings: the margins

narrowly fuscous brown. Wings below rich green. Fore wings

paler tinged, except on costa and apex ; the inner margin dark

grey. Hind wings: a faint paler line on discocellular; a similar

irregular outer line, black, outwardly edged with white from

vein 2 to inner margin; a few red scales at anal angle, sur-

mounted with black and white; a terminal black line at anal

angle.

Expanse 20 mm.

Hab. Poas.

THECLA LEOS, sp.n. (Pl. LII. fig. 7.)

3. Palpi black and white. Head, collar, and patagia black ;

white lines around eyes. Thorax and abdomen above metallic

blue, underneath whitish. Wings brilliant dark blue. Fore

wings: apex and outer margin black, narrowest at tornus; a

large black patch filling the end of cell from before middle

followed beyond cell by a small slate-coloured spot. Hind wings :

the termen narrowly black; the inner margin grey. Wings

below clear bright brown, Fore wings: outer line fuscous brown,

outwardly edged with white, faintly outbent, slightly Iunular ;

354. MR. WILLIAM SCHAUS ON

inner margin greyish; a faint dark marginal line at tornus.

Hind wings: outer line fine, fuscous, outwardly edged with

white, more broadly on costa, vertical from veins 8 to 6, appearing

incurved on account of the broader white edging, slightly outset

below 6 and below 5, inset below 4, outset below 3, inset and

horizontal below 2, downbent on submedian, angled and upbent

to inner margin; a fine subterminal black line from vein 3 to

inner margin, and darker spots at apex; a marginal red spot

with black point and terminal white line between 2 and 3; a

black spot with white irrorations below vein 2; a black spot at

anal angle surmounted by a reddish shade.

Expanse 24 mm.

Hab. Guapiles, La Florida.

I do not know any species with which I can compare this.

In British Museum.

THECLA MELMA, sp.n. (PI. LIT. fig. 4.)

6. Palpi black, the base of joints white. Body black; some

white on vertex, and white lines close to eyes. Wings deep blue.

Fore wings: costa, apex, and outer margin black, the apical

space reaching to near cell; a black line on discocellular. Hind

wings: margins very narrowly black, slightly broader at apex.

Wings below bright greyish bistre. Fore wings: an outer

vertical fine fuscous line, outwardly edged with white from near

costa to vein 2; a terminal fine brown line; cilia fuscous.

Hind wings: a white line on discocellular; outer line black out-

wardly edged with white, outset on costa, almost vertical from

veins 7 to 4, then inset and downbent to vein 2, forming the

usual W to inner margin; subterminal lunules indicated by

ereyish shading from vein 7 to vein 2, followed between 2 and 3

by a large red space, outwardly broken by a triangular black

space ; some reddish scales above vein 3; below vein 2, marginal

white irrorations ; a terminal white line from vein 5 to sub-

median ; anal lobe black inwardly edged with white.

Expanse 27 mm.

Hab. Guapiles; also Bugaba, Panama.

THECLA GUAPILA, sp. n. (PI. LII. fig. 1.)

3d. Thorax and base of abdomen metallic blue, abdomen

otherwise fuscous above, yellowish white below. Head brown,

with white lines at eyes. Wingssilky blue. Fore wings: costal

margin, apex to near glandular spot, and outer margin, narrowly

at tornus, black; a small black spot at end of cell followed by

an oval brownish glandular spot. Hind wings: apex narrowly,

and cilia black. Wings below dark brownish slate-colour. Fore

wings: inner margin ‘oreyish ; a fine outer fuscous line, almost

vertical from below costa to vein 2, outwardly faintly pale- -edged ;

faint traces of a dark subterminal line. Hind wings: outer “line

interrupted, dark brown, outwardly edged with white, inbent on

COSTA: RIGAN BUTTERFLIES. 355

costa, vertical from veins 7 to 2, but inset between 3 and 4, angled

below vein 2, and inbent to inner margin; subterminal darker

spots, edged with whitish irrorations, except at apex, inset be-

tween veins 2 and 3, followed by a red marginal spot, containing

a small black spot ‘outwardly ; ; below vein 2 only a marginal

brown spot irrorated with white; below submedian a line in-

wardly edged with white and parallel with outer line; a terminal

white line between veins 2 and 3; anal lobe black.

Expanse 28 mm.

Hab. Guapiles.

In British Museum.

THECLA AMPHRADE, sp.n. (PI. LIT. fig. 10.)

@. Wings fuscous brown, the hind wings, and inner area of

fore wings below cell glossed with slate-blue; cilia light brown.

Hind wings: the terminal black line inwardly edged with white

near anal angle; a small red-brown spot at anal angle; tails

black tipped with white. Wings below light brown, the post-

medial line dark brown outwardly edged with black and then

white, very indistinctly on fore wing. Fore wings: the line

vertical from costa to vein 2; Beha indistinct paler subterminal

shade, crossed by a faint dark line; a terminal fine dark line.

Hind wings: the line vertical to vein 4, constricted and slightly

oblique to vein 2, upcurved and outangled on submedian; a sub-

terminal paler shade as on fore wing, the faint line crossing it

more distinct followed on inner margin, between veins 2 and 3 and

just above 3, by reddish shades; a black marginal spot between

veins 2 and 3 and at anal angle; a black spot irrorated with

white below vein 2; the terminal dark line inwardly edged with

white.

Expanse 23-25 mm.

Hab. Guapiles.

Near 7’. politus H. Druce.

In British Museum.

Subfam. PrerInz.

ARCHONIAS INTERMEDIA, sp. n. (PI. LIT. fig. 8.)

3. Palpi yellow, somewhat fuscous at base. Head and thorax

black ; three whitish points on vertex. Abdomen brown ; sub-

lateral yellow line. Fore wings black, markings yellow-white ;

an oblique spot at end of cell, medially constricted ; two diffuse

spots beyond cell; elongated postmedial spots above and below

vein 3, and geminate streaks below 2, all followed by marginal

spots, ‘small at tornus, elongated towards costa ; a yellow and

reddish streak along inner margin, not reaching ‘base or tornus.

Hind wings bright brownish red, ‘the costal and outer margins

broadly black, the inner margin finely so; the veins black; cell

edged with black above and below; marginal small white spots.

Underneath duller, the hind wings browner, with a brown up-

Proc. Zoo. Soc.—1913, No. XXY. 25

356 MR. WILLIAM SCHAUS ON

curved streak on basal half of costa; the subcostal and median

broadly shaded with black ; the marginal white spots larger.

2. The markings on fore wings reduced, except the marginal

spots; the hind wings reddish brawn, the costa with only a black

shade medially ; the marginal spots lve ger, yellow, irrorated with

reddish. Underneath similar to the male,

Expanse, ¢ 60 mm., 2 65 mm.

Hab. Carillo.

The fore wings are like A. dismorphites Btl., the hind wings like

A. nigrescens G. & 8. All these forms will no doubt be found to

gradually intergrade. The males are difficult to capture, flying

very high in a circle, and rarely settling.

In British Museum.

PIERIS LIMONA, sp. n. (PI. LIT. fig. 5.)

3g. Antenne black tipped with white. Head and thorax

fuscous, the latter clothed behind with white hairs. Palpi white,

the third joint streaked with black. Abdomen fuscous grey

above, whitish below. Wings white, the veins broadly edged

with chalky white. Fore wings: a fuscous grey streak on basal

third of costa ; a small black spot on discocellular ; apex narrowly

black; marginal fuscous shades, not reaching tornus. Hind

wings: cilia black. Fore wings below: a large black spot on

discocellular ; apex greyish. Hind wings below yellowish white,

the veins dark.

©. Fore wings above with a large black spot on discocellular,

and a fuscous streak in cell at subcostal reaching it; the apex and

termen more broadly black, its inner edge somewhat lunular ;

underneath the black margin replaced by a pale lilacine orey

shade. Hind wings below “Vilineine white, the veins brown ; a

postmedial br oneal shade just beyond cell across veins 3 and 4.

Expanse, ¢ 75 mm., 9 78 mm.

Hab, Limon.

Near P. diana Feld.

In British Museum.

ACMEPTERON POASINA, sp.n. (PI. LIT. fig. 3.)

g. Antenne black. Palpigrey. Head and collar dark brown.

Thorax and abdomen fuscous irrorated with white. Fore wings

black ; median, fold, submedian, and vein 2 all basally streaked

with white ; spots llacine colhthe ¢ a medial streak on costa, spot

beyond cell, an outer spot above vein 3, and one on costa, also. a

small subterminal spot above 6; sometimes traces of spots on

either side of upper discocellular ; cilia greyish brown. Hind

wings iridescent silky brown tah a large ‘medial dull brown

spot; from cell behind to inner margin and termen at vein 5

black, crossed by a grey fascia; cilia yellow. Wings below as in

A. cinerascens Salv., but the markings all more intense, the anal

angle and inner margin bright yellow crossed by purple-brown

strie,

COSTA RICAN BUTTERFLIES. 357

2. Wings black, the markings white. Fore wings: an oblique

fascia from middle of costa consisting of coalescing spots; three

oblique spots from costa before apex; the outer half of inner

margin with a white streak. Hind wings: a large white space

occupying the greater portion of the wing, outangled on vein 5,

its hind edge straight; the base of cell and a streak below it

black. Fore wings below: a white space on basal half from

within cell to inner margin; costa basally lilacine; the medial

fascia as above but with a yellow streak on costa; apex lilacine

white, shaded with yellow on costa; termen finely yellow, pre-

ceded by light brown and greyish white shading. Hind wings

below white: the base, outer half from inner margin to vein 4,

and terminally between 4 and 5 lilacine white irrorated with

brown; the shade in cell extending somewhat, and upbent to

costa. ; subterminal brown irrorations from costa to vein 6; inner

margin narrowly bright yellow, termen finely similar pr eceded by

& narrow inter rupted brown shade.

Expanse, ¢ 60 mm., 2 67 mm,

The male can be distinguished from 4. cinerascens Salv. by the

narrow grey fascia on hind wings above. The female differs in

the broader black margin, the lilacine shading, and brown irrora-

tions on hind wings. aN cinerascens female has the hind wings

below bluish white with faint traces of darker shadings. If the

two forms ogcurred in different localities I should not separate

them specifically, but they fly together and no intergrades were

found.

In British Museum.

ENANTIA LUA COSTARICENSIS, subsp. n. (PI. LIT. fig. 2.)

3. Like #. lua Hew., but the inner edge of black margin on

hind wings near apex is straight, not dentate ; underneath the

spots are ‘lar ger and more confluent. The wings are also broader.

Expanse 62 mm.

Hab. Cachi.

Fam. HESPERIIDA.

TELEMIADES MNEMON, sp. n. (PI. LIV. fig. 1.)

3. Palpi white below, black above with yellow scales at tip of

second joint. Head and collar black mottled with yellow. Thorax

and abdomen brown. Fore wings dark brown, the base to near

middle thickly irrorated with yellow-brown hairy scales ; similar

sealing subterminally, expanding at tornus, also a few similar

scales” beyond cell; medial space below cell tinged with blue-

black ; spots semihyaline yellowish ; a large spot in end of cell,

its outer edee incurved ; a narrow vertical postmedial spot be-

tween veins 2 and 3, and a small spot above 3, slightly outset ;

three small outer spots below costa, and two subterminal between

veins 4 and 6. Hind wings with only the costal margin, a post-

medial macular shade, and terminal spots fuscous brown, other-

wise overlaid with yellow-brown hairy scales. Fore wings below

25*

358 MR. WILLIAM SCHAUS ON

dull fuscous brown; the inner margin tinged with grey ; some

yellow scaling at base, below costa, and in cell. Hind wings

below dull brownish yellow; the outer margin broadly dark

brown ; some brownish-yellow scaling subterminally from vein 3

to inner margin.

Expanse 28 mm.

Hab. Zent district.

NASCUS PHINTIAS, sp. n.

3. Palpi white tipped with black above. Head and collar

fuscous mottled with olive-brown ; white points laterally on head.

Thorax and abdomen shaded with orange-brown. Fore wings

fuscous brown, the base and inner margin shaded with orange-

brown ; spots semihyaline whitish; a spot at end of cell, out-

wardly produced in front; a large postmedial spot between

veins 3 and 4, and a smaller one below 2; an outset large spot

between veins 3 and 4, and a point between 4 and 5; a curved

outer series of spots between veins 6 and 10. Hind wings

orange-brown, the costal margin broadly fuscous, the termen

narrowly so; a fuscous line on discocellular; a broader post-

medial line not reaching inner space. Wings below brown-

black. Fore wings: the base shaded with yellow-white; short

whitish streaks above cell-spot. Hind wings: the basal half and

inner margin for two-thirds pale yellow ; a thick black line on

discocellular.

Expanse 55 mm.

Hab. Guapiles.

The female of this species is figured in the Appendix to the

‘Biologia’ as a variety of WV. phocus Cr., but the discovery of the

male proves it to be a distinct species.

CYCLOS#MIA SUBCHRULEA, sp.n. (Pl. LIV. fig. 14.)

Body fuscous brown. Wings dark greyish brown, the markings

fuscous brown ; a postmedial and marginal broad line, somewhat

macular, also a similar medial line on hind wings. Fore wings:

the costa and termen broadly shot with green, the veins more

vividly so towards apex, but only noticeable in a certain light; a

large black spot in cell at end, faintly edged with paler brown,

and containing two bluish-white points. Fore wings below dark

brown shot with green; the inner margin shaded with grey; a

fuscous shade in end of cell; a postmedial fuscous shade. Hind

wings below bright blue; the costa, apex narrowly, and cilia

dark brown.

Expanse 31 mm.

Hab. Sixola, Guapiles.

In British Museum.

ECHELATUS LUCINA, sp.n. (PI. LIV. fig. 8.)

Body fuscous brown, the vertex and collar glossed with green.

Wings fuscous brown. Fore wings glossed with purple ; an ante-

COSTA RICAN BUTTERFLIES. 359

medial and a postmedial fuscous shade, the latter slightly out-

curved below costa, vertical below vein 4. Hind wings with

fainter purple gloss, except a streak below cell from base to

termen ; a fuscous shade at end of cell, and a similar postmedial

shade. Wings below paler brown, the lines as above, dark brown ;

the antemedial on fore wings replaced by a dark brown shade

from near base to middle, except on inner margin, which is whitish

grey at base.

Expanse 37 mm.

Hab. Guapiles.

AYE CANACE, sp. n. (PI. LIV. fig. 17.)

3. Body and wings fuscous brown. Fore wings: a semi-

hyaline point medially between vein 11 and costal vein ; a similar

outer point between veins 8 and 9, and one below 8 slightly inset.

Hind wings: outer margin broadly blue from vein 6 to inner

margin ; cilia fuscous brown. Fore wings below duller. Hind

wings below blue ; inner margin whitish blue ; costal margin and

apex to vein 6 dull fuscous brown; a similar downbent streak

from vein 2 to anal angle.

Expanse 23 mm.

Hab. Carillo.

Boa SODALIS, sp.n. (Pl. LIV. fig. 12.)

2. Palpi brown mottled with buff and white. Body fuscous

brown. Wings brown slightly irrorated with lighter brown and

buff. Fore wings: a fuscous shade on discocellular followed by a

faint paler brown shade; three outer white hyaline spots below

costa, straight and outbent; three similar postmedial vertical fine

streaks between submedian fold and vein 4; a faint subterminal

paler brown shade cut by darker veins. Hind wings: a faint

subterminal paler brown shade narrower than on fore wings.

Wings below duller brown ; the subterminal shade on fore wings

narrow; hind wings irrorated with buff-brown; a small pale

brown spot in cell, a streak on discocellular, some small post-

medial spots near inner margin, and the subterminal spots all

very indistinct.

Expanse 29 mm.

Hab. El Alto.

Nearest B. cylindus G. & $.; smaller, the spots also smaller ;

the outer spots in a straight line and oblique.

Boia MAcHUCA, sp. n. (PI. LIV. fig. 4.)

3. Body above fuscous brown; some greyish white hairs in

front of antenna; some yellowish white hairs on abdomen.

Throat white; fore femora grey. Abdomen below greyish -

brown. Fore wings fuscous brown; outer margin faintly paler,

forming a subterminal line with darker marginal spots on inter-

spaces ; three white points outwardly between veins 6 and 9.

360 MR. WILLIAM SCHAUS ON

Hind wings dark brown ; large postmedial and marginal fuscous

brown shades, all indistinct and confused. Fore wings below

dark grevish brown ; a broad darker postmedial shade; the costa

tinged with iridescent green; the white points as above. Hind

wings below dark brown; some white irrorations on basal half,

and more thickly aiong inner margin; traces of a darker post-

iwnedial shade, and small marginal spots.

Expanse 26 mm.

Hab. San Mateo.

In British Museum.

BUuTLERIA LETHHA, sp.n. (PI. LIV. fig. 15.)

3. Palpi black mottled with yellowish hairs. Body above

black ; some orange hairs on abdomen; abdomen below yellow.

Fore wings fuscous brown ; a large orange-yellow spot just beyond

middle between veins 2 and 4, extending into end of cell, and

slightly below vein 2; a small oblique similar spot outwardly

below costa; cilia dark brown. Hind wings black; cilia and a

large spot occupying middle of wing orange-yellow. Fore wings

below deep yellow: cell to near end black, extending at base

below cell; a broad black fascia beyond cell, not reaching costa,

outbent above vein 4, and downturned to inner margin at tornus.

Hind wings below yellow; some black scaling at base of inner

margin.

Expanse 20 mm.

Hab. Poas.

Near B. lalage Godm.

In British Museum.

BUTLERIA Lysis, sp.n. (PI. LIV. fig. 2.)

3. Body above black: abdomen below yellow. Wings dark

brown, spots yellow. Fore wings: a spot between veins 2 and 3,

just beyond middle, surmounted by a triangular spot in cell; a

point between veins 3 and 4 postmedially, and a short oblique

line outwardly below costa. Hind wings: alarge spot in middle

of wing from within cell. Fore wings below fuscous; costal

margin yellow-brown, the extreme edge black; termen finely

yellow expanding at apex; the yellow discal spot extending

below vein 2 and expanding along submedian. Hind wings

below whitish yellow, the veins brown except the discocellular

and base of vein 5, the brown on veins 2 and 4 suffusing some-

what towards cell; cilia brown.

Expanse 23 mm.

Hab. Vurrialba.

Near b. faula Godm.

LERODEA ? RUPILIUS, sp.n. (Pl. LIV. fig. 10.)

2. Palpi pale brown tipped with white. Body above and

wings dark brown; body below whitish grey. Wings: spots

COSTA RICAN BUTTERFLIES. 361

semihyaline white; outer half of cilia greyish white with small

dark brown spots at veins. Fore wings: a large spot at end of

cell, and one postmedially between veins 2 and 3. a smaller out-

set spot between 3 and 4; three outer small. spots obliquely

between veins 6 and 9; a more opaque spot above submedian

beyond middle. Hind wings: a postmedial spot below vein 6,

crossed by vein 5. Fore wings below similar, the costa irrorated

with yellowish and white, the apex and termen to vein 2 with

white; the spot above submedian replaced by a large white

shade. Hind wings below thickly irrorated with greyish white ;

small whitish spots antemedially and postmedially below costal

vein; a siailae spot at end of cell; a small postmedial spot

between veins 2 and 3, and a linear spot below vein 2; the spot

below vein 6 as above ; the postmedial spots connected by a fine

brown shade, and a similar faint shade below end of cell.

Expanse 36 mim.

Hab. Guapiles.

The only species I know approaching this is one in the Godman

Collection labelled ‘“ phyllas Mab.” Without a male the position

is uncertain.

STYRIODES, gen. nov.

3. Antenne two-thirds as long as costa. Fore wings: costal

margin slightly sinuous; apex acute; termen rounded below

apex ; vein 2 from middle of cell; 3 near lower angle; 4 and 5

from lower angle, the latter upcurved at base ; 6 to 9 evenly

apart ; 10 and 11 wider spaced ; a downcurved brand below

median from vein 3 to below vein 2; a small medial brand above

submedian. Hind wings: anal angle slightly lobed; vein 2

beyond middle of cell; 3 and 4 from lower angle apart; 6 and 7

from upper angle.

Type of genus, Styriodes lyco Schaus.

SrYRIODES LYCO, sp.n. (PI. LIV. fig. 19.)

3. Body and wings above fuscous brown. Fore wings: brands

black; cilia tipped with yellow-ochre, wider at tornus. Hind

wings: cilia tipped with yellow-ochre, except at anal angle.

Wings below dark reddish brown, the inner margin of fore wings

dull dark brown; a fuscous ‘borin shade in ical behind and

below it.

Expanse 28 mm.

Hab. Poas,

RHINTHON LUCTATIUS, sp.n. (Pl. LIV. fig. 9.)

3. Body above black, the head shaded with dark metallic

green ; abdomen below with two yellowish white lines. Wings

fuscous brown. Fore wings: a very small greyish white post-

medial spot between veins 2 and 3, and one between 3 and 4

outset. Wings below not quite so dark. Fore wings: the two

spots whiter, better defined; a postmedial greyish brown shade

362 MR. WILLIAM SCHAUS ON

below vein 2. Hind wings: postmedial buff points between

veins 2 and 4.

Expanse 44 mm.

Hab. Guapiles, also Santa Rosa, Mexico.

In British Museum.

CoBALUS NIGRANS, sp.n. (PI. LIV. fig. 5.)

3. Palpi and thorax fuscous grey mottled with pale yellow ;

abdomen below with two fine whitish lines. Wings black-brown.

Wings below duller. Fore wings: a faint greyish white post-

medial line between veins 2 and 3, and a small outset spot

above 3; tornus shaded with fuscous grey. Hind wings: an

orange point in cell at end; similar outer points between veins 3

and 4, and just below 6.

Expanse 30 mm.

Hab. Guapiles.

CoBALUS LAUREOLUS, sp. n. (PI. LIV. fig. 3.)

3. Body fuscous clothed with dark brown hairs; the palpi

mottled with some whitish yellow hairs; abdomen below greyish

brown. Fore wings: veins medially tinged with fuscous. Hind

wings: basal half rather darker shaded. Wings below paler

brown. Fore wings: basal third except costa shaded with

fuscous; a triangular shade on costa near apex preceded by faint

greyish points between veins 6 and 9; tornus faintly paler

shaded. Hind wings: base, a postmedial fascia, and large mar-

ginal spots from apex to vein 2 darker brown ; a terminal darker

brown line.

Expanse 35 mm.

Hab. Guapiles.

CoBALUS LATERANUS, sp. n. (PI. LIII. fig. 3.)

3S. Palpi fuscous inwardly shaded with white. Body and

wings fuscous brown. Fore wings: spots semihyaline, white; a

large irregular spot at end of cell; a large postmedial spot be-

tween veins 2 and 3, and a small outset spot above 3; three small

outer spots below costa, outbent, the largest between veins 6 and

7; a small postmedial white spot above submedian. Fore wings

below fuscous, the costa, apex, and termen shaded with olive-

brown; the spot above submedian larger. Hind wings below

dark olive-brown ; a white spot in end of cell; a white point

subterminally between veins 3 and 4.

Expanse 32 mm.

Hab. San Mateo.

The fore wings above are like C. gabina Godm.

CoBALUS PINDAR, sp. n. (PI. LIV. fig. 18.)

3. Palpi olive-yellow irrorated with brown. Body and wings

above dark brown; abdomen below yellowish bistre. Fore

COSTA RICAN BUTTERFLIES. 363

wings: spots semihyaline whitish brown ; a large spot between

veins 2 and 3, its outer edge incurved ; a small spot between 3

and 4; three small spots between veins 6 to 9 in a straight out-

bent line from costa ; cilia tipped with bistre grey. Hind wings:

a yellow postmedial hyaline point between veins 3 and 4. Fore

wings below fuscous brown, the costal margin broadly, and cell

shaded with yellow-brown, the spots as above; a whitish post-

medial shade above submedian. Hind wings below brown; a

small faint fuscous shade at discocellular, and similar spots post-

medially from veins 2 to 7, the one between 3 and 4 showing the

semihyaline point.

Expanse 29 mm.

Hab. Port Limon, Juan Vinas.

In British Museum.

CoBALOPSIS LATONIA, sp.n. (PI. LIV. fig. 16.)

3. Body fuscous brown; palpi above whitish grey; abdomen

below pale buff. Wings dark brown. Fore wings: a fine

short whitish inbent streak below discocellular ; a small white

postmedial spot between veins 3 and 4, also above and below

vein 7. Hind wings: the veins fuscous. Fore wings below

fuscous in cell and ‘postmedially below vein 3, the inner margin

irrorated with greyish ; costa, apical space, and termen yellowish

brown. Hind wings below yellowish brown; a fuscous shade near

inner margin expanding at anal angle; five bluish white post-

medial spots from veins 2 to 7.

Expanse 33 mm.

Hab. Juan Vinas, Cachi.

In British Museum.

CENUS NAUSIPHANES, sp.n. (PI. LIV. fig. 6.)

3. Palpi mottled whitish grey and dark brown. Body fuscous

brown ; abdomen below whitish grey; legs yellow-brown. Wings

dark brown. Fore wings: the brand fine, black, inbent from

vein 3, and downcurved to submedian ; some yellowish scaling on

costa ; cilia tipped with grey. Hind wings: cilia yellow-grey.

Fore wings below fuscous at base, in, and below cell; costa yellowish

brown ; terminal half dull brown. Hind wings dull brown; a

broad medial whitish shade, widest on costa and not reaching

inner margin which is simply irrorated with whitish grey.

Expanse 25 mm.

Hab. Poas.

In British Museum.

MEGISTIAS XANTHO, sp. n. (PI. LIV. fig. 13.)

3. Body fuscous mottled with brown haus. Wings fuscous’

brown. Fore wings: a yellow-brown streak on costa from base

to just beyond middle ; a similar streak above submedian medially

extending towards base; a few similar scales outwardly beyond

364 MR. WILLIAM SCHAUS ON

cell, outbent along vein 6, vertical to below 5, inbent, expanding,

forming a spot between 4 and 3, and a larger spot between 3

and 2; cilia light brown spotted with black. Hind wings: some

yellow-brown hairs at base and on inner margin ; a broad similar

postmedial shade not reaching margins ; the spots on cilia narrow,

barely traceable. Fore wings below fuscous ; veins on costa, and

terminally, also a streak in cell, yellowish ; a postmedial yellowish

spot between veins 3 and 4, a vertical outer streak below costa,

and streak subterminally from veins 6 to 4; a terminal yellowish

line; base of cilia and spots black. Hind wings below greyish

tinged with lilacine, the veins yellowish; an outcurved dark

brown medial fascia, and large terminal spots, not reaching inner

margin.

Expanse 20 mm.

Hab. Juan Vinas.

In British Museum.

Voraves cotiso, sp.n. (Pl. LIII. fig. 6.)

3. Body above and wings dark brown ; throat whitish ; thorax

below greyish; abdomen below yellowish white. Fore wings:

costal margin to beyond middle bright olive-brown ; a similar

streak above and below submedian to middle, followed above

submedian by a small yellow spot; a yellow streak below median

from veins 2 to 8; a small spot between 3 and 4; an outer yellow

point between veins 6 and 7. Hind wings shaded with bright

brown, and olive-brown hairs. Fore wings below fuscous brown ;

yellow marking between veins 2 and 4 as above ; costa and veins

terminally yellow, the streaks longest at apex. Hind wings

below dull dark brown, the veins all yellow; a streak through

cell, and a broad yellow streak below cell and vein 2.

Expanse 24 mm.

Tab. San Mateo.

MoLo NEBROPHONE, sp. n. (PI. LIV. fig. 7.)

S$. Palpi orange-brown tipped with black. Body fuscous

partly clothed with golden-brown hairs; abdomen below yellowish

with a black ventral line. Fore wings black, except base of

costal margin to near middle, cell, inner margin to beyond

middle, postmedial spots from submedian to vein 4, a small outer

spot above 4, and spots above and below vein 7, which are orange-

brown; a slight fuscous streak in cell. Hind wings orange-

brown; costal margin broadly black with a medial inbent pro-

jection; inner margin broadly black ; a broad black streak below

cell to middle of wing; a fuscous shade on termen towards anal

angle. Fore wings below deep yellow; some black on base of

cell; a broad black shade below cell, upbent on discocellular ;

termen black from vein 2 to inner margin; apex shaded with

reddish, the outer spots at vein 7 larger. Hind wings

below yellow on basal half; the inner margin broadly black

COSTA RICAN BUTLERELIES. 365

irrorated with yellow-brown; outer half reddish with oblong

outer yellow spots from vein 6 to below vein 2; subterminal

fuscous shading between veins 2 and 4; a darker streak just

beyond cell between 4 and 6.

2 fuscous brown ; a small yellow spot at upper angle of cell ;

traces of some postmedial yellowish spots. Fore wings below

fuscous; costal margin yellowish ; apex tinged with dull reddish ;

« yellow spot at end of cell; a large postmedial yellow spot below

vein 2, and smaller spots above 2 and 3; small yellow outer spots

above and below vein 7. Hind wings below as in male, the ter-

minal space duller red, limited by a yellow streak before the

fuscous inner space.

Expanse, d 27 mm., 9 33 mm.

Hab. Cayrillo.

The female from the Bejuco river, Panama.

Near MW. herewa Hew.

Moto? APELLA, sp. n. . (Pl. LITT. figs. 4, 5.)

3. Palpi, head, collar, and patagia orange-brown. Thorax

and abdomen fuscous, the latter with orange-brown hairs

laterally; underneath yellowish. Wings black, the cilia and

markings orange-brown. Fore wings: the costal margin broadly

orange-brown, entering cell, narrowing towards black apical

space; a fascia from vein 8 narrow and outangled between

veins 5 and 6, then broader and expanding to middle of inner

margin, cut by black veins. Hind wings: base and inner margin

clothed with orange-brown hairs; a broad postmedial fascia, not

reaching apex or inner margin. Wings below brownish yellow ;

some terminal black points. Fore wings: the outer margin

broadly darker with subterminal black shadings from vein 3 to

inner margin; a black space on basal third below cell; a fine

black line on discocellular.. Hind wings: antemedial, postmedial,

and terminal darker shadings on interspaces.

2. Wings above tinged with brown, the fascia much reduced,

more macular, yellowish; the costa of fore wings with some

yellowish irrorations. Wings below dark olive-brown, the veins:

paler ; the postmedial fascia on fore wings broad, as in male, but

yellower. Hind wings with postmedial dull yellowish spots om

interspaces.

Expanse, ¢ 37 mm., @ 45 mm.

Hab. Cavrillo.

The male above is very much lke J/.? humeralis Mab., but

differs considerably on the underside of hind wings.

In British Museum.

CARYSTUS SUBRUFESCENS, sp.n. (PI. LIV. fig. 11.)

2. Palpi whitish buff. Head light brown. Palpi shaded with

dull reddish brown. Abdomen fuscous brown, the hairs at base

dull greyish green. Wings fuscous brown. Fore wings: costal

366 MR. WILLIAM SCHAUS ON

margin at base shaded with reddish; spots semihyaline whitish

yellow ; a streak in end of cell above median, and a spot below it

to vein 2, its outer edge incurved; an elongated postmedial spot

above vein 4, and a smaller spot below it; small outer spots

between veins 6 and 8; a small yellowish spot above submedian

just beyond middle; cilia yellowish at tornus. Hind wings:

cilia tipped with pale yellow. Fore wings below black; the

costal margin, apical space beyond cell, and termen to vein 2

brownish red ; a yellowish white shade medially above submedian.

Hind wings below brownish red; veins finely pale yellow; a

fuscous shade from base before fold, expanding at anal angle.

Expanse 36 mm.

Hab. Guapiles.

Near C. commodus Plotz, distinguished by the pale veins on

hind wings below.

EPInosis, gen. nov.

S$. Palpi upturned, hairy. Antenne more than half the length

of fore wing, thickened towards end which is curved and pointed.

Wings broad. Fore wings: outer margin evenly curved; vein 2

from middle of cell; 3 close to lower angle; 4 from lower angle;

5 from below middle of discocellular, which is slightly oblique ;

6, 7, 8 close together; a narrow brand below median from

vein 3 to 2. Hind wings: veins 3 and 4 from lower angle

slightly apart ; 6 and 7 from upper angle.

Type of genus, Pamphila angularis Moschl.= Hesperia parvi-

puncta Méschl. fide Godman, (Pl. LITT. fig. 1.)

A common species in Costa Rica not mentioned in the ‘ Biologia’,

and agreeing perfectly with other specimens from the Guianas.

FAuGa ? HERMIONE, sp.n. (Pl. LIV. fig. 20.)

¢. Palpi black, the second joint with a lateral white shade.

Body above fuscous with orange-brown hairs; abdomen below

yellowish white with a fine interrupted medial and outer black

line. Fore wings black; base of costa shaded with brown ; sub-

costal broadly orange-brown, not reaching apex, beyond cell with

short upbent lines; a broad postmedial orange-brown shade

inbent from vein 4 to submedian, its edge uneven, and joined by

a similar streak below cell; cilia orange-brown partly spotted

with black, entirely black at apex. Hind wings and cilia orange-

brown, the margins black; a black streak from base near inner

margin. Wings below orange-brown. Fore wings: a black

streak through cell, and on interspaces above and below vein 5;

tornus black ; subterminal black spots on other interspaces, and

streaks on costa towards apex. Hind wings: a terminal black

line; a streak below cell and vein 2, and one at apex ; subterminal

black spots above and below vein 3.

COSTA RICAN BUTTERFLIES.

The female has the orange - brown on fore wings

restricted.

Expanse 26 mm.

Hab. Carillo.

In British Museum.

367

EXPLANATION OF THE PLATES.

Pirate L.

Fig.1. Napeogenes hemisticta. Fig. 6. Huptychia drymo.

2. Ithomia bolivari. 7. Hresia sticta.

3. Pteronymia fumida. 8. 6 CHa Bx

4, Huptychia agnata. 9. Phyciodes phlegias, ?.

5. Catargynnis dryadina.

Pxuate LI.

Fig. 1. Hurygona leucon, 3. Fig. 6. Victorina steneles, ab. stygiana.

2. Hupetoieta poasina. 7. Hurygona matuta.

3. Hurygona leucon, 2. 8. BS mystica.

4. Eubagis hecuba, 6. 9. Phyciodes dora.

5. 35 sp oF 10. Perisama barnesi.

Prate LII.

Fig. 1. Thecla guapila. Fig. 6. Thecla hypocrita.

2. Enantia lua costaricensis. Yo by 9 UGOBs

3. Acmepteron poasina, 2. 8. Archonias intermedia, 3.

4. Thecla melma. 9. Thecla subflorens.

6. Pieris limona, 2. 10. 4, amphrade.

Prats LIII.

Fig. 1. Epinosis angularis. Fig. 8. Eurygona leucophryna.

2. Corrachia leucoplaga. 9. Hermathena oweni.

3. Cobalus lateranus. 10. Phyciodes niveonotis, 2.

4, Molo? apella, 3. 11. Charis subtessellata.

Suns 33 2 12. Mesosemia albipuncta.

6. Vorates cotiso. 13. Charis turrialbensis.

7. Heliconius galanthus, ab. sub-

rufescens.

PuatEe LIV.

Fig. 1. Telemiades mnemon. Fig. 11. Carystus subrufescens.

2. Butleria lysis. 12. Bolla sodalis.

3. Cobalus laureolus. 13. Megistias xantho.

4. Bolla machuca. 14. Cyclosemia subcerulea.

5. Cobalus nigrans. 15. Butleria lethea.

6. Enus nausiphanes. 16. Cobalopsis latonia.

7. Molo nebrophone, : 17. Ate canace.

8. Echelatus lucina. 18. Cobalus pindar.

9. Rhinthon luctatius. 19. Sturiodes lyco.

10, Lerodea ? rupilius. 20. Falga ? hermione.

368 MR. C. TATE REGAN ON

27. A Collection of Fishes made by Professor Francisco

Fuentes at Haster Island. By C. Tare Reaay, M.A.,

F.Z.8.

[Received March 3, 1913: Read April 8, 1913.]

(Plates LV.—LX.*)

INDEX. Page

Acanthistius fuscus, sp. n. seechipiaeetaeieeaee aca ae MeO OS,

Ganellopsicemims te teenrcre teen ccichen sieoteeriinos riences tee 369

Wabrichehysmfuentest spe sre acs-ccuneesseasdekeeee ee eee OL

AlGMOIDSES FOUNALOWP, So Wo goo cc0800 28000 a00000 900160700000 860 006 371

Pseudomonacanthus paschalis, sp. ne ...... 0. ccc ceecee eevee =—872

IBAChySteliusronientale spo meee ereeeeeae cee OTS

Easter Island lies in 27° 6'S. and 109° 17' W.; it is very isolated,

distant about 2500 miles from the coast of Chile and nearly

1000 miles from Ducie, the nearest island of the Paumotu

Archipelago. A small collection of fishes made at Easter Island

in April, 1911, by Professor Fuentes, is of considerable interest.

Tt includes examples of eleven species : seven belong to genera

or species of the tropical Pacific, three are new and are related

to species known from Norfolk Isiand or from New South Wales,

and one belongs to a new genus whose affinities are with Zephreops —

of New South Wales and the southern coasts of Australia. No

Chilean species are represented, and it is of interest to note that

in the case of Chilean genera, Acanthistius and Labrichthys,

relationship is with the Norfolk Island species and not with the

Chilean ones.

SERRANID &.

1, AcANTHISIIUS FuscUs. (PI. LV.)

Acanthistius cinctus (non Giinth.) Kendall & Radcliffe, Mem.

Mus. Comp. Zool. xxxv. 1912, p. 107.

‘¢ Capuco.”

Depth of body 2% in the length, length of head 22. Snout 12

as long as diameter of eye, which is 54+ in the length of head ;

interorbital width 7. Lower jaw a little projecting : maxillary

extending to below middle of eye, its width distally nearly ?

diameter of eye; occiput, interorbital region, cheeks, and opercles

covered with ciliated scales; rest of head naked, or with minute

imbedded scales; 9 or 10 gill-rakers (and some rudiments) on

lower part of anteriorarch. Scales strongly ciliated, 105 in a longi-

tudinal series above the lateral line, 55 in the lateral line, 17

from base of spinous dorsal to lateral line. Bioveal XIII 15;

fourth to sixth spines longest, # length of head, ee soft

rays. Anal III 8; second : spine longest and strongest, - + length

* For explanation of the Plates see p. 373.

EPO VOM pe ly exe

A.A, Searle, del et lith. Huth,imp.

PSEUDOMONACANTHUS PASCHALIS.

Pent

eee

ers

“meant

AERA t

ie Hy ye 3

iui Is

P7196. Pl Lix.

AH. Searle, del et Lith. Huthimp.

ANAMPSES PULCHER.

“Ss Gil waned! ONT

aittmwWitlad SpAHoOAINVNV

ISH LNANY SAH LAO aVY 1

isp)

lem

‘des “qa

‘THAT ‘Tad @16ET SZ a

aes

en aN

3 i ‘ ene i Rares Peete ume er

BUC Se A teed hou te pe age ; eral lessee at

Pee ents: et ‘3 ng Pease 7 i z Ss aeeh lar ented fy

Saye ka <4 ' :?

Rei

Nae

ata,

eeu

12 Zl Qls), Pw IVAN

AH. Searle del et lith, ee Huth imp,

GIRELLOPS NEBULOSUS.

1 Zor), WSN) IE, IENWAL;

A.H.Searle,del et lith. Huth,imp.

KUHLIA MUTABUNDA.

J 4 hSy, LOS), JELLY,

A.H.Searle del. etlith.

Huthimp.

AGAN Pols LoS PUSGuS:

FISHES FROM EASTER ISLAND, 369

of head. Pectorals and pelvics subequal, a little more than

length of head. Caudal rounded. Dark brownish.

A single specimen, 230 mm. in total length.

This species is very closely related to A. cinctus Giinth. from

Norfolk Island, described from a specimen of nearly. the same

size. Comparison shows that 4. cinctus differs in the larger head

(22. in the length), higher dorsal fin (longest spines } the length

of head), longer second anal spine (3 the length of head), longer

and stronger maxillary (extending to below posterior } of eye, its

width 7 the diameter of eye), and the banded coloration.

Die

KUHLIIDA.

2. KuHLIA MUTABUNDA Kendall & Radcliffe. (Pl. LVI.)

Mem Mus. Comp. Zool. xxxy. 1912, p. 105, pl. iii. f. 1.

‘“¢ Mahori.”

Depth of body 23 in the length, length of head 31. Snout 2

diameter of eye, which is 22 in length of head; interorbital width

33. Lower jaw a little prominent; maxillary extending to below

anterior + of eye; cheek with 4 series of ctenoid scales; angle and

lower edge of preoperculum finely denticulated; 25 gill-rakers on

lower part of anterior arch. 50 scales in the lateral line, 6 between

sheath at base of anterior dorsal spines and lateral line, 13 from

lateral line to pelvic fin. Dorsal X11; fourth and fifth spines

longest, 4 length of head; last 3 length of head, a little shorter than

seventh, considerably shorter than longest soft rays. AnalIII 11;

third spine a little longer than second, as long as tenth dorsal ;

base of fin a little more than $ length of head or 11 its distance

from caudal. Pectoral 2 length of head. Caudal deeply forked.

Caudal peduncle 12 aslongas deep. Silvery, back darker ; caudal

blackish posteriorly. —

A single specimen, 190 mm. in total length.

Related to A. humilis De Vis, from Queensland and the Fiji

Islands, and to K. sandvicensis Steind., known from the Sandwich

Islands and Society Islands.

GIRELLIDS.

GIRELLOPS, gen. nov.

Related to Tephrwops, but with uniserial teeth.

3. GIRELLOPS NEBULOSUS Kendall & Radcliffe. (Pl. LVIL.)

Girella nebulosa Kendall & Radcliffe, Mem. Mus. Comp. Zool.

RAV, OO ple lites 2 pl. tive tt. Vo 2.

‘“* Maahaki.”

Depth of body 22 in the length, length of head 4. Diameter

of eye 4 in length of head, interorbital width 2. Scales on cheeks,

temporal region, and upper) edge of operculum; rest of head

naked, Teeth in jaws fixed, uniserial, incisor-like, the anterior

370 MR. C. TATE REGAN ON

ones truncated, the lateral ones obtusely tricuspid; palate toothless ;

maxillary extending to below nostrils. Depth of preorbital 2

diameter of eye. Gill-membranes broadly united, forming a fold

across isthmus. Dorsal XVI 12, scaly at the base ; spines stout,

subequal from the fifth to the last, which is 3 the length of head

and a little shorter than the anterior soft rays. Anal III 10;

second and third spines subequal, 4 length of head; second or

third soft rays + as long as head. Caudal slightly emarginate.

Pectoral as long as head. Seales ciliated, 78 in a longitudinal

series, 12 or 13 between sheath at base of anterior dorsal spines

and lateral line. Dark greyish, with traces of darker spots and

markings.

The above description from a specimen of 270mm. A young

example, 43 mm. in total length, has the same number of scales

and fin-rays, but all the teeth are tricuspid, and the markings

obscurely indicated in the adult take the form of 7 dark cross-

bars and scattered spots on the body, and longitudinal bands on

the dorsal fin. The nearest ally of this species appears to be

Tephreops richardsonit, which ranges from New South Wales to

Western Australia.

CARANGIDS2.

4, CARANX CHEILIO Snyder.

Caranx guara (non Bonn.) Kendall & Radcliffe, Mem. Mus.

Comp. Zool. xxxv. 1912, p. 99.

“« Poopo.”

Originally described from a specimen 30 inches long obtained

in the market at Honolulu; the example now recorded from

Easter Island is 12 inches long.

PoMACENTRID A.

5. Pomacentrus inornatus. (PI. LVI. fig. 1.)

““ Cototi.”

2 Pomacentrus niomatus De Vis, Proc. Linn. Soc. N. 8. Wales,

vill. 1883, p. 451.

2 Pomacentrus inornatus Seale, Bull. U.S. Fish. Bureau, xxv.

UQOS; jos BBs

Pomacentrus jenkinst Kendall & Radcliffe, Mem. Mus. Comp.

Zool. xxxv. 1912, p. 132.

Depth of body 14 in the length, length of head 32. Snout

a little shorter than eye, the diameter of which is 33 in the

length of head; interorbital width 3. Mouth subterminal.

Head scaly, except lips and extremity of snout; 2 series of scales

on cheek and 1 on lower limb of preoperculum. Preorbital

entire; suborbitals and preoperculum denticulated. 28 scales

in a longitudinal series, 20 in the lateral line, 3 between middle of

spinous dorsal and lateral line. Dorsal XIII 16; spines subequal

FISHES FROM EASTER ISLAND. By!

from fourth to last, which is 4 length of head; middle soft rays

nearly as long as head. Anal II 13. Pector: i] as long as bead,

Caudal deeply emarginate. Caudal peduncle much deeper than

long. Brownish; scales black-edged; vertical fins blackish ;

a small blackish spot above axil of pectoral.

A single specimen, 100 mm. in total length.

De Vis’s description is so incomplete that it is impossible to

identify his species with certainty. He described it as probably

from the South Seas, and Seale has recorded this, or a related

species, from Samoa. P. jenkinsi; from the Sandwich Islands, as

described and figured by Jordan and Evermann, seems to have

a broader and more convex interorbital region, deeper preeorbital,

and less deeply emarginate caudal fin.

LABRIDZ.

6. CHEILIO INERMIS Forsk,

“ Ure-ure.”

A species of wide distribution, ranging from Africa to the

Western Pacific.

7. JULIS UMBROSTIGMA Riipp.

Sa acojues

This also occurs heowlivont the tropical Indo-Pacific.

8. LABRICHTHYS FUENTESI, sp. n. (Pl. LVIIT. fig. 2.)

Pseudolabrus inscriptus (non Richards.) Kendall & Radcliffe,

Mem. Mus. Comp. Zool. xxxv. 1912, p. 137, pl. v. f. 2, pl. vi. f. 1.

** Cootea,”

Depth of body 23 in the length, length of head 33. Snout 132

as long as diameter of eye, one is 5d in the length of head

and a little less than the interorbital width. Upper jaw with a

posterior canine on each side and a pair of strong anterior

canines; lower with 2 pairs of anterior canines; cheek with

5 series of scales. 25 scales in the lateral line. Dorsal IX 11;

seales extending on to basal part of fin. Anal IIT 10. Pectoral

2 length of head. Caudal subtruncate. Body with about

8 irregular cross-bars ending in blotches on basal part of dorsal

fin ; head and upper part of body with numerous small pale

spots or vermiculations; soft dorsal and anal with broad

marginal dark bands including numerous pale spots with dark

centres ; base of pectoral fin blackish.

A single specimen, 165 mm. in total length.

This species is nearest to L. luculenta NRichards., from Norfolk

Island and Australia.

9. ANAMPSES PULCHER, sp.n. (Pl. LVIIJI. fig. 3 & Pl, LIX.)

“ Mouri.”

Depth of body 2? in the length, length of head 34. Diameter

Proc. Zoot. Soc.—1913, No. XX VI, 26

BD MR. C. TATE REGAN ON

of eye 7 in length of head, 25 in fenguln of snout. Interorbital

region strongly convex, its width + the length of head. <A

naked strip running backwards from ‘occiput to origin of dorsal

fin. 28 scales in the lateral line. Dorsal 1X 12. Anal IIT 12

Pectoral # the length of head. Caudal subtruncate. Each scale

with a blue spot surrounded by a dark ring; head with blue

horizontal stripes or series of spots; one through upper part of

eye; another from end of snout through lower edge of eye, a

third across the cheek ; vertical fins with spots like those on the

body and with a narrow pale blue or white margin; base of

pectoral blackish.

A specimen of 205 mm., from Haster Island, differs from one

of the same size of A. cwruleopunctatus Riipp. of the tropical

Indo-Pacific, especially in the broader and more convex inter-

orbital region, the incompletely scaled nape, and the different

coloration of the head, A. cewruleopunctatus having vertical

stripes running from eye to suboperculum.

A second example of A. pulcher, 165 mm. in total length,

from Tahiti (Wragge), is extremely similar to the type, but has

the interorbital region less convex, and the naked median strip

on the nape less elongate, not quite reaching the dorsal fin.

BLENNIIDS.

10. SALARIAS ARENATUS Bleek.

Alticus striatus (? Cuv. & Val.) Kendall & Radcliffe, Mem. Mus.

Comp. Zool. xxxv. 1912, p. 134.

‘¢ Patuki,”

A species of the Kast Indian Archipelago and Islands of the

South Pacific.

BALISTID.

11. PSEUDOMONACANTHUS PASCHALIS, sp. n. (PI. LX.)

“ Coreba.”

Seales small, irregular, covered with minute spines. Depth

of body 27 in the length. Snout produced, with upper profile

a little convex, more than 3 times as long as eye and a little less

than 4 the length of the fish. Lower angle of gill-opening below

anterior margin, base of pectoral below middle of eye. Dorsal

spine nearly as long as snout, above posterior 4 of eye, antero-

posteriorly compressed, with a series of 24 small barbs on each

side; soft dorsal of 35 rays, elevated anteriorly; anal rays 34.

Caudal rounded. Greyish, with numerous scattered small dark

spots.

A single specimen, 300 mm. in total length.

This species is related to the Australian P. ayraudi Quoy &

Gaim.

FISHES FROM EASTER ISLAND. 373

EXPLANATION OF THE PLATES.

Prater LV.

Acanthistius fuscus.

PLATE LVI.

Kuhlia mutabunda.

PuatE LVII.

Girellops nebulosus.

(a) Teeth. (6) Young.

Prater LVIII.

Fig. 1. Pomacentrus inornatus.

2. Labrichthys fuentesi.

3. Anampses pulcher; head of specimen from Tahiti.

Puate LIX.

Anampses pulcher.

Prats LX.

Pseudomonacanthus paschalis.

APPENDIX.

[It was not until the above was in the press and the plates

had been drawn, that I saw Kendall and Radcliffe’s recent memoir

on fishes from the eastern tropical Pacific. I have altered my

paper only by taking their names for the new species of Kuhlia

and Girellops and by inserting references to their work when

our determinations did not coincide. They obtained nine of the

eleven species, all but Anampses pulcher and Pseudomonacanthus

paschalis. They also got a number of other species from Easter

Island mostly known previously from the tropical Pacific, viz.

Myripristis pralinius Cuv. & Val., Holocentrum punctatissimum

Cuv. & Val., Decapterus sancte-helene Cuv. & Val., Cyphosus

cinerascens Forsk., Pseudupeneus multifasciatus Quoy & Gaim.,

Thalassoma purpurewm Forsk., Teuthis umbra Jenkins, Kellogella

oligolepis Jenkins, Alticus variolosus Cuv.& Val., and Xanthichthys

lineopunctatus Hollard. ‘The identification of a species of

Monacanthus with the Japanese M. cirrifer Schleg. seems open

to question, and still more so the determination of an Eel as

Gymnothorax dovi Giinth., as the type from Panama appears to

differ in many respects from the specimens from Easter Island,

which, from the data given, seem more like the Indo-Pacific

G. meleagris Shaw.

Finally, the resemblance to Norfolk Island is increased by the

addition of two species—the one, 7’rachypomamacracanthus Giinth.,

apparently not distinct from the Norfolk Island form, the other

a Bathystethus, requiring a new specific name.

BAtHYSTETHUS ORIENTALE, Sp. 0.

Platystethus cultratum (non Schneid.) Kendall & Radeliffe,

Mem. Mus. Comp. Zool. xxxv. 1912, p. 97, pl. ii. £. 3.

Closely related to B. cultratum, but head longer, 4 in length

26%

374 MR. C. TATE REGAN ON

of fish (42 in B. cultratum), eye larger, 35 in length of head or

nearly equal to its distance from edge of lower limb of pre-

operculum (43 in head and 14 in its distance from lower edge

of preoperculum in ZB. cultratwm), and fin-rays more numerous.

Dorsal VIII, I 27; anal III 32 (VIII, I 23-25 and IIT 28-30

in B. cultratum).

The type from Easter Island measures about 200 mm. in total

length (to end of middle caudal rays), the examples of B. culiratum

from Norfolk Island that I have examined 170 to 180 mm.|

28. A Revision of the Fishes of the Genus Kuhlia.

By C. Tate Regan, M.A., F.Z.8.

[Received March 4, 1913: Read April 8, 1913.]

(Text-figures 68 & 69.)

INDEX. Page

Kuhlia cerulescens; sp. 1. ......0.0-deccevee eee eeesse ee 316

RO TOAUGOG On CID 1G ceosudeededbaeecbsocdecoue onc nooosanmncas, | SU

TRG, SDUCTIIGIS, SDs We 306600.39500500000 000 cens0n000000acc00a | SI)

I have been led to undertake a revision of the species of the

Indo-Pacific genus Auhlia Gill, owing to the difficulty I

experienced in determining the correct name for a fish of this

genus from Haster Island (Kuhlia mutabunda).

To the synonymy of Auhlia as given by Boulenger (Cat. Fish,

i. p. 36) may be added Boulengerina (non Dollo) Fowler (Proce.

Acad, Philad. 1906, p. 512), and Safole Jordan (Proc. U.S. Nat.

Mus. xlii. 1912, p. 655).

These names are intended to distinguish the more silvery

and more strictly marine species with more numerous gill-rakers

(Safole) from the spotted forms that enter rivers and have fewer

gill-rakers (Kuhlia). There is but little justification for such

a generic division. . boninensis is intermediate between the

two groups in the number of gill-rakers; A. marginata may

sometimes be unspotted, and in the form and coloration of the

caudal fin is nearer to K. humilis than to A. rupestris, and

K. splendens, a silvery, unspotted species, with numerous gill-

rakers, is described below from the fresh-waters of Rodriguez.

Synopsis of the Species.

I. Lateral line of 40 to 45 scales (+ 4 or 5 on base of caudal

fin); 16 to 19 gill-rakers on lower part of anterior

arch ; body usually spotted.

A. Caudal fin slightly emarginate ; middle rays } or ¢ as

long as longest; lobes rounded. Dorsal X 11.

Anal III 10. :

1. Maxillary extending to below middle of eye (young)

Oe loenv Oral (Exh) Goo coconaons xono00900800020 940009 0005000

2. Maxillary not or barely reaching middle of eye in

adult.

1. rupestris.

FISHES OF THE GENUS KUHLIA. 375

Upper opercular spine nearly as strong as lower ; dorsal origin

in advance of pelvies.., 53 Seavstea asec harseceenpeierae :

Lower opercular spine ins the nornsteg agra origin above

base of pelvics .......... Job soapp>aanmounghooehseMCLA OCA IOce

BG. Caudal fin more Heonly 6 emar roinattes suave rays $

to 2 as long as longest ; lobes pointed. Dorsal X

10-12, Anal III 11-13.

Maxillary extending beyond middle of eye ............0000:0020 4. wnvilliz.

Maxillary not or barely reaching middle of eye .................. 5. marginata.

II. Lateral line of 46 to 56 scales (+ 4 or 5 on base of caudal

fin) ; no spots on body.

A. 21 or 22 gill-rakers on lower part of anterior arch ... 6. boninensis.

B. 24 to 29 gill-rakers on lower part of anterior arch.

1°) Dorsal xe 13=145 > Amal WW V2=W3) eoe..s 22k seis 7. caudovittata.

2. Dorsal X11. Anal III 11; caudal fin plain, usually

with dark posterior margin.

a. Middle caudal rays more than + as lone as

longest ; 4 scales from sheath at base of spinous

dorsal to nearest part of lateral line.

Kye a little shorter than postorbital part of head .............. 8. splendens.

Hye as long as postorbital part of head ................60.0.se0cc0 ee 9. humilis.

6. Middle caudal rays less than 3 as long as longest ;

5 or 6 scales from sheath at base of spinous

dorsal to nearest part of lateral line.

Anal nearly as long as ne much longer than caudal

peduncle eer. s-s-- ener ... 10. sandvicensis.

Anal } length of head, a en ener: thant aida Peannicls .. IL. mutabunda.

8. Dorsal X 9-11. Anal III 10-11. Caudal fin with

five blackish bands, one median and two on each

WO bey ee Se: i Settee do astiiec adem. satis satcnasth aceoncs eee bas MUNG.

. cerulescens.

1. KUHLIA RUPESTRIS.

Oentropomus rupestris Lacep. Hist. Nat. Poiss. iv. pp. 252,

273 (1802).

Kuhlia rupestris (part.) Bouleng. Cat. Fish. i. p. 36 (1895).*

Kuhlia rupestris hedleyi Ogilby, Proc. Linn. Soc. N. 8. Wales,

Sul, WD 06 WO

Depth of body 23 to 3 in the length, length of head 23 to 33.

Snout ?# to 13 diameter of eye, which is 37 to 54 in length

of head; interorbital width 3 to 38. Maxillary extending to

below middle of eye (young) or beyond. 16 to. 19 gill-rakers on

lower part of anterior arch. 40 to 44 scales in lateral line, 4 or

43 from scaly sheath at base of spinous dorsal to highest part of

lateral line, 9 to 11 thence to ape of pelvic fin. Dorsal X 11 ;

fourth or fifth spine longest, 1 to more than 2 length of head ;

last as long as sixth, seventh, or eighth, 7 to more than 2 2 length

of head. Anal III 10; third spine usually shorter than tenth

dorsal; base a to 4 length of head, longer than caudal peduncle.

Pectoral 4 to = length of head. Caudal peduncle as long as, or

longer than, deep. Bluish silvery ; each scale usually with a

dark spot ; each lobe of caudal and soft dorsal with a blackish

* The synonymy is as given by Boulenger, except for Dules fuscus of Sanvage,

which is a distinct species.

376 MR. C. TATE REGAN ON

or brownish spot in young, these spreading to form broad marginal

bands in adult ; soft dorsal and upper and lower margins of

caudal often narrowly edged with white ; soft vertical fins usually

with one or more series of spots.

Indo-Pacific, from Africa to the Paumotu Archipelago. Twenty-

six specimens, 85 to 400 mm. in total length. :

2. KUHLIA CHERULESCENS, sp.n. (Text-fig. 68.)

Kuhlia rupestris (part.) Bouleng. Cat. Fish. i. p, 36 (1895).

Depth of body 3 in the length, length of head 3}. Snout ? dia-

meter of eye, which is 32 in length of head; interorbital width 33.

Maxillary extending to below middle of eye; upper opercular

spine nearly as strong as lower; 7 gill-rakers on lower part of

anteriorarch. 41 scales in lateral line, 44 from sheath at base of

Text-fig. 68.

Head of Kuhlia cerulescens.

spinous dorsal to highest part of lateral line, 95 thence to base of

pelvic fin. Dorsal X 11; origin in advance of base of pelvies ;

fifth spine longest, 2 length of head; last as long as seventh,

3 length of head. Anal III 10; third spine 3 length of head ;

base ?# length of head or 12 as long as caudal peduncle. Pectoral

less than #2 length of head. Caudal slightly emarginate, with

rounded lobes. Caudal peduncle 1+ as long as deep. Bluish

FISHES OF THE GENUS KUHLIA. 377

silvery; a dark spot at base of each scale ; cheeks and opercles

spotted ; soft dorsal and anal with longitudinal bands or series of

spots; caudal with broad dark band posteriorly.

Solomon Islands.

A single specimen, 290 mm. in total length, from Stirling

Island.

This species is near K. sawvagii, but has the coloration of

K. rupestris. Sauvage’s figure of the former agrees with the

example in the British Museum collection, and differs from the

type of K. cerulescens in the much smaller upper opercular spine

and the more posterior origin of the dorsal fin.

3. KUHLIA SAUVAGII, sp. n.

Dules fuscus (non Cuv. & Val.) Sauvage, Hist. Madag., Poiss.

p. 149, pl. xv. f. 4 (1891).

2 Moronopsis fuscus Steind. Sitzungsb. Akad. Wien, Ixxxu.

1881, p. 240.

Depth of body 22 in the length, length of head 33. Snout

2 as long as diameter of eye, which is 3 in the length of head and

equal to the interorbital width. Maxillary extending to below

anterior + of eye; lower opercular spine much stronger than

upper; 16 or 17 gill-rakers on lower part of anterior arch. 44

scales in lateral line, 4 between sheath at base of spinous dorsal

and arch of lateral line, 10 thence to base of pelvic fin. Dorsal

X 11; origin above base of pelvics; fifth spine longest, 2 length of

head ; last nearly as long as seventh, 4 length of head. Anal IT

10; base nearly as long as head or nearly twice as long as caudal

peduncle ; third spine less than 3 length of head. Pectoral ?

length of head. Caudal slightly emarginate. Caudal peduncle

a little longer than deep. Silvery, back darker; sides with

scattered rounded or erescentic reddish-brown spots; soft dorsal

and anal spotted ; caudal with dark membrane and pale rays.

Madagascar.

The above description is based on a single specimen, 150 mm.

in total length, from Imerina, Madagascar, which evidently

belongs to the same species as the example of 300 mm., also from

Madagascar, figured by Sauvage. This larger fish has, of course,

shorter spines, a smaller eye, ete., and in it the maxillary almost

reaches the vertical from the middle of the eye.

Dr. Pellegrin has kindly examined the types of Dules fuscus

Cuv. & Val., two examples only 4 inches long, and writes that the

maxillary extends slightly beyond the vertical from the middle of

the eye. There can be little doubt but that these are specimens

of K. rupestris.

4, KUHLIA URVILLII.

Kuhlia urvillii Bouleng. Cat. Fish. i. p. 38 (1895).

This species is based on a coloured figure in Dumont D’Urville’s

‘Voyage au Pole Sud,’ which represents a fish of 160 mm. very

similar to K. marginata, but with the last dorsal spine longest of

378: MR. C. TATE REGAN ON

all, 2 the length of head, and with the maxillary extending to

below the posterior part of the eye.

5. KUHLIA MARGINATA.

Dules marginatus Cuv. & Val. Hist. Nat. Poiss. ii. p. 116,

pl. 1. (1829).

Dules maculatus Cuv. & Val. op. c. vol. vii. p. 475 (1831).

Dules malo Cuv. & Val. t. ¢. p. 479.

Dules mato Lesson, Voy. Coquille, Zool. ii. p. 223 (1831).

Dules leuciscus Jenyns, Zool. Beagle, Fish. p. 17 (1842).

Kuhlia marginata Boulenger, Cat. Fish. i. p. 38 (1895)

(with synonymy).

Kuhlia malo Kendall & Goldsborough, Mem. Mus. Comp.

Zool. xxvi. 1911, p. 280, pl. 1. f. 2.

Kuhlia maculata Kendall & Goldsborough, ¢. c. p. 281, pl. 11.

ity The

Depth of body 22 to 31 in the length, length of head 8 to 3.

Snout 4 to ? diameter an eye, which is 22 to 34 in length of

head ; interorbital width 3 to 32. ee extending to below

anterior 1 of eye. 16 to 18 gill-rakers on lower part of anterior

arch. 40 to 45 scales in lateral line, 4 or 44 from sheath at base.

of spinous dorsal to highest part of lateral line, 8 or 9 thence to

base of pelvic fin. Dorsal X 10-12; fifth spine usually longest,

4 to 2 length oe head ; last as long as sixth, seventh, or eighth,

Pi to more than 2 2 length of head. Anal III 11-13; third spine

shorter than tenth dor sal; base nearly as long as ‘head, nearly

twice as long as caudal ’ peduncle. Pectoral 2 to # 2 length of

head. Caudal moderately emarginate, the middle rays 4 to 2

as long as longest. Caudal peduncle longer than deep. Silvery,

usually dark spotted above lateral line and sometimes below it ;

soft dorsal and anal dark edged, at least anteriorly, often dark

spotted basally; caudal with dark posterior margin and with

dark spots or a band parallel to posterior edge ; soft dorsal, anal,

and upper and lower margins of caudal often narrowly edged

with white ; pelvics sometimes spotted. .

Malay Archipelago and South Pacific.

Numerous examples, 75 to 160 mm. in total length, including

the types of D. leuciscus, kindly lent to me by Mr. L. Doncaster.

Some examples: are almost covered with spots, others are

silvery with scarcely a trace of spots, the variation in this respect

recalling our British Trout; probably the silvery examples are

marine, whilst the strongly spotted ones may have been for a

long time in fresh water.

6. KUHLIA BONINENSIS.

Kuhlia caudovittata (part.) Bouleng. Cat. Fish. i. p. 41 (1895).

Dules marginatus boninensis Fowler, Proc. Acad. Philad. vin.

1906, p. 510, fig.

Depth of the body nearly 3in length, length of head 33. Snout

FISHES OF THE GENUS KUHLIA. 379

2 diameter of eye, which is 3 in length of head; interorbital

width 33. Maxillary extending to below anterior 7 of eye.

22 gill-rakers on lower part of anterior arch. 52 scales in lateral

line, 5 from sheath at base of spinous dorsal to arch of lateral

line, 12 thence to base of pelvic fin. Dorsal X 11; fifth spime

longest, a little more than } length of head; last as long as

eighth, 4 length of head. Anal II] 12; base as long as head

or 14 as long as caudal peduncle. Pectoral 3 length of head.

Caudal forked, the middle rays 3 as long as the longest. Caudal

peduncle 13 as long as deep. Silvery ; back darker ; caudal with

dark posterior edge and pale intramarginal band.

Kastern Pacific.

Here described from a specimen of 190 mm. from Tahiti:

this is not so deep in the body as Fowler’s type, a specimen of

about 115 mm. from the Bonin Islands. Otherwise, making

allowance for differences due to size, the two examples agree

perfectly.

7. KUHLIA CAUDOVITTATA.

Holocentrus caudovitiatus Lacep. Hist. Nat. Poiss. iv. pp. 332,

367 (1802).

Dules caudovittatus Cuv. & Val. Hist. Nat. Poiss. iii. p. 117

(1829) and vii. p. 475 (1835); Sauv. Hist. Madag., Poiss. p. 151,

pl. xvii. f. 3 (1891).

- Kuhlia caudovittata (part.) Bouleng. Cat. Fish. i. p. 41 (1895).

Evidently related to K. humilis, sandvicensis, etc., but dis-

tinguished by a small head, the subequal ninth and tenth dorsal

spines, and the increaséd number of soft rays, 13 or 14 in the

dorsal, 12 or 13 in the anal.

Madagascar and Mauritius.

Total length 225 mm.

8. KUHLIA SPLENDENS, sp. n. (Text-fig. 69 B.)

Kuhlia mato (part.) Bouleng. Cat. Fish. i. p. 40 (1895).

Depth of body 24 to 22 in the length, length of head 3 to 33.

Snout 2 to 2 diameter of eye, which is 3 in length of head;

interorbital width about 34. Maxillary extending to below

anterior 1 of eye. 26 or 27 gill-rakers on lower part of anterior

arch. 48 to 50 scales in lateral line, 4 from sheath at base of

spinous dorsal to arch of lateral line, 11 thence to base of pelvic

fin. Dorsal X 11; fourth or fifth spine longest, nearly = length

of head: last as long as or longer than eighth, nearly 3 length of

head. Anal III 11; third spine a little shorter than tenth

dorsal; base a little shorter than head or 12 as long as caudal

peduncle. Pectoral 2 length of head. Caudal deeply emarginate,

but middle rays more than 2 as long as the longest. Caudal

peduncle longer than deep. Silvery; back darker ; caudal with

blackish posterior edge.

Four specimens, 120 to 180 mm. in total length, from

Rodriguez (fresh water) and Mauritius.

380 ' MR. C. TATE REGAN ON

A small stuffed specimen from South Africa also belongs to

this species.

Text-fig. 69.

mo\\

Heads of (A) Kuhlia humilis and (B) Kuhlia splendens, in each case

from a specimen 120 mm. in total length.

9. Kuuuia HuMILIS. (Text-fig. 69 A.)

Kuhlia humilis De Vis, Proc. Linn. Soc. N. 8. Wales, ix.

1884, p. 396; Ogilby, Ann. Queensland Mus. x. 1911, p. 46, pl. vi.

ie dl.

Kuhlia malo (part.) Bouleng. Cat. Fish. i. p. 40 (1895).

Kuhlia proxima Kendall & Goldsborough, Mem. Mus. Comp.

Zoolexxvin | QUES 282 replete ta: 2.

Depth of body 22 to 3 in the length, length of head 3 to 3}.

Snout 3 to 2 diameter of eye, which is 2} to 2 in length of head ;

interorbital width 31 to 3}. Maxillary extending to below

anterior } of eye.. 25 gill-rakers on lower part of anterior arch.

46 to 50 scales in lateral line, 4 from sheath at base of spinous

dorsal to arch of lateral line, 10 or 11 thence to base of pelvic fin.

Dorsal X 11; fourth or fifth spine longest, 2 length of head ;

tenth a little shorter than seventh, more than ? length of

head. Anal JII 11; third spine as long as or a little longer

than second, as long as tenth dorsal; base of fin nearly 3 as long

as head or 1} as long as caudal peduncle. Pectoral 3 length of

head. Caudal deeply emarginate, but middle rays more than 4 as

long as longest. Caudal peduncle 14 as long as deep. Silvery ;

back darker; caudal with a rather broad dark posterior margin.

Queensland ; Fiji Islands.

Three specimens, 90 to 120 mm. in total length, from Fiji, are

undoubtedly A. prozima, but this seems to be a synonym of the

Queensland A. humilis. The species is scarcely distinct from the

preceding, but has a larger eye, the anal fin a little shorter, and

the blackish margin of the caudal fin broader.

FISHES OF THE GENUS KUHLIA, 381

10. KUHLIA SANDVICENSIS.

Moronopsis argenteus, var. sundvicensis Steind. Sitzungsb, Akad.

Wien, xxiv. 1876, p. 205.

Moronopsis sandvicensis Steind. ib. xevi. 1887, p. 56, pl. i. f. 1.

Kuhlia malo (part.) Bouleng. Cat. Fish. i. p. 40 (1895).

?Kuhlia wenura Jord. & Everm. Bull. U.S. Nat. Mus. xivi.

1836, p. 1015.

Depth of body 22 to 24 in the length, length of head 31 to 32.

Snout 2 to 4 diameter of eye, which is 22 to 3 in length of head ;

interorbital width 31 to 4, Maxillary extending to below anterior

edge or anterior 1 of eye. 25 to 28 gill-rakers on lower part of

anterior arch. 48 to 52 scales in lateral line, 5 or 6 from sheath at

base of spinous dorsal to arch of lateral line, 11 to 13 thence to base

of pelvic fin. Dorsal X 11; fifth spine longest, 3 to length of

head; last a little shorter than seventh, + to 4 length of head.

Anal III 11; third spine as long as tenth dorsal; base of fin

nearly as long as head or 12 to 14 as long as caudal peduncle.

Pectoral 2 length of head. Caudal deeply forked, middle rays

less than 4 as long as longest. Caudal peduncle 14 to 1} as long

as deep. Silvery; back darker; caudal with narrow blackish

posterior edge; dorsal and anal sometimes similarly margined.

Eastern Pacific.

Six specimens, 90 to 240 mm. in total length, from the Sand-

wich Islands and Tahiti.

11. KUHLIA MUTABUNDA.

Kuhlia mutabunda Kendall & Radcliffe, Mem. Mus. Comp.

Zool xexmye Ol, p05) poles teal

This species from Easter Island, described above (p. 369,

Pl. LVI), differs from A. sandvicensis especially in the shorter

anal fin.

12. KUHLIA TENIURA.

Dules teniurus Cuv. & Val. Hist. Nat. Poiss. ii. p. 114

(1829).

Kuhlia teniura Bouleng. Cat. Fish. i. p. 39 (1895) (with

synonymy).

Kuhlia sterneckii Steind. Sitzungsb. Akad. Wien, cvii. 1898,

p. 461, pl.

This species is at once distinguished by the coloration of the

caudal fin, with five black bands, a median longitudinal and two

on each lobe. I am unable to regard Steindachner’s A. sternechit

as distinct, for the relative length of the ninth and tenth dorsal

spines varies considerably, and they are equal in 4 or 5 of the

20 specimens I have examined.

The species ranges throughout the Indo-Pacific, from Africa to

the Revillagigedo Islands.

382 MR. R. I. POCOCK ON

29. The Affinities of the Antarectie Wolf (Canis antarcticus).

By Re Pocock, ERS. eS ie ZS.) Curatenol

Mammals.

{Received and Read April 8, 1918.]

(Text-figures 70-74.)

INDEX. Pave

1. Published views on the affinities of C. antarcticus............... 383

2. Skull characters of C. antarcticus and C. latrans ............... 3884

3. Hxternal characters of C. antarcticus and OC. latrans ......... 391

The story of Canis antarcticus has been told by Darwin *, by

Hamilton Smith 7, and more recently by Mr. Rupert Vallentin =

from first-hand information, and by several authors indebted

either to Darwin’s or Hamilton Smith’s account, or to the

accounts of travellers who visited the Falklands before Darwin’s

time. References to the literature down to 1890 may be found

in Mivart’s ‘Monograph of the Canide,’? published in that

year. According to Mr. Vallentin, Canis antarcticus became

extinct in 1876, without leaving a trace of its former existence

in the Falkland Islands; and since all the known material of the

species appears to be preserved in London and Paris, I have

attempted to supply the want expressed by Allen §$ by figuring a

skull of one of the specimens in the British Museum. I have

not, however, given detailed measurements of the skull, because

these may be found in Mivart’s monograph and in the paper by

Huxley mentioned below.

Some six or seven years ago, when trying to identify some

South American dogs exhibited in the Zoological Gardens, I took

the incidental opportunity of looking at the skulls of a few of

the species of Neotropical Canidze contained in the» British

Museum, to learn, if possible, something of their affinities to

one another and to the better known species inhabiting North

America and the countries of the Old World. Amongst the

species examined were Canis antarcticus, the so-called Wolf of

the Falkland Islands, and Canis latrans, the Coyote or Prairie

Wolf, which ranges roughly from Canada to Mexico. The

examination was made without any intention on my part of

adding to the literature of the subject, with which I was only

acquainted in a very general way; and after satisfying myself

that C. antarcticus was related to certain Neotropical forms,

of which C. thous (= cancrivorus) may be taken as an example,

and that the affinities of C. latrans lie with some of the so-called

jackals and wolves of the Old World, I was contented to let the

matter rest.

* In Waterhouse’s Zool. of H.M.S. ‘ Beagle,’ Mammalia, p. 7, 1839.

+ In Jardine’s Nat. Libr., Mammalia, ix. p. 252.

{ Manchester Memoirs, xlviii. p. 45, 1904. This paper is quoted hy

Mr. Lydekker, and some of the interesting and puzzling points connected with

C. antarcticus are discussed in ‘The Field,’ Oct. 1, 1904.

§ Rep. Princeton Univ. Exped. Patagonia, iii. pt. 1, p. 153, 1905.

me THE ANTARCTIC WOLF, 383

But in the summer of 1912, Lreceived for review from the Editor

of ‘ Nature’ a copy of Dr. R. F. Scharft’s volume, ‘ Distribution

and Origin of Life in America,’ 1911; and when I found it

definitely stated therein that C. antarcticus is closely related to

C. latrans, and when I saw the obvious difficulties in which

Dr. Scharff was involved in his attempt to explain, on geo-

graphical grounds, this singular aftilation, I ventured to reassure

him by remarking, in effect, that his belief was devoid of morpho-

logical foundation.

Now, an author who compiles a volume on zoology of the size

and scope of the ‘ Distribution and Origin of Life in America’

cannot be expected to verify all the statements of earlier and con-

temporary writers. Nor in the present instance could Dr. Scharff

be justly criticised for not travelling to London to examine for

himself the preserved material of C. antarcticus, of which, I take

it, there is no specimen in Dublin. Very naturally, therefore,

he trusted to the verdict of others, and promptly replied to my

remark with a request for my reasons for making it. But since

I could not ask the Editor of ‘ Nature’ to give me the necessary

space for justifying the statement I had made, I pledged myself

to do this elsewhere, and the matter that follows is an attempt

to redeem that promise.

The acknowledged source of Dr. Scharfi’s opinion about the

mutual affinities of C. antarcticus and C. latrans was the following

passage in Mr. Lydekker’s ‘ Geographical History of Mammals,’

1896 :—“ Of the two indigenous mammals, the most remarkable

is the Falkland Island Wolf (Canis antarcticus), which differs

markedly from all the Canide of the mainland and is apparently

closely allied to the North American Coyote (C. latrans)” (p. 140).

I therefore wrote and asked Mr. Lydekker if he would kindly

tell me his reasons for this conclusion, and he informed me that

he took it from Prof. Huxley’s classic paper upon the cranial and

dental characters of the Canidz, published in the ‘ Proceedings’ of

this Society, 1880, pp. 238-288. Upon looking up this paper

I find the following passages referring to the two species under

discussion and bearing upon the question at issue :—

(1) ..... But sometimes there is a well-defined though com-

paratively narrow sagittal area, from the centre of which

a low sagittal crest rises. This is well seen in some

Jackals, and especially in C. antarcticus (p. 250).

(2) In the large size of the upper molars..... C’. antarcticus

presents the closest approximation to some specimens of

C. latrans (p. 266).

(3) From the range of variation of C. cancrivorus it can hardly

be doubted that the examination of more extensive

materials will prove the existence of an uninterrupted

series of gradations from C. vetulus to C. antarcticus and

C. jubatus (p. 266).

(4) Seven crania of C. latrans, when measured, exhibit a con-

siderable range of variation, though probably less than

a larger series would show. But, as they are, I must

confess myself unable to find an important break in the

384 MR. R. I. POCOCK ON

series of gradations of cranial and dental structure

between Canis latrans and C. antarcticus on the one

hand, and C. latrans and C. occidentalis on the other.

..... IT may further remark that I can discern no

difference of the slightest importance between skulls of

C. latrans and those of some of our domestic dogs

(pp. 272-273).

(5) In the genus Canis we have .....as a lowest section the

species of the C. cancrivorus and C. vetulus type (an-

swering pretty much to the Aguarra dogs of Hamilton

Smith), the Sacaline section (C. aureus, C. anthus,

C. mesomelas, C. antarcticus, C'. latrans), and the Lupine

section (C. dwpus and all its varieties) (p. 286).

Whether these paragraphs justify Mr. Lydekker’s statement *

that C. antarcticus differs markedly from all the Canidz of the

mainland of South America and is apparently closely allied to

C. latrans, and Dr. Scharff’s extension of this to the effect that

C’. antarcticus is certainly closely related to C. latrans, must be

left to individual judgment.

Paragraph 1 merely points out one resemblance between

C. antarcticus and some jackals. Paragraph 2 similarly points

out one resemblance between the two species, but contains no

suggestion of affinity between them. Paragraph 3 may be

interpreted as suggesting affiliation between the extreme forms

of South American Canide represented by C. vetulus and

C. jubatus, with C. antarcticus lying midway between them.

Paragraph 4 is more precise and states that there is no im-

ortant structural break between C. antarcticus and C. latrans,

and that the latter similarly intergrades with C. occidentalis

and C. familiaris. Paragraph 5, on the contrary, definitely

associates C. antarcticus and C. latrans, and at the same time

severs the former from the group typified by C. vetulus and the

latter from the group typified by C. lupus or occidentalis, an

arrangement not easy to reconcile with the views expressed by

paragraphs 3 and 4,

After reading Prof. Huxley’s paper rather carefully for

enlightenment on this subject, I must confess that I cannot

form any clear idea as to his views of the athnities of the species

he discussed, except in a broad sense.

If the substance of paragraphs 4 and 5 afford some justifi-

cation for Mr. Lydekker’s declaration respecting the relationship

between CO. antarcticus and C. latrans, it must be admitted that

paragraph 3 does not support the contention that C. antarcticus

is quite unrelated to the species of Canide inhabiting the South

American mainland. However that may be, the conclusions

forced upon me by the examination of five crania of C. ant-

arcticus and twelve of C. latranst in the British Museum and

* Tn the article in ‘The Field’ (Oct. 1, 1904), above referred to, Mr. Lydekker

evinces less assurance on these points; but he evidently could not bring himself to

reject the authority of Huxley’s opinion.

+ I use this term in its old-fashioned and broad sense, disregarding the species or

subspecies recently dismembered from C, latrans by American systematists.

THE ANTARCTIC WOLF.

Text-fig. 70,

L By

Z ;

SUNY Aer

AOS

NY)

Bia

)

TMT

A. Dorsal view of posterior part of skull of Canis latrans.

B. Dorsal view of skull of C. antarcticus. ;

0.¢., occipital crest; s.c., sagittal crest; s.a., sagittal area.

The figures of the skull of C. latrans are from a specimen (¢) in the British

Museum from Assiniboia (2.8.22), and those of C. antarcticus trom a specimen in

the British Museum (69,2%24.3). ]

386 MR. R. I. POCOCK ON

the Museum of the College of Surgeons are:—(1) that C. ant-

arcticus and C. latrans are not closely allied; (2) that C. antarcticus

is more nearly related to the C. thous (= cancrivorus) group of

South American Canidee than to C. latrans; (3) that C. latrans

must be affiliated with such Old World species as C. pallipes,

C. lupaster and C. anthus, and not with C. antarcticus. The

first and third of these conclusions are borne out by the

external characters of the two species concerned. My reasons

for these conclusions are as follows :—

The sagittal area and sagittal crest—As Huxley and Mivart

have shown, the skull of C. antarcticus has a well-marked lyriform

sagittal area which, according to the evidence of available crania,

persisted throughout life, although in one of the three specimens

in the British Museum it is decidedly narrower than in the two

others. In the skulls of C. latrans that I have seen there is no

distinct lyriform sagittal area, but in adult skulls there is a

median cariniform sagittal crest varying in height with age.

Even in two young skulls, in both of which the sphenoidal and

occipital sutures are open, while one still retains a milk canine

behind the permanent canine, there is no lyriform sagittal area.

The significance of this depends upon the fact that the young of

many species of Canidee of corresponding age or older show a

stronger or weaker lyriform area corresponding with the sinuosity

of the upspreading temporal muscle on each side, although in the

young of no species of dog in which the adult possesses a carini-

form sagittal crest does the lyriform sagittal area show, I believe,

the development and definition it exhibits in the adult of

C. antarcticus. However that may be, if C. antarcticus and

C. latrans were closely related, we should at least expect to see

a well-defined lyriform sagittal area in the skulls of subadult

individuals of C. latrans killed before the temporal muscles had

reached the summit of the cranium. But, as has been said, this

area is remarkable for its indistinctness in immature skulls of

that species.

The occipital crest.—In C. antarcticus the occipital crest, when

viewed from above, is transversely truncated and not angular;

when viewed from the side it only overhangs the vertical portion

of the supraoccipital to a small extent; and when viewed from

behind it forms a truncated angle. In C. latrans this crest is

angularly produced backwards in the middle line, overhangs the

occipital area to a much greater extent, and is more acutely

angled from behind. It varies in shape and development in this

species, but never, so far as I have seen, resembles that of

C. antarcticus (text-figs, 70 & 71).

The malar bone. Th Canis antarcticus the anterior portion of

the malar bone is marked by a strong masseteric ridge traversing

approximately the middle of its ane surface; the sinless ior edge

of the bone close to the maxilla is expanded convexly to afford

additional support to the masseter muscle ; its upper edge close to

the maxilla is somewhat out-turned, forming a very appreciable

hollow on the subjacent portion of the eee above the first

THE ANTARCTIC WOLF. 387

molar tooth. In C. latrans the masseteric crest of the malar is

low down on its external surface, the inferior edge of the bone is

scarcely at all expanded, so that the area for the attachment of

Text-fig. 71.

BoLa:

A. Occipital region of skull of Canis antarcticus.

B. Occipital region of skull of C. latrans.

S.a., sagittal area; s.c., sagittal crest.

the masseter is much narrower than in (. antarcticus, and the

upper edge of the malar is not noticeably out-turned, so that the

hollow on the maxilla beneath it is less pronounced (text-figs. 72

& 73, pp. 388-9).

Upper carnassial tooth.—In C. antarcticus the antero-external

Proc. Zoot. Soc.—1913, No. XX VII. 27

388 MR. R. I. POCOCK ON

cusp has the front border more rounded and the very fine crest

that runs down it is defined on the inner side by a very indistinct

groove. The antero-internal cusp is wider and rises further hack

and has no distinct little crest running inwards towards the

Wits

tyre

in ee

J My

m., masseteric ridge or malar bone.

Text-fig. 72.

Side view of skull of Canis latrans.

antero-external cusp. In C. latrans the crest traversing the

anterior edge of the antero-external cusp is more pronounced

and is defined by a distinct groove, the two combining to make

the edge of this cusp more cutting than in C. antarcticus. The

THE ANTARCTIC WOLF. 389

antero-internal cusp is narrower and set distinctly more forwards

than in C. antarcticus, and there is a delicate crest running along

its surface towards the base of the antero-external cusp (text-

fig. 74, A, B, p. 390).

Lower carnassial tooth—The main cusp is higher and more

pointed in C. antarcticus than in C’. latrans, and the little cusp at

its base on the inner side is much lower, so that it stands on a

little higher level than the internal cusp of the talon. In C.latrans

m., masseteric ridge or malar bone.

Mexct=tio~ Wa.

Side view of skull of Canis antarcticus.

this cusp is comparatively high up the main cusp of the tooth and

is considerably above the inner cusp of the talon (text-fig. 74,

CDM p9390):

There are other minor differences both in the skull and teeth.

The palatine bones, for instance, extend farther forwards with

relation to the upper carnassials, and the margin of the posterior

nares is also farther forwards with relation to the posterior

molars in C. latrans than in C. antarcticus: the incisor teeth are

21 *

390 MR. R. I. POCOCK ON

smaller and the crowns of the cheek-teeth are higher with relation

to their breadth in C. antarcticus than in C. latrans. But apart

from these, the principal differences mentioned above are quite

sufficient to disprove the claim that the two species are closely

related. According to modern standards of classification they are

subgenerically, if not generically, distinct.

But the characters above described tell us more than that.

Taking C. latrans first, it is obvious that in the cariniform sagittal

crest, the angularly produced occipital crest, the position of the

masseteric ridge on the malar bone, and in the points alluded to

in connection with the upper and lower carnassials, the species

falls into line with the large wolves like C. occidentalis and lupus,

Text-fig. 74.

A. Vertical view of upper carnassial of Canis latrans.

B. Vertical view of upper carnassial of C. antarcticus.

C. Internal view of Jower carnassial of C. latrans.

TI). Internal view of lower carnassial of C. antarcticus.

and with (. pallipes and C. lupaster, which, according to fancy,

may be called large jackals or small wolves. These resemblances

explain Mivart’s dismissal of the cranial and dental characters of

C. latrans with the remark, ‘‘The skull possesses no distinctive

characters, nor have we been able to detect any in the shape of

the teeth.”

On the other hand, the skull of C. antarcticus, with its lyriform

sagittal area and truncated occipital crest, agrees in the main with

the skulls of certain species or subspecies of South-American dogs

in the British Museum labelled C. thous (=cancrivorus), rudis,

sclatert (=microtis), parvidens, urostictus, gracilis, and fulvipes.

And in the skull of a dog, perhaps referable to C. gracilis, which

THE ANTARCTIC WOLF. 391

eame from Mar del Plata and died in the Gardens, the above-

described crests on the upper carnassial are not better developed

than in C. antarcticus, and the masseteric ridge on the malar bone

shows a decided approximation to the condition seen in that

species. This latter character is still better marked in the skull

of another South-American dog, the exact locality of which is

unknown, but which was a different species * from the Mar del

Plata example, and the crests on the carnassial exhibit the same

feebleness of development. But it may be noted that in both

these skulls the positions of the cusps on the upper and lower

carnassials are more latrans-like than antarcticus-like, so that in

this respect at least they serve to bridge over the difference

between those two species; a fact in keeping with the idea that

C. antarcticus is a specialised form of the group of South-

American dogs above alluded to, but specialised in a direction

away from that taken by C. latrans and its allies.

The external characters of C. antarcticus and CO. latrans also

afford no justification for the claim of close relationship between

them. In the first place the ears of C. antarcticus are very small,

smaller indeed comparatively, I believe, than in any wild species

. the dog family, with the exception perhaps of C. sclatera,

C. (Nyctereutes) procyonoides, and Vulpes (Alopex) lagopus. In

C. latrans, on the contrary, they are as large as in most, at all

events, of the species of Canis. An idea of their length in the

two species may be gathered from the measurement of a specimen

of each of approximately the same size given by Mivart, the ear

of OC. antarcticus being 6°5 cm. (= about 24 inches) and that of

C. latrans 14 em. (=about 54 inches), or more than twice as long.

It may be added that the measurement of 2 inches 9 lines assigned

by Waterhouse to the ear of (. antarcticus confirms Mivart’s

statement.

As regards colour ©. latrans varies from grey to greyish fawn,

mixed with black above, and shows the characteristic clouded or

patchy coloration caused by the running together of the bands of

the individual long coarse hairs of the ‘back and sides seen in so

many of the so- called wolves and jackals. One of these long

coarse hairs, pulled at random from a skin, measured about

3 inches long, the black tip being 3 inch (12 mm.) and the

whitish area below it 1 inch (25 mm.). The whole of the ventral

surface from the chin to the root of the tail is usually white or

whitish, and always apparently markedly paler than the back and

sides, though sometimes the continuity of the light tint is inter-

rupted on the throat by an infusion of fawn. There is no dark

patch above the hock on the hind leg, and the tail matches the

back approximately in colour throughout, the tip and the gland-

spot being blacker than the rest.

In C. antarcticus the coat is thick and soft, and comparatively

short, with none of the long coarse hair seen in C. latrans. One

* The South-American dogs of this groun are in such 2 systematic muddle that

it is very difficult to identify specimens without a complete revision of the whole

series,

392 ON THE ANTARCTIC WOLF.

of the longish hairs pulled at random from the back measured

only 14 inches (37 mm.) in length, the dale apical tip being

1 inch (6 mm.) and the pale ‘eral below it + inch (3 mm.) in

length. The prevailing colour of the body is brown, relieved by

the. fine speckling due to the narrow pale band on the individual

hairs just described. The lower side is white only on the

posterior portion of the belly and on the upper end of the throat,

the-chin and lower jaw being white stained with a fuscous tint.

Apart from the areas described the ventral surface is brownish.

There is, moreover, as Mivart said, a fuscous patch above the

hock of the hind leg and the tail is particoloured, its basal

portion being like the back, its tip white, and the intermediate

area blackish, the colour of this area gradually blending proximally

with the brownish basal portion, but- being quite black distally

and sharply defined from the white ter minal area,

Perhaps it may rightly be claimed that these differences in

the length of the ears and in colour do not count for much in

themselves. That may be so. Nevertheless, if C. antarcticus and

C. latrans were only known from their skins, it is quite certain

that the latter would be placed in the same category with such

species as C. pallipes and C. lupaster, and that C. antarecticus

would be excluded therefrom. The latter would be difficult to

classify; but there is one significant colour-feature connected

with the species. ‘This is the presence of the dark patch above

the hocks; and the interest of this lies in the circumstance that

it is a very common feature in various species of the smaller

South-American dogs and occurs in some of the species of Vulpes,

hike V. chama*.

There is one other little point that may be referred to.

Darwin says he was informed that the cries of C. antarcticus

resembled those of the South-American species C. azare. I have

never heard C. azare bark or howl, but the keeper in the Gardens

informs me that examples of wild dogs from Mar del Plata and

Cordova, which are closely allied to and perhaps only racially

distinct from C. azare, bark after the manner of foxes. On the

whole, however, they are silent dogs in captiv ity, and, like the

foxes, never succumb to the temptation of joiming in the howling

concerts in which the dingos, jackals, prairie wolves, and laree

wolves in the Gardens indulge. and lhe they seem unable to

resist contributing to. Personally I believe that voice in mammals

is often a good guide to affinity; and, in the present case, the

voices of C. antarcticus and C. latrans bear out wy opinion of

the relationship of these species to others, shown by structural

characters.

Finally, if the conclusions above put forward are correct,

Huxley’s classification, expressed in par. 5 (p. 384), must be

emended by transferring C. antarcticus to the lowest section of

* The presence of this patch in some of the primitive Canide is well worth more

attention than it has received. I do not know what it may mean, any more than I

know what the pale area behind the shoulder, observable in many Canidx, both wild

and domesticated, may mean.

“WCIOVEODOUOVIVHA AO WTTIHLVd

‘onoyud “Sg mM a

UXT “iel Cll “S “A al

ON AVIAN OSTEOLOGY. 393

dogs, containing C. thous (=canerivorus) and vetulus, and by

uniting his Sacaline and Lupine sections. Possibly these sections

contain groups worthy of subgeneric, if not of generic, recog-

nition: C. antarcticus and C. thous, for example, may be sub-

generically or generically separated by the structure of the

mandible; but I do not see how C. latrans is to be distinguished

other than specifically from such forms as C. pallipes, anthus, or

even lupus.

30. On the Patella in the Phalacrocoracide.

By Dr. R. W. Saurezpr, C.M.Z.S.

[Received April 14, 1913: Read May 20, 1913.]

(Plate LXL*)

While recently employed in preparing a detailed account of the

skeleton of Harris’s flightless Cormorant (Vannopterum harrisi),

in which the osteology of that species is compared with that of

a number of others of the family, I became interested in the

morphology of the patellz of those birds.

Many years ago J published a number of papers on the skeleton

in the Cormorant, in some of which the patella of the Phala-

crocoracide was referred to and figures given of it. But my

material, at the time to which I refer, was very limited— in fact

J think there were but the skeletons of one or two species of

those birds at hand, and this included the skeletons of two or

three young ones.

In the higher groups of birds the patella, when present, is

usually small, and offers but little of value to the avian taxonomist.

This, however, is not the case when we come to examine into the

osteology of some of the groups occupying lower positions in the

system, and especially is this true of many of the Pygopodine

forms and their allies near and remote.

Owen paid but scant attention to this bone of the skeleton in

Aves, devoting less than half a paragraph to it, thus :—‘‘ The

chief of the sesamoid bones in the hind limb is the patella: it is

of unusual size in the Penguin, is ossified from two centres, and

articulates with the procnemial process of the tibia: it coexists

with the long rotular process in the Loon, fig. 34,7; it is large

and of an angular form in the Musk-duck (Biziwra): in the

Merganser the patella is largest and deeply notched; in the Coot

it iselongate. In most aer al birds a patella is wanting je hhere

is no reference made here either to a Grebe or a Cormorant! and

* For explanation of the Plate see p. 402.

+ Shufeldt, R. W. “Osteology of the Cormorant,” Science, Dec. 7, 1883, p. 739 ;

Feb. 8, 1884. vol. iii. No. 53, p. 143; ibid. Apr. 18, 1884, No. 63, pp. 474, 475.

oe Gonter ning some of the forms assumed hy the Patella in Birds, » Proc. U.S. Nat.

Mus. 1884, vii. pp. 324-331. Numerous text-figures.

Osteology of the Steganopodes,” Mem. Camegie Museum, Pittsburgh, Pa., Apr.

1903, vol. 1. No. 3, Art. 3, pp. 15-70. Plates and many text-figures.

t Owen, Richard. Comp. Anat. and Phys. of Vertebrates, vol. ii. p. 88, London

1866.

394 DR. R. W. SHUFELDT ON

Professor Owen evidently considered the smal, scale-like bone

found in the tendon of the extensor femoris, at its insertion,

to be the patella in that bird. When I first examined this

question, this was likewise my opinion, and in a paper on the

patellee in birds, published in 1884 in the ‘ Proceedings of the

United States National Museum,’ I so figured it (fig. A, p. 328)

for Colymbus septentrionalis (Gavia stellata). This opinion was to

some extent qualified later on, when I stated, with respect to the

Loons (“ Urinatoride”) that they possess ‘‘ only a very small,

flake-like sesamoid, which occurs in the tendon of the extensor

femoris muscle at its insertion, and probably the true patella has

coossified in the adult with the elongated cnemial process of the

tibio-tarsus”*. Possibly some avian osteologist has published on

this subject, but if so, | have not seen the work; and never

having been so fortunate myself as to have come into possession

of the skeleton of any Loon, secured at the right time to demon-

strate the exact composition of the tibio-tarsus in that bird, I am

still in doubt on the question. However, it is quite possible—

indeed quite likely—that the true patella in Loons (Gavia) is, in

the adult, completely codssified with the great elongate cnemial

process of their tibio-tarsi. The moulding of the patella on the

back of this process in Grebes, especially in very old birds, is

wonderfully close,—so close in some ligamentous preparations as

to deceive the eye upon casual examination.

To settle this interesting poimt—if it has not as yet been

settled—will require the examinations of the skeletons of Loons

including those of individuals of the genus of all ages.

At present [ am inclined to think that the patella in Gavia,

in the adult, has been indistinguishably fused with the cnemial

process of alg tibio-tarsus, for the reason that it is on the road to

such a fate in the Grebes ( tolymbidee), and that in all such birds

as Hesperornis—an ancient ancestor of the Loons—the Penguins,

the Cormorants, and some others, the patella is very large.

Then, finding it large in a Gr abe, one would naturally look for

the same in eh forms as Loons, especially when one considers

the relationship of these two families.

The statements now being made are, in a way, prefatory, leading

up to what I have to say on the patella of the Cormorants. It

must be borne in mind in this connection that the patella in

Penguins is very large, and grooved obliquely across the anterior

face for the tendon of the ambiens muscle 7.

* “Concerning the Taxonomy of the North American Pygopodes based upon their

Osteology,” Jour. Anat. & Phys. London, Jan. 1892, pp. 199-208. The lines quoted

are trom page 202. In this paper I also give two figures of the skeleton of the thigh

and leg of a Grebe, in which the patella is included. One of these figures was repro-

duced by Coues in his fifth edition of the ‘ Key,’ without acknowledgment (vol. 11,

p. 1052, fig. 712).

+ Coues, E. “Material for a Monograph of the Spheniscide,” Proc. Acad. Nat,

Sci. Phila. xxiv. 1872.

Watson, Morrison. “Anatomy of the Spheniscidie” (Rep. Scient. Results of

Vovage of H.M.S. ‘ Challenger,’ Zoology, vol. vii. pl. vii. figs. 9 & 10, 1883).

Shuteldt, R. W. Proc. U.S. Nat. Mus. 1884, p. 326, fig. 1. Reproduces from Watson

figures of patellze of Aptenodytes pennantii and Ludyptes chrysocome. Comments

AVIAN OSTEOLOGY. 395

Personally I have not examined the patella in Hesperornis with

the view of substantiating Marsh’s statement that it is ‘‘ perforated

by a large foramen for the tendon of the ambiens muscle, agreeing

in this respect with the patella of the Gannet (Sula bassana

Briss.).” His several figures of the patella of Hesperornis regalis

do not show this “large foramen” (‘Odontornithes,’ p. 23,

plates xv. and xx. figs. 1-3), while I have yet to see the patella

of Sula bassana presenting any such perforation for the ambiens

muscle as Marsh describes.

However this may be, the fact remains that very radical dif-

ferences exist with respect to the fate and disposition of the

patella and cnemial process of the tibio-tarsus in Grebes, Loons,

Hesperornis, Penguins, Gannets, Cormorants, and many other

different species of water birds. Great differences also exist in

the morphology of this sesamoid and the aforesaid apophysis.

This brings us to a point where the patella of the Phalacro-

coracidee may be discussed. For this purpose I have before me

the skeletons of several species of Cormorants, all belonging to

the Collection of the U.S. National Museum, to which institution

I am indebted for the loan of them. It will not be necessary to

name these species here or to give their museum numbers, as this

information is fully set forth in the explanation of Plate LXI.,

which appears at the end of this article. As to the 19 figures on

the plate, they are reproduced, natural size, from my own grouping

and photograph,—the latter having been made direct from the

specimens.

Among avian osteologists the general opinion has prevailed that

the patella in Cormorants is a big trihedral one, in some way

“perforated” for the ambiens muscle.

Garrod found the ambiens muscle present in the family Phala-

crocoracide ; and, in noting that Meckel had not done so, he states

that “it is peculiar in that [in the Cormorant] it runs through

the substance of the large triangular patella, in a bony canal” *.

In another paper this gifted ornithotomist is still more positive,

and in speaking of the “‘ Steganopods,” he says: ‘“ The ambiens is

of fair size; it deeply grooves the large ossified patella ; and some

of the fibrous ligament overlapping this groove shows traces of

ossification ; so that in aged birds the groove may be converted

into a foramen, as is always the case in Phalacrocorax, where the

made thereon. Thereisacurious slip in this paper with respect to the description of

the patella of the Gannet (Sula bassana). On page 362 I state that “ Professor Marsh

tells us that the patella of Sula is pertorated by a large foramen for the passage of

the tendon of the ambiens muscle.” On pages 329 and 330 I quote the entire para-

graph from Marsh’s ‘ Odontornithes’ (p. 93) where he gives this description of the

patella of Sula and states on page 327 that “I fail to find any such foramen in the

patella of the specimen of Suda before me, although it has a shallow, oblique groove

across its anterior face that seems to correspond with the one described when speaking

of the patella of the Penguin.” This statement is supported by a correct figure of

the patella in Sula bassana (No. 16643, Coll. U.S. Nat. Mus.), and I must believe

that the description left us by Prof. Marsh is another of those curious slips of his m

the osteology of modern birds.

* Garrod, A. H. “On Certain Muscles of the Thigh of Birds, and on their value

in Classification,” Pt. I, P. Z, 8. 1873, pp. 636, 637 (footnote).

396 DR. R. W. SHUFELDT ON

thus formed foramen is far from superficial” *. This is a very

interesting statement, and will be commented upon further on in

this paper.

Forbes, in describing the anatomy of Biziwra, makes the state-

ment that “The ambiens muscle is large, and peculiar in that its

tendon perforates the large-sized triangular patella, just as it does

in Phalacrocorax and the extinct Hesperornis” y.

When referring to this sesamoid in the Cormorants, Coues says,

in the fifth edition of his ¢ Key’ (p. 961), that “ There is a bulky,

free patella, coexistent with a short cnemial apophysis or rotular

process of tibia, but perfectly distinct therefrom, as in Grebes.”

This somewhat ambiguous description is made clearer by an illus-

tration of the patella in ‘“‘ Phalacrocorax bicristatus,’ which he

borrowed from a previous paper of mine. Nothing is said with

regard to the ambiens or the “ perforation ” in the patella, so this

note is quite useless for present purposes.

From my own various accounts of the patella in Cormorants

I select the following as best suited to throw light upon the

facts I propose to here set forth.

In my above-cited article on the patella in birds (Proe. U.S.

Nat. Mus. 1884, p. 330, fig. 7) IT remark: “In No. 41 of ‘Science,’

I presented a lateral view of the leg-bones of a Cormorant (Phala-

crocorax), Showing the form of the patella in these birds. The

Same specimen is given here, only an anterior view is chosen

instead of the lateral one.

‘“This form is a particularly interesting one, and I am not

aware of any bird at present that can show a similar condition of

the parts in question.

‘On the face of it, it almost looks as if a patella had developed

of a size equal to the rotular process, and subsequently the two

became thoroughly united, and formed one large patella, articu-

lating as shown in the drawings I have made of it. Ido not say

that this is actually the case, nor were the young of this spe-

cimen, which also belong to the collections at the Smithsonian

Institution, of a proper age to determine exactly the manner in

which this great bulky patella was developed.”

Apart from the suspicion expressed in this last paragraph,

which will be put into more exact terms further on, there are but

two opinions extant, at present,in regard to the development of

the patella in the Phalacrocoracide. They are the opinions of

Garrod and of Forbes, and have been set forth in their own words

above.

There is but one construction that can be placed upon Forbes’s

view of a Cormorant’s patella, and that view cannot be success-

fully defended. It will be noted above that, in his calling

* Garrod, A. H. “Notes on the Anatomy of Plotus anhinaa,” P. Z.S. 1876,

p. 340, pl. xxviii. figs. 5, 6. The figures give the patella, indicating the position of

the foramen.

+ Forbes, Wm. Alex. “ Note on some points in the Anatomy of an Australian

Duck (Biziura lobata),” P. Z. S. 1882, p. 457.

I regret to say that I have never had the opportunity of examining the patella in

this duck, and I doubt if we have a skeleton of it in the United States.

AVIAN OSTEOLUGY. 397

attention to the fact that the tendon of the ambiens muscle

passes through the patella in Cormorants, he uses the word

“* nerforates ” "(emphasizing it by the use of italics). Surely

could not have meant that the tendon really did “ perforate ”

bore its way through the patella as the Cormorant grew sail

developed? And, after finding its way through the “bone (or

cartilage) that it duly made attachment at its point of insertion ¢

No such thing has ever occurred in anatomical development, and

Forbes was altogether too good an anatomist to have set up any

such ridiculous explanation. This being true, and disregarding

entirely any such theory or such another possible suggestion as

that, in subadult life, the patella of a Cormorant, forming first

in elementary cartilage, does, when in that condition, surround

the tendon of the ambiens muscle, and afterwards, when the

patella has completely ossified in old birds, the aforesaid tendon

runs through a foramen, which passes through the middle of

it. Barring this, too, as I say, as being a totally untenable sug-

gestion, we are left with but two other possible ways by means of

which the ambiens comes to pass through the foraminal canal in

the patella of Cormorants.

Tn discussing these, we are at once confronted with Garrod’s

opinion in regard to it—an opinion which had never, apparently,

occurred to Forbes. Garrod took the view that, inasmuch as

in anhinga (Plotus) the anterior face of the patella is somewhat

deeply grooved, and as ligament stretches across that groove to

confine ‘the tendon of the ambiens muscle which passes through

it, and as this same ligament sometimes ‘“‘ shows traces of ossi-

fication,’—hence in Cormorants this also takes place, and the

fibrous ligament in them becomes, in old individuals, thoroughly

ossified, and we thus find the tendon of the ambiens passing

through the bone.

Such an opinion will not hold for an instant in the face of the

necessary material to examine into it.

Now in some Cormorants the patella is very large and thick,

and the aforesaid foraminal passage, passing through it from side

to side, is found but slightly in advance of the middle and above

the centre of the bone; so that, had it been a “groove” in the

young bird spanned by fibrous ligament, and this ligament sub-

sequently ossified in the adult, that groove must have been a very

deep one, and the patella in old birds would, through its form

alone, after ossification was complete, exhibit the manner as to

how it had come about. Moreover, in connection with this it is

very clear that, were the foramen in the patella in Cormorants

formed by a groove in front of it being covered over by fibrous

ligament in the subadult bird, which ligament later in life ossified,

that ossification would be smooth on its anterior face, which is by

no means the case, as one may see by a study of the figures in the

plate accompanying this paper. For instance, a fibrous ligament,

stretching across a deep, narrow groove as a retaining band for

the tendon of a muscle, would not, in ossifying, take on any other

shape beyond that possessed by the flat ligamentous band; for

398 DR. R. W. SHUFELDT ON

instance, as I say, such as has occurred in the patella shown in:

Pl. LXI. fig. 11 and others. It must stand to reason, then, that

Garrod entertained an untenable opinion in this matter.

Coming to the second way in which ‘this foramen could have

been formed, it is clear that, in young and subadult individuals,

the tendon of the ambiens muscle could be confined by an

additional piece of bone or cartilage being placed in front of it,

and this added part subsequently ossify and not only form, in

the adult, a foraminal passage for the aforesaid muscle or its

tendon, but largely add to the bulk of the patella. It might, too,

—from the thoroughness of the coéssification—cause the atrophy

of the muscle and its tendon at that part of the course of the

latter across the front of the true patella; and such a result would

be brought about by the entire sealing up of the foraminal

passage, which not only actually takes place in the patelle of

some Cormorants, but again proves Garrod to have been wrong

when he stated that such a foramen or foraminal passage was

always present in the patella of Cormorants. See Pl. LXI.

fig. 3—-Phalacrocorax penicillatus—where it has been sealed up

entirely, leaving not so much as a trace or a suspicion of its ever

having been present there.

In my opinion, the foraminal passage for the ambiens muscle,

passing transversely through the patella in the Phalacrocoracide,

when present, 1s formed as set forth below, which formation can

be demonstrated by the material figured in the plate accompanying

this paper. When the foraminal passage is entirely absent it

has been absorbed, atrophy of the ambiens probably having

ensued.

It would appear that in young Cormorants the ambiens passes

in a groove, of the required depth only, obliquely across the

anterior face of the patella, which latter is very closely adpressed

against the posterior surface of the long enemial process of the

tibio-tarsus,—a condition that persists throughout life in Grebes

(Colymbide). Codssification between the patella and the upper

two-thirds of the cnemial process of the tibio-tarsus, in the case

of the growing Cormorant, next sets in, which, owing to the

morphology of the parts involved, would, in time, depreciate

the action of the knee-joint. This is clear when we come to

consider the origins and insertions of the various muscles about

the knee-joint anteriorly, posteriorly, and laterally *. These,

during their continued action in locomotion—especially during

the act of swimming—tend to overcome the aforesaid danger,

militating against the complete freedom of action of the knee-

joint. Codssification steadily proceeds ; the individual grows ; a

constant tugging is exerted during locomotion at the tibio-tarsal

enemial apophysis. This eventually results in its becoming

dissociated from the bone to which it belongs along a transverse

line, at a short distance above the level summit of the tibio-

tarsal shaft, thus leaving the lower third of the cnemial

* Shufeldt, R. W. ‘The Myology of the Raven,’ p. 187, figs. 51-53, London

1890.

AVIAN OSTEOLOGY. 399

‘process where it occurs in all birds when such an apophysis is

present *.

In the Phalacrocoracide, then, the patella is a compound bone,

consisting of the true patella posteriorly, and the upper two-

thirds, more or less, of the cnemial process of the tibio-tarsus

anteriorly, which latter has become dissociated and thoroughly

codssified with the former. By this union, a foraminal passage

is formed for the ambiens muscle, and this, in some species, may

gradually, but entirely, disappear, whereupon, in time, the muscle

meets its usual fate.

For very obvious reasons, the patella is very large in Harris’s

flightless Cormorant (Vannopterum harrisi); indeed, in so far as

I have been enabled to discover, itis larger in this species than in

any other existing form of the entire family (Pl. LXI. figs. 1 & 2).

It has an average height of 25 millimetres, the longest diameter

of its base being 19 or 20 millimetres, and is nearly a square in

outline. Posteriorly, it is flat and triangular, the acute angle

being above. Above its middle, and nearer its posterior surface

than its anterior margin, it is pierced, transversely, by the foramen

for the ambiens, which is of considerable size. Externally, its

exit is almost flush with the surface of the bone, while on the

inner aspect it is situated at the base of an extensive concavity.

Anteriorly, this compound bone exhibits a uniform longitudinal

excavation which, when the patella is articulated as in life, is the

continuation with the similar excavation between the pro- and

ectocnemial processes of the tibio-tarsus. On the anterior face

of this patella, at the lower-internal angle, there is a transverse,

triangular facet which is intended for articulation with the

superior border of the cnemial crest of the tibio-tarsus. In fig. 2

we have the mesial or inner aspect of the left patella of

NV. harrisi, from the same skeleton (No. 19719), where the

perforating foramen for the ambiens is seen at the base of the

aforesaid extensive concavity. A study of these two patelle

reveals the nature of the composition of the bone as a whole, as

described above.

Phalacrocorax penicillatus possesses only a medium - sized

patella, which is elongate and wedge-shaped. It has the general

form of the patella in WVannopterum: but the transverse fora-

minal passage for the ambiens has been entirely absorbed,—not

a vestige of it remaining in an individual of the age to which

this patella belonged. The union of the two parts composing the

bone is very complete, and all traces of their origin have been

entirely obliterated,—that is, beyond the anterior rotular channel,

and the here extensive facet below it for articulation with the

cnemial process of the tibio-tarsus.

This is a most excellent example of the patella in a Cormorant

* This is the part referred to by Coues in his ‘Key’ (5th ed. vol. ii. p. 961),

where he describes the patella in the Phalacrocoracide in the following words:

“There is a bulky free patella, coexistent with a short cnemial apophysis or rotular

process of tibia, but perfectly distinct therefrom, as in Grebes.” It is very evident

from this definition that Coues had never given the patella of a Cormorant any very

close study,

AOO DR. R. W. SHUFELDT ON

showing the conditions described as they occur in a very old

bird (figs. 3 & 4), and they are equally well shown in the patella

of P. punctatus (figs. 5 & 6), where the foraminal passage has

likewise entirely disappeared. The rotular channel in front is

very narrow and pointed above, as in a Grebe or Loon, and there

is a rounded notch on the superior border of the cnemial process

of the tibio-tarsus, which marks the limitation, externally, of the

facet for articulation with the patella on that border (fig. 6,

where this “notch” is plainly seen). This patella gives barely

any hint as to the two parts of which it is composed.

This is to a less degree true of the patella of Phalacrocorax

magellanicus (figs. 7 & 8); for here again we find the foraminal

passage in its very last stages of ultimate absorption, while the

continuation of the rotularl channel on the anterior face of the

patella, and the evidence of this part of the patella once having

belonged to the tibio-tarsus, is very complete (fig. 7, viiol

should be compared with fig. 6, the former in no way recalling

the form of these bones in the Grebe).

Phalacrocorax urile (Nos. 19655 & 18982) has, in old individuals,

a rather bulky patella (figs. 9, 10, & 11), in which, in the

specimens selected, the foraminal passage is reduced to capillary

proportions, and the sutural traces of the elements composing

it have become nearly obliterated. Fig. 10, which is from the

right limb of P. wrile, No. 18982 of ale Wollleetion of the U.S.

National Museum, shows the minute entrance to the foraminal

assage; while in fig. 11, it being the left limb from the same

skeleton, the opposite opening is in view,— posterior to two

other foramina which occur here. Fig. 11 is given on the plate,

to the exclusion of the patella of Phalacrocor ax carbo, for the

reason that in no Cormorant, other than P. wrile, do we find,

on this anterior view, a better example of the interenemial

channel on both the big sesamoid and the tibio-tarsus, and the

line between them where they eventually parted company.

Phalacrocorax carbo (No. 18850, Coll. U.S. Nat. Mus., not

figured) has a patella that is an interesting one in several

particulars. On its imner aspect the surface is flat and smooth,

the minute opening of the almost entirely absorbed foraminal

canal being situated far posteriorly upon it. In size and general

form the patella of this Cormorant is much like the one shown

in figs. 18 & 19 of P. pelagicus; while it likewise possesses

characters peculiarly its own. It has the usual trihedral form

and makes rather more than the average articulation with

the tibio-tarsus in extent, especially on the proximal margin of

the procnemial process. Its posterior face is triangular and flat,

while the external one is pierced near its middle by the outer

opening of the foraminal passage. Anteriorly, the usual inter-

enemial channel is present, it being continuous with the same on

the tibio-tarsus. On the patella, it is unusually narrow, and

faces very much to the outer side.

Phalacrocorax carbo (No. 18851) has the patella almost

exactly like that in P. uwrile (figs. 10 & 11.) The former,

AVIAN OSTEOLOGY. 40]

however, in this particular individual, has completely lost the

foraminal passage, while otherwise the morphology is quite the

same.

One of the most interesting forms of the Patella among

Cormorants is found in Phalacrocoraa vigua (figs. 12 & 13); for

it is not only very small and chunky, but the facet on its

infero-anterior aspect for the tibio-tarsus is relatively, as well as

actually, large. On the inner aspect, the big foraminal opening

is situated at the base of a fossa, as in No ‘annopterum harris,

while on the external side it is sical again, and has, leading

down from it to the lower border of the bone, a deep groove,

which not only indicates the suture between the two elements

composing this bone, but apparently the continuation of the

groove for the ambiens as well.

Passing to Phalacrocorax auritus (No. 19262), the patella of

which species is shown upon two views in figs. 14 & 15, we find

that that sesamoid is rather small for a Cormorant of its size.

In it, the foraminal passage has gone entirely, while the bone,

upon the other hand, shows better than any other Cormorant’s

patella [ have examined, the two parts of which it is composed.

These are very plainly to be seen in the figures,—indeed, equally

as well as in the specimens themselves.

There is a large patella in Phalacrocorax albiventris (No. 18487,

figs. 16 & 17), where again we find, in the adult bird, ‘ihe

foraminal passage almost obliterated, having been reduced to a

capillary calibre throughout. Sutural traces between the bone

are faint, while anteriorly the rotular excavation is broad and

deep, being but slightly wider at the top than it is at the distal

border.

A most instructive patella is found in Phalacrocorax pelagicus

(No. 19032, figs. 18 & 19). Once more we find the foraminal

passage reduced to a very small calibre, though the entrances on

the inner and outer surfaces of the bone are still in evidence,

though very small. What is to be specially noted among other

things is, however, the position of the foraminal aperture on

the inner aspect (fig. 18); this is but 4 millimetres from the

posterior margin of the patella, while it is 9 millimetres from

the anterior margin, which is sufficient to render the theory of

the “anterior” groove being filled in by an ossified fibrous liga-

ment—oridieulous!

In its amalgamation and coossification of its parts, this patella

became to a large degree twisted,—-a twisting that cannot well

be appreciated in the ficures. Still, the bone fits most poeetcly

on the superior border of the cnemial process of the tibio-tarsus

its lower border, antero-posteriorly, being no thicker than is that

of the leg- pone, while everything in “its morphology plainly

indicates its origin. On the front of the patella, the rotular

channel is soune 0 shallower than usual, and is as broad above

as it is below, where it, in all particulars, directly continues the

corresponding channel separating the pro- and _ ectocnemial

apophyses of the tibio-tarsus.

402 ON AVIAN OSTEOLOGY.

The obliquity of the foraminal passage for the ambiens is

easily made out, its higher aperture being on the inner aspect of

the bone, and situated, as I have pointed out above, far back

from the antero-internal border of the bone. To best appreciate

the fact that this sesamoid in P. pelagicus is formed as I have

demonstrated it to be in Cormorants of all species, it should be

viewed from above rather than from either side or in front.

EXPLANATION OF PLATE LXI.

[All the figures are reproductions of photographs made by the author direct from

the specimens. Each bone is of natural size, and from an adult individual. They

are from skeletons in the Collections of the U.S. National Museum, the Museum

numbers of which are given under the figures. |

Fig. 1. Right patella of Nannopterum harrisi; antero-lateral aspect. (No.

19719.)

. Left patella of NV. harrisi; mesial or inner aspect. (No. 19719.)

. Right patella of Phalacrocorax penicillatus, antero-lateral aspect, or the

same view in which fig. 1 is shown. (No. 18535.) Note that the

foraminal passage for the ambiens has been entirely absorbed.

. Left patella of P. penicillatus ; inner aspect. (No. 18535.)

. Left patella of Phalacrocorax punctatus; inner aspect. (No. 18282.)

. Left tibio-tarsus of P. punctatus ; anterior view, with the patella articulated

in situ. (No. 18282.)

. Right tibio-tarsus, anterior view, of Phalacrocorax magellanicus, with

patella articulated in situ. (No. 18438.)

. Left patella of P. magellanicus; inner surface. (No. 18438.) Note that

the exit of the foraminal passage points to the fact that it is im the

very last stages of its ultimate disappearance. This is an interesting

link in the chain of evidence on the real composition of this bone in the

Phalacrocoracide.

9. Proximal extremity of tibio-tarsus and fibula of right pelvic limb of

Phalacrocorax urile; adult. (No. 19655.) Outer aspect, with patella

and femur articulated in sitz.

10. Inner aspect of the bones composing the knee-joint of P. wrile; partly

ligamentous. (No. 18982.) Right pelvic limb.

11. Left patella, tibio-tarsus and fibula (superior moieties), anterior aspect ;

left pelvic limb of P. wrile. (No. 18982.) Note the complete union

and the continuation of the rotular channel of the tibio-tarsus on the

anterior face of the patella.

12. Right patella of Phalacrocorax vigua; outer surface. (No. 18479.)

Note groove leading down from the foraminal opening.

13. Left tibio-tarsus of P. vigua (No. 18479), with patella articulating

im situ.

14. Bones of the knee-joint of Phalacrocorax auritus (“P. dilophus,”

No. 19262.) Right pelvic limb, inner aspect. Femur somewhat

displaced, but patella properly articulated.

15. Bones of the knee-joint of P. awritus (No. 19262) ; left pelvic limb, outer

aspect and articulated in situ. Note the line of demarcation between

the true patella and the anterior part of the bone, which originally

belonged to the tibio-tarsus.

16. Anterior aspect of the proximal extremities of the left tibio-tarsus and

fibula of Phalacrocorax albiventris, with the patella duly articulated

above them. (No. 18437.) The rotular channel is very broad in this

species and rounded above.

17. Right patella, inner surface of P. albiventris. (No. 18437.) The foramen

is almost closed up.

18. Bones entering the knee-joint of Phalacrocorax pelagicus, adult, nat. size.

Right pelvic limb, outer aspect. Ligamentous preparation with femur

slightly luxated. Patella in situ. (No. 19032.)

19. Skeleton of the knee-joint of P. pelagicus. (No. 19082.) Left limb,

outer aspect; patella i situ.

do bo

co sn DO

ON THE METAMORPHOSIS OF THE AXOLOTL. 403

31. Experiments on the Metamorphosis of the Mexican

Axolotl (Amblystoma tigrinum), conducted in the

Society’s Gardens. By E. G. BouLEncsEr, F.Z.S.,

Curator of Reptiles.

| Received and Read May 20,1913. ]

(Text-figures 75 & 76.)

During the past year I have been experimenting on points relat-

ing to the metamorphosis of the Mexican Axolotl (Amblystoma

tigrinum), and have succeeded in obtaining the transformation of

a number of specimens. ‘This paper deals with the methods

employed, and gives a detailed account of the external changes

undergone by the animal during the process, a subject which,

with the exception of Duméril’s (1) brief account, has not, so far

as I am aware, been previously treated.

Before describing my experiments, it may not be out of place

to give a short account of the history of the animal. Although

suspected by Cuvier to be but the larva of some unknown air-

breathing salamander or newt, the Axolotl of Mexico was

considered for many years to be one of the Perennibranchiata,

such as Proteus, Vecturus, and Siren. In the year 1863 a number

of specimens were imported from Mexico to France, where some,

kept in the Jardin des Plantes in Paris, bred, and the young were

successfully reared. The conclusion was not unnaturally drawn,

that the Axolotl, having bred in the branchiate condition, could

not possibly be anything but a perfect aquatic animal. It was,

not, however, until over two years later that the subject assumed

a different aspect, for some individuals of the second generation

lost their gillsand the dermal folds of the back and tail, developed

eyelids, and yellow spots on the skin, and taking to land, changed

into a land salamander, already well known from North America

as Amblystoma tigrinum.

A few years later Weismann (2) tried to solve the question

as to whether it were possible to force the larve, if brought into

conditions which rendered the use of the gills difficult and that

of the lungs easy, to change into Amblystomes, and he therefore

experimented with several broods which were placed in shallow

water, and thus compelled to breathe air more frequently.

Although he met with no success Weismann was not discouraged,

and came tothe conclusion that the failure of his experiments was

due to his having been unable to bestow the necessary care and

attention on the animals: he therefore asked Mlle. de Chauvin,

a lady who had already done much careful experimental work,

to take a number of larve, just out of the egg, rear them, and

make an attempt to bring them to the perfect condition. When

the Axolotls were about 6 months old, Mlle. de Chauvin placed

Proc. Zoo. Soc.—1913, No. XX VIII. 28

404 MR. E. G. BOULENGER ON THE

them in large glass vessels which were so disposed and the

water so restricted that at one spot only could they dive quite

under, while everywhere else they came into contact with the

air. The water was then gradually reduced. Within a few

days a change took place, the creatures leaving the water in from

four to fourteen days, the complete metamorphosis following

about ten days later. Mlle. de Chauvin (3) summarized her results

as follows :—‘ From what I have said, the correctness of the view

suggested by Weismann must be established, namely that most

Axolotl larvee, if not all, complete their metamorphosis, if in the

first place they come out of the ege healthy and are properly

fed, and in the second place meet with arrangements which

force them to change from breathing under water to breathing

above water.”

Dr. J. H. Powers (4) at Doane College, Nebraska, has more

recently conducted numerous experiments on the metamorphosis

of Nerth American examples of the Axolotl, and he hes come

to the conclusion that the metamorphosis is not due, as was

thought by Mlle. de Chauvin, to a direct response to changes in

conditions of environment, compelling them to resort to aerial

respiration, but to checked nutrition, and that a careful study of

Mlle. de Chauvin’s methods and results seems to cast a doubt

upon the conclusion that enforced air-breathing caused the

metamorphosis. The following is a passage from Dr. Powers’s

paper on the subject of this lady’s experiments :—‘‘ Fearing that

her charges would die, as indeed they sometimes did, she

always prepared them for the trying ordeal of metamorphosis by

raising the temperature of the water in which they were kept

and feeding to the maximum for several days, to which she

ascribes no other importance than giving the animals increased

strength. The Axolotls were then brought immediately into

water sufficiently shallow as to force them, at least part of the

time, to breathe air. In this latter condition the experimenter

complains again and again that it was next to impossible to

induce the Axolotls to take any food whatever. Thus in these

experiments we have high feeding followed by practical starva-

tion, and it seems that no control experiments were instituted to

determine what the effects of over and under nutrition might

have been with Axolot!s still in abundance of water. Yet most

interesting is it to note that even the varying factors of nutrition

seem to have been wholly neglected in the final interpretation of

the results.”

Dr. Powers’s field-notes show that metamorphosis occurs

rarely, if ever, as the result of enforced air-breathing through

the drying up of ponds, and that in spite of repeated search at

appropriate times and places, no Axolotls have been found trans-

forming on the mud of drying ponds.

Dr. Gadow (5), who not long ago visited the lakes near Mexico

City in which this creature lives, and where it is said to retain its

branchiate condition, has been able to refute the theories framed

METAMORPHOSIS OF THE AXOLOTL. 405

by various zoologists, as to why the Axolotl does not transform

in those localities. The reason he gives is that the unfailing

abundance of food and water, and the immerse hiding- places

amongst the reeds under the banks, constitute fon these

Batrachians a real paradise where they remain, in spite of the fact

that there is nothing to prevent them from leaving the water.

Dr. Gadow’s explanation appears to be in keeping with the results

of both Mlle. de Chauvin’s and Dr. Powers’s experiments.

The conflicting conclusions arrived at made it highly desirable

that further experiments should be undertaken. The lack of

success obtained by the majority of those who have attempted to

force the Axolotl to transform, has led many zoologists to believe

that the change is due to some congenital disposition, possessed

only by certain individuals, and that Mlle. de Chauvin and the

few others who have succeeded in obtaining Amblystomes from

the larval form were specially fortunate in the choice of their

subjects. Therefore, when I commenced my experiments, I was

by no means sanguine of attaining any definite results.

In August of last year I obtained five Axolotls with very fully

developed gills and fins, ranging in length from 105 mm. to

138 mm., and therefore probably from six to nine months old.

The enforced breathing of air, by the gradual absorption of

the water, and by the gradual elimination of the necessary amount

of oxygen from the water, starvation and irregular feeding, and

increased temperature having all been given or suggested as

means of obtaining the metamorphosis, sr placed the animals

under the following conditions :—

Nes. 1 & 2.—In shallow water so that after the first week

only the top of the animal’s head and fins remained un-

covered, thus forcing the creature to make free use of its

lungs. These I kept at a uniform temperature of between

75° and 80°.

Nos. 3 & 4.—Under similar conditions, but at a uniform

temperature of between 55° and 60°.

No. 5.—In deep water from which the necessary amount of

oxygen was eliminated by the gradual substitution of

boiled water and by the introduction of decomposing

vegetable matter.

Asat the time I could not obtain any more Axolotls of the size

required, I decided to abstain from experimenting with the

feeding, until I had either obtained negative results with those I

was about to experiment on or had secured more suitable spe-

cimens. Nearly a month later I was fortunate enough to obtain

six further suitable specimens. At the same time some of the

first brood showed signs of metamorphosing in spite of the fact

that they had fed with the greatest regularity. I therefore decided

not to abstain from offering them food as success seemed likely

without resorting to starvation, and placed the individuals of

the second brood, which shall be referred to as numbers 6, 7, 8, 9,

28*

406 MR. E. G. BOULENGER ON THE

Text-fig. 75.

STAGE f/f

STAGE 2

elt ot

STAGE 3

STAGE 4

First stages in the metamorphosis of Amblystoma tigrinum.

METAMORPHOSIS OF THE AXOLOTL. 407

Text-fig. 76.

ee ce STAGE 5

/| nt

Ih ‘

Ss Ze

STAGE 6

pee, SINGS 7

STAGE 8

Final stages in the metamorphosis of Ainbiystoma tigrinum.

408 MR. FE. G. BOULENGER ON THE

10, and 11, under similar conditions to those of brood 1. Nos. 6

and 7 were forced to breathe air, being placed in shallow water,

at a high temperature; Nos. 8, 9, and 10 were placed under the

same conditions at a normal temperature, and No. 11 was placed

along with No. 5 in foul water, from which the oxygen was

removed by the addition of boiled water. In all, therefore, I

had eleven specimens under artificial conditions with the object

of forcing the metamorphosis, and as a result succeeded in

bringing six individuals to transform into the Amblystome

stage, all six specimens being those induced to make free use of

their lungs by being placed in shallow water; three of these were

kept at a temperature of between 75° and 80°, three at between

55° and 60°. The time required for the entire metamorphosis

was from 12 to 16 weeks, periods which all to a remarkable degree

exceeded those taken by Mlle. de Chauvin’s specimens. ‘Three in-

dividuals kept under identical conditions died in from 33 months

to over 5 months, but not until after having undergone a change,

which, although considerable, did not necessarily suggest, as will

be referred to later, that, but for their death, the entire metamor-

phosis would have taken place. The two specimens kept in deep

water from which the oxygen was eliminated died in the one

case after 19 weeks, in the other after 22 weeks. These speci-

mens, but for a slight reduction in the size of their gills, did not

undergo any change, the fins being at the time of their death as

well developed as at the commencement of the experiment. It

is worthy of note that these specimens, in spite of the fact that

they must have felt exceedingly uncomfortable, did not rise to

the surface for air, even just prior to their death, more frequently

than individuals kept in oxygenated water. Another point of

interest lies in the fact that both specimens grew rapidly during

this period, No. 5 from 136 mm. to 155 mm., No. 11 from

112 mm. to 120mm. The specimens kept in shallow water, in

the case of those that died, remained stationary, while those that

transrormed actually decreased im size.

Text-figures 75 and 76 (pp. 406 and 407) represent eight stages

through which my Axolotls passed in the course of their

metamorphosis.

Stage 1 represents the perfect larval form, the condition which

all eleven were in before being placed in shallow water. A few

days after the animals had been placed in shallow water, the gills

with their fringes began to shrink, and the dorso-caudal fin

which lopped over to one side, to the left side in all my specimens,

became reduced in size, the change being plainly visible in from

a fortnight to three weeks (Stage 2).

In from 10 to 20 days later the gills were only half their

normal size, while the fin of the back was represented by a mere

ridge ; that of the tail, although having undergone further reduc-

tion, was still well developed on the upper surface and flopped

over to one side; on the lower surface, however, it showed con-

siderable reduction (Stage 3). It was not, however, until from

five to seven weeks later (Stage 4) that the metamorphosis proper

METAMORPHOSIS OF THE AXOLOTL. 4.09

took place, when the part of the dorsal fin nearest to the head

began to be entirely absorbed ; at the same time the head became

shorter, assuming a more Salamander-like shape, while a swelling

above the eye foreshadowed the advent of the upper eyelid; the

gills at this stage measured but from 3 to 5 mm.; the fin of the

upper edge of the tail, although now much reduced, still lopped

over to one side; the lower fin, but for a rudiment at the extreme

end, had entirely disappeared.

Stage 5 represents the conditions seven to eight days later,

when the dorsal fin had retreated to the middle of the back;

the caudal fin on the lower surface had entirely disappeared ; the

head had quite altered in aspect, having become much shorter,

giving the eyes the impression of having been shifted forwards.

At this stage the upper eyelid had become completely formed,

the lower eyelid being incomplete.

In from seven to nine days later (Stage 6), the fin of the back

had receded to a level with the hind limbs, while on the upper

surface of the tail, where it still lopped over, it measured only

1 mm. Both eyelids were now formed. At this stage the

animals raised their heads completely out of the water, and

the opportunities they were given of getting on land were

taken advantage of. Stage 7 represents the creatures about a

week later, when nothing remained of the fin but a mere rudi-

ment bordering the extreme end of the upper surface of the

tail. The gills were represented by three knobs. Stage 8, the

perfect form, the gill slits having closed up and the tail having

become more or less roundish, was attained five to eight days

later. In the case of both Duméril’s and Mlle. de Chauvin’s

specimens the yellow spots appeared several days prior to the

complete metamorphosis; in all my specimens, however, it was

not until after they had attained the perfect Amblystome con-

dition that the yellow spots put in an appearance, these being

situated on the sides of the body, tail, limbs, and throat; none

appeared on the back.

As I was desirous of ascertaining whether the metamorphosis

could be checked and the shrinking gills and fins made to undergo

fresh development, and if so, at what stage it could no longer be

checked, two specimens were taken out of the shallow water and

replaced in deep water as they reached successive stages in their

development. No.6 was kept ata temperature of between 75° and

80°, and No. 9 at a temperature of between 55° and 60°. Both

these Axolotls were placed in shallow water and then induced to

breathe air frequently by means of their lungs on October 7th.

Stage 2 was reached on October 20th; they were then replaced

in deep water. The gills grew immediately and the fins developed,

Stage 1 being reached again on October 28th and November 5th.

They were then once more put into shallow vessels, with the

result that the gills and fins again began to shrink, Stage 2 being

attained on November 15th and 20th, Stage 3 on December 5th

and 26th. On the latter dates they were again transferred to

deep water, Stage 2 being attained on December 20th and

January 10th. The metamorphosis was continued on placing the

410 MR. HE. G. BOULENGER ON THE

Axolotls in shallow water, Stage 3 being reached on January

10th and 25th, and Stage 4, the stage when: the front part of the

fin becomes totally absorbed, on February 12th and 28th. The

Axolotls were then returned to deep water: the development at

this stage, however, could no longer be checked and both creatures

completed their metamorphosis in the water.

From the latter experiment, showing that when replaced in

deep water the previous stages were resumed in about half the

time required to reach them, the conclusion may I think be

drawn that the natural tendency undoubtedly is for the animal

to remain an Axolotl, and that compulsion is needed to bring

about metamorphosis.

It will be noted that at Stage 4, when the front part of the

fin becomes absorbed, the metamorphosis can no longer be checked,

and that this is therefore to be regarded as the critical stage, at

which the further development must proceed whatever the

conditions under which the animal is placed.

During the course of all my experiments, with the exception

of the last stages of the metamorphosis, the Axolotls fed with

regularity twice a week, and in this respect [ experienced none

of Mile. de Chauvin’s difficulties.

Below will be found detailed accounts of the conditions under

which my eleven specimens were kept, and of the changes which

took place under the circumstances.

Specimen No. 1.

Placed in shallow water at a temperature of 75°-80°.

Stage 1. August 17th.—Length 138 mm.

September Ist.

September 15th.

October 17th.

October 24th.

November 2nd.

. November 8th.

November 14th.—Length 131 mm.

Total length of period—12 weeks.

DID OF wb

Specumen No. 2.

Placed in shallow water at a temperature of 75°-80°.

Stage 1. August 17th.—Length 119 mm.

. September Ist.

September 15th.

. November 9th.

November 16th.

November 21st.

November 29th.

December 4th.—Length 113 mm.

Total length of period—15 weeks

BONIS Ot pe bo

METAMORPHOSIS OF THE AXOLOTL. 41

Specimen No. 3.

Placed in shallow water at a temperature of 55°-60°.

Stage * August 17th.—Length 124 mm.

ees September 25th.

. October 10th.

November 12th.

. November 20th.

. November 27th.

. December 5th.

. December 11th.—Length 118 mm.

Total length of period—16 weeks.

OID ONE wb

Specimen No. +

Placed in shallow water at a temperature of 55o=608:

Stage 1. August 17th.—Length 105 mm.

Be SD: September 25th.

» 3 October 10th.

Died January 22nd.—Length 105 mm.

Specumen No. 3.

Placed in foul water from which the oxygen was removed by

the gradual addition of boiled water.

* Stage 1. August 17th.—Length 136 mm.

Died January 4th.—Length 155 mm.

Specumen No. 6.

Placed in shallow water at a temperature of 75°-80°, and

replaced in deep water as each successive stage was reached.

Stage 1. October 7th.—Length 134 mm.

» 2. October 20th.

Replaced in deep water.

» Ll. October 28th.

Replaced in shallow water.

», 2. November 15th.

», 3 December 5th.

Replaced in deep water.

» 2. December 20th.

Replaced in shallow water.

,, o January 10th.

,» 4. February 12th.

412 MR. E. G. BOULENGER ON THE

veplaced in deep water.

Stage 5. February 18th.

6. February 25th.

7. March 3rd.

8. March 10th.—Length 129 mm.

br)

Specimen No. 7.

Placed in shallow water at a temperature of 75°—-80°.

Stage 1. October 7th.—Length 98 mm.

2. October 31st.

» o& November 25th.

Died January 29th.—Length 98 mm.

Specimen No. 8.

Placed in shallow water at a temperature of 55°--60°.

Stage 1. October 7th.— Length 96 mm.

. October 20th.

. November 10th.

January 3rd.

. danuary 10th.

. January 19th.

. January 27th.

. February 4th.—Length 91 mm.

CON rE eo bt

Total length of period—164 weeks.

Specimen No. 9.

Placed in shallow water at a temperature of 55°-60°, and

replaced in deep water as each successive stage was reached.

Stage 1. October 7th.—Length 117 mm.

» 2. October 25th.

Replaced in deep water.

1. November oth.

Replaced in shallow water.

2. November 20th.

3. December 26th.

Replaced in deep water.

2. January 10th.

Replaced in shallow water.

3. January 25th.

4, February 28th.

METAMORPHOSIS OF THE AXOLOTL. 413

Replaced in deep water.

Stage 5. March 6th.

6. March 13th.

: 7. March 19th.

8. March 27th.—Length 113 mm.

Specimen No. 10.

Placed in shallow water at a temperature of 55°—-60°.

Stage 1. October 7th.—Length 105 mm.

2. October 31st.

, 3 November 30th.

Died March 24th.—Length 105 mm.

Specimen No. 11.

Placed in foul water from which the oxygen was removed

by the gradual addition of boiled water.

Stage 1. October 7th.—Length 112 mm.

Died March 22nd. ete b20) sonar

The results of these experiments show, I think, in the first

place that, in accordance with Mlle. de Chauvin’s observations,

and contrary to those of Dr. Powers, the Mexican Axolotl will,

with perhaps a few exceptions, transform into the Amblystome

stage 1f placed, when about six months old, under conditions

which force it to make frequent use of its lungs; secondly, that

starvation, irregular feeding, and temperature have no influence

on the metamorphosis; thirdly that, as no change occurs when

the Axolotl is placed in poorly oxygenated water, owing to the

fact that it will not under the circumstances rise to the surface

and make use of its lungs, the quantity of oxygen in the lakes

of Mexico can have little bearing on the explanation of the

phenomenon of neoteny; and lastly, that there is a critical stage

in the metamorphosis.

References.

. A. Dumtrit.—Comptes Rendus, lx. (1865), and Nouv. Arch.

Mus. 11. (1866).

. A. Weismann.—Zeitschr. wiss. Zool. xxv. (1875).

. Mute. pe CuAuvin.—Zeitschr. wiss. Zool. xxvii. (1876).

. J. H. Powrrs.—Amer. Nat. xxxvii. (1903).

. H. Gapow.—Nature, Ixvii. (1903).

Ce & 0

A414. SURGEON J. C. THOMPSON ON THE

32. Contributions to the Anatomy of the Ophidia.

By Josep C. THompson, Surgeon, United States Navy *.

[Received April 25, 1913; Read June 3, 1913.]

(Text-figures 77 & 78.)

INDEX. Page

Suggestion as to the Possible Origin of the so-called Tracheal

Notes upon the Anatomy of :—

AGAUSOOHNTEIS. WOIGOUOP ccacasosgcnn 569600 500509886 dovoadsd00b5 vovaen | Sli)

Cycloconusnliineatusmerce een ee ee ee ee Le

CRIS VGUCE, OPMGED cos odacsvccssodabooescaysoansssdocososcdexacqese | CHAO)

(CHROGTUIS: COPTAGDIEEB oso 000000 bodvoebadeco.ceese0 Sag acncoDsbocasendeae , SP

The Intromittent Organ in :—

PolyodontophisiOiwittatus cence seeeessctese eee eeee eee Ee

LUO HUCKONOGIS GUNCUUNFO Bonorecrnensses000250esoo00GscK0aA0aeo8008 43)

JEAUSCOIDS. CHUGRUGIGES ocaads coogucssoes8 00650005 6300a0 90000 000ccqana | GH)

Cycloconusilineatuserccn ce ee eet ee eee ee eee EO

SVad@reS (DOWUCECIS soc conccucoboodesccde dav occ sue sédooeooeovscccdan0c0 4A

Systematic :

SET D 14 S MEENA OVE es.c 2c sence teesaeeee eae eR a ee

Suggestion as to the Possible Origin of the so-called

Tracheal Lung.

It is suggested that the tracheal membrane has been developed

as a device originally contributing to the required flexibility

of the tube. The cartilaginous rings being interrupted behind,

or on one side, permit that the structure be subject to a much

greater pressure, especially when bulky morsels are being

swallowed, than would be the case were the rings complete.

The tracheal membrane, being continuous with the lung, appears

to have afforded a convenient foundation for the extension of

pulmonary tissue when the need for an increased breathing

surface has arisen, or when the lung has been encroached upon

by other organs.

In the Hydrophide an increased pulmonary area is required

both to give buoyancy and to enable the sea-snakes to remain

some time beneath the surface. The most specialized species in

the genus Hydrophis Daudin, those in the fasciatus group, possess

a lung that extends nearly to the vent, and a tracheal lung of

due proportion.

In the Acrochordine, the members of which are of aquatic

habits, there is also a highly developed tracheal lung. Owing to

the abnormal size of the heart the respiratory tissue on the

trachea is widely separated from the lung itself.

In the Viperide the elaboration of venom requires a large

liver. These serpents as a rule have short and thick bodies, and

the liver invades the region just caudad of the heart. As a

* Communicated by Dr. F. E. Bepparp, M.A., F.R.S., F.ZS.

ANATOMY OF THE OPHIDIA. AlL5

result of this the crowded pulmonary tissue seeks the direction

of least resistance, which is along the tracheal membrane.

In recording the position of the viscera it has been found

advantageous to adopt the following routine with the view of

correlating the data obtained. The serial number of the gastro-

stege that underlies the anterior tip or the posterior extremity

of an organ is taken as the external landmark. This number is

reduced “to a percentage, the total number of the ventral shields

in the specimen being used as the base. From this procedure

there results a set of figures which will materially aid in com-

paring one species with another or in learning the extent of the

variation which the individuals of a given form exhibit.

It appears that whenever a serpent varies widely in an

important character from one of the natural groups, a further

investigation regularly results in the finding of several additional

structural features that are well worth making a matter of

record. The single species constituting the family of Xenopeltidee

may be taken as an illustration of this general statement.

Some Notes wpon Anatomy.

XENOPELTIS UNICOLOR Reinhardt. (Text-figs. 77 & 78.)

Specumen. No. 16750, California Acad. Sci. Singapore.

Female ; total length 480, tail 58 mm.

Squamation.—Scales in 15 rows anteriorly, and the same

posteriorly, an oblique series commencing at one gastrostege

Text-fig. 77.

ae,

A B Cc

Teeth of Xenopeltis unicolor.

A. Maxillary bone viewed at a right angle to the outer edge, and showing the

oblique fashion in which each tooth is set in the jaw.

._B. Profile view of the same teeth, showing the lateral cutting-edges of the cusp.

C. Palatine tooth viewed from below, and showing a stout and blunt tip which

exhibits a tendency to assume the arrow-head shape of the cusps.

terminates at the ninth ventral shield to the rear. Gastrosteges

177. Anal divided. Urosteges 32 pairs, the second entire. Loreal

and preocular absent. Postoculars 2. Anterior temporals 2,

posterior 3. Supralabials 8, the fourth and fifth entering the

eye. Infralabials 9.

416 SURGEON J. C. THOMPSON ON THE

Anatomy.—There are two dental characters in this species that

are unparallelled among the Ophidia. The most striking is the

shape of the individual tooth, and the fact that the teeth borne

on the palatine bone are decidedly the largest. Each tooth is

set in the alveolar ridge with the anterior surface directed

forward and outward at an angle of 45° with the long axis of the

bone. When a tooth is viewed from the anterior surface, it

appears a trifle constricted at the middle; towards the extremity

it flares out laterally, and terminates like a blunt spear-head,

with two cutting-edges that meet at a right angle. When

viewed from the side, this anterior cutting-edge is seen as a

narrow, beveled cusp, and from its base the tooth is continued

backwards as a horizontal process, the length of which is equal

to the broad diameter of the tooth. The edges of the cusp and

the tip of the backward—directed point are covered with brownish

enamel.

The hypapophyses are present on the anterior vertebre and

absent on the posterior. They have a moderately long base.

In the writings of Cope * this form is credited with possessing

a coronoid bone. By Mr. Boulenger, however, it is placed in

the category of those in which this structure is absent. Two

specimens have been examined and no vestige of the bone can

be found.

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :—

Total number of gastrosteges...... 177 100 p. cent.

ANDERS O18 INGA GE oasanogageodbooseoqdac 52 29-4

Wiversambertor tip) s.-5:- ere eeere 64 39°]

PE DOSLeLIOn end ann. abt 6 116 65°3

Gall-bladder, middle ............... 129 73

Kidney, right, anterior tip ...... 146 82°5

As » posteriorend ... 155

BS left, anterior tip ...... 150 85

ss » posteriorend ... 161

iDleo-Geacell Waly) Bocdsesonsocbnocsocne 157 88°8

The heart is large, and its position is indicated on the thoracic

walls by a distinct bulging. Over the base of the organ there

are two areas in which the scales are enlarged; each area is

triangular, with the base along the edge of the ventrals and the

apex at the upper border of the fourth row. In the first row are

five scales, each of which is one-third larger than those before or

behind the heart. The scales in the seven dorsal rows are not

altered. The increase in the diameter of the body over the heart

is compensated for by an increase in the size of the scales in the

outer rows, and not by the interpolation of an additional series.

The tracheal rings are complete from the glottis to the ninth

gastrostege. From this point the rings are interrupted, and the

* Croc., Liz. & Sn. N. Amer 900, p. 731.

ANATOMY OF THE OPHIDIA. 417

dorsal wall of the trachea is formed by the tracheal membrane

which joins the ends of the incomplete rings. From the 9th

to the 21st gastrostege the ends of the rings are in con-

tact and the membrane lies collapsed above them. From this

point to the base of the heart, the membrane widens and the

ends of the rings are permanently separated. Dorsal to the

heart the tracheal membrane winds around to the right side of

Text-fig. 78.

Fi SCO

ise

Base of the lungs of Xenopeltis unicolor.

Viewed from below; the object being te display the bridge uniting the two lungs,

the termination of the trachea, the long free apex of the left lung, the adherent

apex of the right lung, with its tiny free tip, and anterior to it the portion of

the tracheal membrane which is lined internally with pulmonary tissue and is

bordered by a branch of the pulmonary artery.

the tube, and 2:5 mm. before the trachea enters the lung it

becomes lined with pulmonary tissue. The two lungs are

adherent for a distance of 10 mm., the connecting bridge

extending from the 53rd to the 57th gastrostege. Viewed from

within, this bridge appears as a septum lined with alveoli and

418 SURGEON J. C. THOMPSON ON THE

dividing one lung from the other. There are two perforations

affording communication between the lungs; the larger, measuring

2mm. in diameter, is at the termination of the trachea, and the

smaller, measuring 1 mm. in diameter, is near the posterior

border of the septum. The trachea ends abruptly 3 mm. behind

the apex of the heart, and on the lower surface of the bridge.

The right lung extends from the 49th to the 119th gastrostege.

Anteriorly, it reaches 8 mm. beyond the border of the bridge;

it is adherent to the right side of the trachea for the distance of

6 mm., and has a free apex 2 mm. long. The trachea communi-

cates with the lung 2°5 mm. behind the point where they join.

The left lung extends from the 50th to the 77th gastrostege.

Anteriorly it terminates in an acutely pointed free apex 8 mm.

long, at the level of the auriculo-ventricular septum. Poste-

riorly the lung terminates in a blunt cone, and towards the

end the walls are a trifle thinner and the alveoli larger. There

are two pulmonary veins. The right courses along the angular

ventral border of the right lung and enters the anterior inferior

corner of the auricle. The left commences at the posterior

border of the bridge and enters the posterior inferior corner of

the auricle. This vein is the thicker of the two. It is formed

of two branches that arise along the mesial side of each lung;

the branch from the right lung is the larger, and it has frequent

anastomoses with the right pulmonary vein.

The liver extends from the 64th to the 116th gastrostege.

There is no trace of segmentation.

The centre of the gall-bladder is at the 129th gastrostege.

The right kidney extends from the 146th to the 155th, and

the left from the 150th to the 16lst gastrostege. Each is

semidivided into nine irregular lobes. The right kidney is

supped by two renal arteries, the anterior entering at the

second and the posterior at the sixth lobe. There is a single

renal vein leaving the anterior tip of the organ, and this promptly

unites with its fellow to form a common trunk.

The lining of the rectum is smooth. The ileo-cecal valve is

at the 157th gastrostege. ‘The rectal cecum is 9 mm. long, and

is directed forwards and lies on the right side of the ileum. Its

walls are thin, and the opening into the cecum is just anterior

and to the right of the valve. Its lumen was filled with fecal

matter, though the cecum and rectum were empty.

Along the dorsal wall of the cloaca there is a longitudinal,

rounded ridge, which terminates in a papilla 1 mm. in length,

and at the level of the posterior border of the last gastrostege.

This ridge is formed by the two parallel tubes of the completely

divided vagina ; these terminate in two separate orifices at the

tip of the papilla. The lining of the vagina is finely and

longitudinally plicate. According to Cope* the vagina in the

Peropoda is undivided and the walls are nearly or quite smooth,

* Op. cit. p. 7Q0.

ANATOMY OF THE OPHIDIA. 419

and in the Colubroidea the organ is bilobate and the walls have

deep longitudinal grooves. This form more closely resembles the

latter.

CycLocoruUS LINEATUS Reinhardt.

Specimen. No. 15240. California Acad. Sci. Olongapo,

Luzon, P.I. Female; total length 503, tail 81 mm.

Squamation.—Scale-rows 17 anteriorly ; the V row suppressed

at the 127th gastrostege on the right and the 125th on the left,

leaving 15 rows which are continuous to the vent. The last

scale in the row that is dropped is before an enlarged scale

in the row below. Anteriorly an oblique series of scales

starting at one gastrostege terminates at the tenth to the rear.

Gastrosteges 163. Anal entire. Urosteges 41, single, Pra-

oculars 2, postoculars 2. Anterior temporals 2, posterior 2.

Supralabials 8; the third to the fifth entering the eye. Infra-

labials 9; the first to the fifth in contact with the anterior

geneials. Anterior larger than the posterior geneials.

Anatomy.—The hypapophyses on the dorsal vertebre are

continuous throughout the column. On the vertebra above the

122nd_ gastrostege the base of the hypapophysis occupies

the posterior one-third of the centrum; the apex is horizontally

truneate, and extends a trifle posterior to the vertical of its

centrum. The caudal vertebra at the level of the 12th urostege

has two triangular hemapophyses the bases of which occupy the

posterior half of the centrum. The inferior zygapophyses are

developed as broad wing-like plates, 1-4 mm. long.

The maxillary bone at the junction of the anterior and middle

third is bent inwards at an angle of 45°, and the teeth are

arranged in two groups. The first set consists of 7 teeth that

are mounted on the anterior one-third of the bone; the first five

are very small and increase in size posteriorly ; the sixth is

nearly three times the size of the fifth, and the seventh is larger

still, being 1:5 mm. long. These teeth are inclined backwards.

The two groups are separated by an interval of 155 mm. The

second set contains 13 teeth ; the first nine are small and nearly

equai; the tenth to the fourteenth increase rapidly until the last

is twice the size of the ninth. These teeth are inclined towards

the median line. The anterior tip of the palatine bone reaches

the interval between the fifth and sixth maxillary teeth; there

are 16 teeth. The pterygoid bone bears 19 teeth; these are on

a line that is convex externally ; the middle teeth being three

times as far from their fellows on the opposite bone as are the

teeth at the extremes. The dentary bone is also bent inwards at

an angle of 45°, and has the teeth in two groups. The first set

is mounted on the inflexed portion of the bone, and consists of

6 teeth; the first to fourth gradually increasing in size; the

fifth and sixth strongly enlarged. The two groups are separated

by an interval which is shorter than the sixth tooth. The second

set consists of 17 very small teeth.

Proc. Zoot. Soc.—1913, No. XXIX, — 29

420 SURGEON J. C. THOMPSON ON THE

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :—

Total number of gastrosteges ... 174 100 p. cent.

INGE OF INCAVAY 5 coasodvocnscscs0e060r 37 21:3

JbmyEIe, QVNESOTE UND) oogconsconsdecoass 44 25°3

7 posterior end ~.......-...... 96 55-1

Grille Glee dogepdocesbsoosesodaceds 114 65°3

Kidney, right, anterior tip ...... 142 81:8

i; } posterion endy i.) 29 85°8

As left, anterior tip......... 144 82°8

a iP posberlom endian tol 86°8

iileo=ccecalliivalivielec en snc-ce steer 154 88°95

The tracheal membrane begins at the glottis, and for a short

distance lies along the right side of the tube. Almost immediately

it broadens, and at gastrostege 14 it is lined with respiratory

tissue. At gastrostege 22 the membrane alters its position so

that it forms the dorsal instead of the right wall of the trachea.

Well above the apex of the heart the alveoli on the membrane

have assumed the same size and shape as those in the lung; there

is no gross difference to indicate the ending of one structure and

the beginning of the other. The trachea terminates abruptly

7 mm. posteriorly to the apex of the heart. There is uo rudi-

mentary lung. Arteriorly the walls of the lung are thick and

lined for the entire circumference with pulmonary tissue ;

posteriorly they become thin and terminate in an air-sac. The

lateral lobes of the liver are distinct. The left lobe is the longer ;

anteriorly it extends beyond the right lobe 6 mm. and posteriorly

jl mm. There are four transverse fissures on the left lobe near

the anterior tip, the rest of the organ is smooth. The anterior

one-third of the rectum is thickly and regularly plicate; the

posterior two-thirds are irregularly folded. There is no body to

the vagina. The organ is bilaterally divided into two separate

tubes that open into the dorsal wall of the cloaca by two separate

orifices, which are 2 mm. apart.

Notes.—This species is described as having scales with apical

pits, and an eye with a round pupil. Over a dozen specimens

have been studied and the pits cannot be made out. The pupil

is usually round in the young and frequently vertically oval in

the adult.

CHRYSOPELEA ORNATA Shaw.

Specimen. No. 16707. California Acad. Sei. Cochin China.

Female; total length 910, tail 237 mm.

Squamation.—The number of scale-rows on the body, the

sequence In which they become suppressed, and the gastrostege

level at which they terminate on each side may be thus

presented —

17 rows, IV row ends, right 137th, left 136th gastrostege, leaving :

yee VALLE ees, 4 Wadia, op Jashiln

13. ,, which are cortinuous to the vent.

ee] 29

ANATOMY OF TIE OPHIDIA. 421

Gastrosteges 227, the last shield divided. Anal entire.

Uvosteges 120 pairs. Supralabials 10; the fifth to the seventh

entering the eye; on the right the fourth is reduced and fused

with the posterior inferior angle of the third. Infralabials 10 on

the left, 11 on the right, the additional shield is between the

corresponding third and fourth, Anterior geneials larger than

the posterior, the right in contact with four and the left with

five infralabials. Gular shields in six pairs,

Anatomy.—Maxillary bone with 20 teeth, the posterior three

with a broad shallow groove; the five preceding these are

also grooved but less distinctly. Palatine bone with 8 teeth,

increasing in size posteriorly ; the anterior tip of the bone

reaches to the interval between the second and third maxillary

teeth. Pterygoid bone with 28 teeth. Dentary bone with

18 teeth, the anterior enlarged and grooved on the external

quadrant.

In terms of gastrosteges the external landmarks of the viscera

are as follows :—

Total number of gastrosteges ...... 227 100 p. cent.

Aexcolshearty Gara ssct-ceiscecat|sce hes 58 25°6

Miversanterlonibip (reece osece os: 73 32°2

5) [DOSS Cha VOSS a Soebgacoeeuae. 112 A9-4

Galll=bladden centre tase. .0.0ssee eee 232 58°2

Kidney, right, anterior tip ...... .. 183 80°8

ps oy) posterior endi= sce 201 89

ee elit era mbenroritipis ween. 196 86°5

cs ye posterior ends. 214 94-2

lilewzceecallt valves sae eeees asec aeee 209 92-2

Waginalcormuay basemen .c eee 216 95

The cartilaginous rings of the trachea are complete from the

elottis to the angle of the jaw; from this to their termination

on the ventral surface of the lung, at the apex of the heart, they

ave interrupted. ‘The tracheal membrane begins with the semi-

rings; it is on the right side of the tube, and is continued eaudad

to the base of the heart. For the greater part of its length it

comprises more than half the circumference of the tube. At

its termination it becomes continuous with the pulmonary pleura.

The elasticity of the rings maintains their free ends in close

apposition, and the membrane lies in a lax state by the side of

the trachea. When the windpipe is dilated, the ends of the

rings are separated and the tracheal membrane becomes a

functioning part of the organ. Under ordinary conditions the

air passes only through that part of the tube that is formed by

the semi-rings. Anteriorly the pulmonary tissue begins at the

level of the auriculo-ventricular septum. In the lung it is

confined strictly to the dorsal half of the circumference.

Posteriorly it is drawn out as a fine streak from which are sent

out transverse septa ; it terminates on the left side of the air-sac,

at the level of the 79th ventral. Posteriorly the lung is continued

as a membranous air-sac; exactly where it ends cannot be

G ¥

bo 4%

423 SURGEON J. C. THOMPSON ON THE

ascertained. There is a small rudimentary lung, a mere vesicle

without pulmonary tissue. ‘The anterior tip of the liver is not

divided into lateral lobes ; posteriorly the left lobe is about three

times as thick as the right, and is 5 mm. longer, The surface is

smooth, there being no transverse fissures. The ileo-czcal valve

is at the 209th gastrostege ; there is no cut-off tube or rectal-

cecum. The lumen of the rectum is longitudinally plicate, with

faint closely-set transverse ridges. The vagina bifurcates at the

level of the 216th gastrostege, the walls are smooth.

Habits.—The observations made by Mr. Shelford * on the aereal

locomotion that this serpent indulges in afforded one of the most

interesting field-notes on a reptile that has been published for

some time. Attention was drawn to the hinge-lines along the

gastrosteges, and to the habit of retracting the middle section of

these shields “ so that the snake became deeply concave along the

ventral surface ”. This serpent has been captured several times

in Luzon, and the same performance witnessed. If the lung in

this species had thick walls for its entire circumference, this

radical change in the shape of the body could not be so readily

brought about. On the contrary, 1t is so modified that when

subject to ventral pressure it merely has the lower membranous

wall stove in and no serious interference with respiration takes

place.

CROTALUS CONFLUENTUS Say.

Specimen. Field No. 8208. U.S. Nat. Mus. Mt. Tamilpais,

alt. 650 M., California. Female; total length 460, tail 45 mm.

Squamation.—The number of scale-rows on the body, the

sequence in which they become suppressed, and the gastrostege

level at which they terminate on each side may be thus

presented :—

25 rows, VI row ends, right 108th, left 160th gastrostege, leaving :

Me egy Vigbes. on sis Fl okd ees Gla 5 i

PHI Pio 8 One EIN, 45 G2! bs ia

20. X ,, intermittent between 124th and 165th __,,

19 ,, which are continuous to the vent.

Ventrals 174. Anal entire. Urosteges 21 entire, the first

and last three paired. Preoculars 2, suboculars 2, postoculars 4.

Supralabials 14 on the right, 13 on the left side; the reduced

count due to the fusion of the fourth and fifth shields.

Anatomy.—The external landmarks of the principal viscera in

terms of gastrosteges are as follows :—

Total number of gastrosteges ... 174 100 p. cent.

Igoe OH INCHED ooooodoonqvsnscdesacoase 70 40:2

ToihyGIey PyOEVAOVe (HN)8), -benccoanabaoonsos 70 40-2

py ee postenioncemdiinn rare. ses. 109 62°5

* “ A Note on ‘ Flying’ Snakes,” P. Z.S. 1906, p. 227.

ANATOMY OF THE OPHIDIA. 423

Kidney, right, anterior tip ...... 144 82°8 p. cent.

> 3, posteriorend ..: 163

3 left, anterior tip ......... 146 84

. » . posteriorend ... 166

Wei@Quaen, GYD eobeogsstanbecsonoddads 164 99-2

The cartilaginous rings of the trachea are complete from the

glottis to the 6th gastrostege. At this point the tracheal

membrane begins on the dor sal quadrant of the tube. Almost

immediately, ‘while still very narrow, it acquires pulmonary

tissue, this rapidly increases and becomes continuous with the

lung, The semi-rings extend along the ventral surface of the

trachea ; viewed from. within they appear as a narrow gutter; they

terminate 15 mm. caudad from the apex of the heart. The walls

of the lung posterior to the heart have a very thin lining of

pulmonary “tissue ; the alveoli are large and shallow in contrast

with those in the trachea, which are small, thick, and superimposed.

In other words, the area of the respiratory tissue in the lung is

much less than it is in the trachea. The liver is deeply divided

into a right and left lobe, and these are partially divided by

shallow and transverse fissures. Anteriorly the left lobe is thick

and rounded, extending three gastrosteges beyond the right, and

overlapping the apex of the heart by 1 mm. The peritoneum

covering the liver and the abdominal walls is pigmented, the

remainder is plain. The vagina is short, it extends over 10

gastrosteges, and is bilobate. The cornua are adherent to each

other at the base for a distance of 15 mm. Externally each

horn gives the appearance of being composed of two adherent

tubes, with the oviduct entering the outer. The lining is

smooth, and is thrown into several longitudinal folds; two of

these are larger than the rest, and their free edges nearly

touching tend to divide the lumen of each horn into an inner

and an outer compartment.

The Intromittent Organ.

Polyodontophis bivittatus Boulenger.—The hemipenis is un-

divided and the sulcus simple. The base is plicate and is followed

by a spinous zone 4 mm. in length. The spines are about ten in

number ; there are two placed opposite to the sulcus that are

much enlarged, being 2°5 mm. in length. The distal 6°5 mm. of

the organ 1s calyculate. The calyces are very small and their

borders bear numerous minute spines. The calyces at the border

of the spinous zone and at the tip are enlarged, the latter have

simple borders.

In Cope’s classification the enlarged basal hooks place this form

in the Natricine. The occurrence of these hooks with a calyculate

apex represents a divergence in the direction of the Colubrine.

Tropidonotus vibakari Boie.-—The hemipenis and the sulcus

spermaticus are undivided. The organ is densely spinous; those

on the middle are a trifle the larger, diminishing in size towards

494. SURGEON J. C. THOMPSON ON THE

the apex and the base. There are a few enlarged basal hooks

situated close to the suleus, one being on one side and four on

the other; of these four the one nearest the base is double the

size of any of the others. The tip of the organ is rounded and

smooth, and the line of demareation between the bare and the

spinous areas is sharp. On either side of the smooth area at the

tip, there are mounted two apical papille, which are one-fourth

the length of the body of the organ. Each papilla is spinous for

three-fourths of its circumference, the inner quadrant being

smooth and continuous with the spineless area at the tip.

Terminal papille similar to those in this species are of com-

paratively rare occurrence. Cope has reported them in the genus

Oligodon Boie, and in the three following species: Jropidonotus

vittatus Linneus, [schnognathus lineatus Hallowell, and Coluber

helene Daudin. Hach of these has been set apart by Cope as

the type of a new genus, established on the ground that the

intromittent organ differed widely from the type prevailing

among the nearest allies. 7. vibakari in this respect is clearly

separated from the Far Eastern representatives of the genus.

These have been placed in the genus Bothrodytes by Cope, and

are characterized by having the organ furcate. The organ in

vibakari most closely resembles that found in Ischnognathus

lineatus Hallowell, and the figure given by Cope* of this

species shows that it differs from vibalari in the one detail of

being less completely covered with spines. Furthermore, this

serpent is of precisely the same mild-actioned and fearless nature

that is characteristic of the species of /schnognathus. In this it

offers a marked contrast with the Far Hastern members of the

genus Tropidonotus. Over a dozen species of these have been

captured and every one showed a nervous, irritable, and snappy

disposition. bain

To allow this: species to remain in a complex genus like

Tropidonotus, after it has been found to differ so radically, 1s not

good taxonomy. On the other hand, to chronicle promptly each

new anatomical discovery under a new generic caption does not

materially aid our studies. It is believed that the dilemma can

be overcome for the time being by strictly adhering to the

nomenclature contained in the ‘Catalogue of Snakes in the

British Museum,’ and at the same time by registering a pro-

visional new generic term. This new term is not to become

current until the entire group to which it belongs has been

re-examined from the standpoint of comparative anatomy.

With this understanding it is suggested that the new genus

Hebius be established for Tropidonotus vibakari Boie, the salient

character distinguishing it from all species of Zropidenotus

being the densely spinous and undivided hemipenis, bearing two

long apical papillee.

Helicops angulatus Linneus.—The total length of the hemi-

penis is 42 mm.; it is divided into two quadripenes, the length

* Op. cit. pl. xx. fig. 12.

ANATOMY OF THE OPLIDIA. 425

of each being 10 mm. The quadripenis is calyculate, and the

cells are arranged in seven rows that trend obliquely forward

from the suleus. he distal half of the body of the hemipenis is

spinous; opposite to the suleus there is a septal fold. The

proximal half is plicate. At the border of the spinous area and

close to the sulcus there are two round, flat-topped, and gristle-

like papille ; these are closely set and one is distal to the other.

The sulcus is divided at a point 12 mm. from the base, this is

well within the plicate area.

Mr. Rosén* writes that the hypapophyses are absent from the

posterior vertebre in H. modestus Giinther and H. leopardinus

Schlegel. In this species they are well developed throughout,

one occurring even on the last dorsal vertebra.

Cyclocorus lineatus Reinhardt.—The hemipenis is of extreme

length and slenderness, being 38 mm. long and 2 mm. in diameter.

It is not divided and the sulcus is simple. The base for the

distance of 4 mm. is plicate; the remainder of the organ, even

to the tip, is spinous. Towards the tip there are three slightly

elevated longitudinal ridges, one 8 mm., the others 5 mm. in

length. The spines are nearly uniform in size, being about

°3 mm. to 4 mm.long. Each is mounted on a broad fleshy base,

and is slightly recurved. When the walls are stretched the

spines are seen to be arranged in longitudinal series.

Spilotes pullatus Linneus.—The structure in this species is

one of the most complicated recorded. The total length of the

hemipenis from the base to the insertion of the retractor muscle

is 77 mm. The organ and sulcus spermaticus are undivided.

From the base to the 20 mm. point are longitudinal plications ;

the folds are thick, at first straight, later becoming wavy. From

the 20 mm. to the 38 mm. point it is densely beset with stout,

blunt spines, each of which is about 2-5 mm. long. From the

38 mm. to the 54 mm. point on each side of the sulcus are rows

of flounces, these trend obliquely forward and acquire partitions

which convert them into rows of calyculi; the cells of these

calyculi increase rapidly in size, the largest being situated

opposite to the sulcus. On each side of the sulcus at the 60 mm.

point are two thick flaps 10 mm. long; these are in the longi-

tudinal plane, hang free in the lumen of the organ, and are

covered with small regular ruches. Between one of these flaps

and the sulcus there is a thin membraneous septum 20 mm. long,

parallel to the sulcus, and hanging free in the lumen. Towards

the extremity of the organ the calyculi change to irregular

flounces; these in turn are converted at the extreme tip into

minute frills.

United States Fisheries Steamer ‘ Albatross ’,

Sausalito, California, April Ist, 1913.

* Ann, Nat. Hist. (7) xv. 1905, pp. 170-171, fig. 1.

426 DR. S. F. HARMER ON THE

33. The Polyzoa* of Waterworks.

By Sipney F. Harmer, Sc.D., F.R.S., F.Z.8.7

[Received March 11, 1913: Read April 22, 1913.]

(Plates LXIL. & LXILL2)

Inpex.

Bionomies : Page

Relation of Polyzoa tothe other Constituents of the Fauna and Flora... 428

Economic importance of Polyzoa .......... Pu Reet ncc ss ea REO,

Records of occurrence of Polyzoa in Branch VWiatonworeel Fae oe cs eo OD

Systematic :

JEGHORICANG GRRECCOWAEE 450.503 090 009200 048 oEneO6 od 600 SB LOD ecanagedosodasedanannocacca . 4lal

Fredericella sultana .......... Pe Recn ee ronoselcescucanmonecenteccdcadaacual | (aiats)

Plumatella fungosa, var. Perales BRE atar toc oni tonicooseconarcoepmog neh aoe 449

Plimatella emarginata, VAL. MUSCOSH 0.60 ce0cceccveeeceecseceeeeeisecessss en, 452) ©

Development of hibernacula of Puludicella .......0.c.0c1eccccecsee eee eeereceseresere 444

Morphology of hibernacula and statoblasts

I. Historical Account.

It is perhaps not generally recognized in this country that,

under certain circumstances, Polyzoa may be of great economic

importance. But it has long been known to those interested in

the subject, particularly in Germany, Holland, Belgium, and the

United States, that a group of organisms, among which certain

Polyzoa play a leading part, may flourish in the pipes of water-

works which are not provided with an efficient filtering apparatus,

to such an extent as to give rise to the most serious inconvenience,

and may indeed finally throw the whole system out of gear. It

is the object of this paper to show, by means of examples which

have come under my notice during the last two or three years,

that this country is by no means exempt from the risks which

have been experienced in various places abroad.

The first important contribution to the subject was an ex-

tremely interesting paper by Kraepelin (85), published under the

title of “ Die Fauna der Hamburger Wasserleitung” ; although,

AS Kraepelin himself points out, a list of 18 species of amines

from the pipes of the same water- supply had been published

by Petersen some nine years earlier. The occurrence of such

organisms as Asellus and Gammarus in the water supplied to

houses in Hamburg was no rare event in that city; while the

pipes were frequently choked by Eels, ‘ Leitungsmoos” and

other organisms. Kraepelin undertook a biological investigation

of the subject, mfluenced partly by the hope of finding blind

Crustacea among the inhabitants of the water-pipes, to which

* [In view of the difference of opinion as to whether this Phylum should be called

Polyzoa or Bryozoa (see Proc. Linn. Soc. 1911, p. 63), I have accepted the preference

of the author.—Ep1ror. |

+ Published by permission of the Trustees of the British Museum.

+ For explanation of the Plates see p. 456.

BP, 4A. S.IGIS, IPL, IAN.

0 20 40 60 80 100 iS)

8. tO).

5.F. Harmer del. j Bale & Danielsson ,L*4 imp.

PALUDICELLA ARTICULATA.

12, Ge S._ IIS), Jet, ILD,

| | ae. 25.

S. F. Harmer, del. ; Bele & Danielsson, L*? imp.

STATOBLASTS OF PHYLACTOLAEMATA.

POLYZOA OF WATERWORKS. 497

light had, of course, no access; and by the further consideration

that he might be able to detect some modifications of structure,

as compared with the ordinary inhabitants of the Elbe, in the

animals which were living under these unusual conditions.

Although the results of the investigation were not of the nature

which he had anticipated as possible, they were nevertheless very

surprising; since he was able to demonstrate that some fifty

genera of animals occurred, often in enormous numbers, in the

system. The examination was made with the assistance of a

specially devised piece of apparatus which was screwed on to

various parts of the water-mains. A column of water was

allowed to escape through this arrangement, and the animals

which were contained in it were filtered off. The results obtained

were very uniform, in whatever part of the system the apparatus

was used, so that it became clear that the organisms found were

not merely a fortuitous collection of individuals which had acei-

dentally been introduced into the supply, but were, on the

contrary, perfectly normal constituents of a definite fauna in-

habiting the water-pipes under a pressure of 23-53 atmospheres.

Among these organisms a conspicuous, and usually the greater,

part consisted of large masses of what had become known to the

workmen as ‘“ Leitungsmoos ”—a mass of entangled tubes which

belonged principally to Polyzoa of the genera Mredericella, Pluma-

tella and Paludicella, but partly to the well-known Hydroid

Cordylophora. Sponges (Hphydatia fluviatilis and Spongilla

lacustris) were frequently found. Colourless forms of Hydra

were discovered; but, as might have been expected, /. viridis

appeared to be completely absent. Several species of Freshwater

Oligocheetes were recorded, as well as “almost incredibly large

numbers” of Leeches (Glossiphonia and Nephelis). Parasitic

worms were represented by Lchinorhynchus, found in large

numbers, in its larval condition, in Asellus; and, in its adult

condition, apparently derived from Eels. Among the Polyzoa

the “ mossy” appearance of the masses of tubes above alluded to

was generally due to Fredericella sultana. Traces of Paludicella

articulata were generally found ; while Plumatella was represented

principally by a form described by Kraepelin as P. princeps, var.

muscosa, and by the Alcyonelloid P. fungosa. Almost every

sample contained hundreds, or indeed thousands, of <Asellus

aquaticus, which were found creeping over the Polyzoa “in ugly

crowds.” The Freshwater Shrimp, Gammarus pulex, was some-

what less numerous, while various forms of Cladocera, Copepoda,

and Ostracoda were nearly always present. Leander | Palemon|

squilla and Mysis chameleon were represented by single finds. A

small water-mite and a few Dipterous larvee and pup were rarely

met with. '

Among Molluscs the commonest species was Dreissensia poly-

morpha; but Bithynia tentaculata was frequent, and other fresh-

water genera—Physa, Limnea, Planorbis, Ancylus, and Spheriwum

(Cyclas)—were also discovered; while Petersen had previously

428 DR. S, F. HARMER ON THE

recorded Vivipara (Paludina), Anodonta and Unio. Planarians

and a Nemertine (Zetrastenma) were also found, although the

method employed for collecting the samples was not well adapted

for the discovery of the smaller organisms. Rotifers and Infusoria

(Vorticella, Stentor, Paramecium, and Acineta) were, however,

discovered.

Even the Vertebrates were not unrepresented in this remark-

able fauna. The Eel, up to a foot in length, was common. As

many as six individuals were sometimes present in one sample,

and Kraepelin estimated that there must have been many

thousands in the system. Other fishes were less numerous ;

but the Stickleback (Gasterostews aculeatus) was occasionally

found ; and on one occasion a Burbot (Lota vulgaris), a foot in

length, was discovered. A young Flounder, several centimetres

long, was found in company with the Prawn (Leander) above

alluded to.

Summarizing his observations, Kraepelin points out that the

constituents of the fauna of the water-pipes consist almost exclu-

sively of forms which feed either on detritus or on one another ;

that in the absence of ordinary green plants, forms which feed on

these are absent ; while animals like insects, which require access

to air, are also absent. For the species which had succeeded in

adapting themselves to their novel environment the conditions

were, on the contrary, exceptionally favourable, and indeed more

favourable than those found normally in Nature. Some of the

factors specially alluded to, in this connexion, were the absence

of carnivorous insects, and the existence of large surfaces suitable

for the attachment of sessile animals.

Kraepelin proceeds to consider the ultimate source of the food

of the pipe-fauna; and he points out that this must consist of

the microscopic organisms and other detritus introduced into the

system by the constant flow of water from the Elbe. The absence

of any filtering arrangements, at that time, allowed these particles

to enter without hindrance. In working up these materials into

new organic substances the Polyzoa played a part of pre-eminent

importance; particularly since these animals formed so large a

proportion of the entire fauna, The carnivorous animals natur-

ally depended for their supply of food on the organisms which

were able to make use of the microscopic detritus. Kraepelin

accordingly arrived at the perfectly legitimate conclusion that if

the food-supply were to be cut off by the introduction of an

efficient filtering arrangement, the pipe-fauna would before long

die out. The soundness of this conclusion was amply demon-

strated by the cessation of the trouble at Hamburg on the

introduction of the remedy suggested ; while the experience of

Water ‘Engineers in many other places has been of a similar

nature.

The presence of Polyzoa in water-pipes is, however, not an

unmixed evil; since, as has been pointed out by several observers,

if floating organisms exist in the water, the Polyzoa serve the

POLYZOA OF WATERWORKS. 429

purpose of removing many of them. But any good they may do

in this way is more than counterbalanced by other inconveniences.

One of the most obvious of these is the diminution of the calibre

ot the water-pipes, which may take place to such an extent as

seriously to interfere with the circulation. Thus Kemna (99,

p- 46) mentions a case which had come under his own observation

of a pipe 60 cm. (about 24 inches) in diameter, which was en-

circled, on its inner side, by a layer of Polyzoa 15 cm. (6 inches)

in thickness. It has further been found by Water Engineers

who have been confronted with this difficulty that, as the Polyzoa

disintegrate, masses of the tubes are torn off from the walls of

the pipes and pass into the circulation. They thus pass to the

smaller pipes of the domestic supply, where they give endless

trouble by choking the ball-taps and strainers of the meters to

such an extent as to stop the supply of water.

These are, however, by no means the only troubles which

result from the presence of Polyzoa and other organisms in the

water-supply. It is the habit of Freshwater Polyzoa, in these

latitudes at least and in the state of nature, to die towards the

approach of winter. There is evidence that even in the sheltered

conditions under which they live in water-pipes protected from

the weather, they are not entirely insensible to seasons; although

it may be noted that Kraepelin (85, p. 12) points out that many

of the constituents of the pipe-fauna were found, in the depth of

winter, at a time when the water containing or ganisms living in

exposed places was covered with ice and snow, in a condition

differing but slightly from that in which he had found them in

the summer. But he adds that the polypides of Pluwmatella were

all dead at this time (December), although in March the hiber-

nacula of Paludicella were splitting (as a preliminary to germina-

tion), and the statoblast-“‘ embryos” of Wredericella were already

escaping from their shells. Owing to the death of the polypides

of the Polyzoa, at the approach of winter, a certain amount of

decomposition of organic material naturally occurs. ‘This sets up

further troubles, partly by tainting the water and partly by pro-

viding material which encourages the growth of Bacteria. The

tainting of the water, which may be due to a combination of

these two factors, is sometimes very serious. Thus De Vries (90,

p- 38) records a case of this kind in which even horses would

not drink the water.

It is not my purpose to discuss the flora of the filtering- beds

and water-pipes—a subject of great biological inter est. with

which I do not feel myself competent to deal—but it is neces-

sary to refer to an illuminating study of this question which was

made, in 1890, by Hugo De Vries, in the memoir just referred

to. ‘The town of Rotterdam is supplied by water from the Maas.

The waterworks were established in 1874, and at first gave no

trouble. In the spring of 1877 serious difficulties began to be

experienced as the result of a sudden invasion of “ Tron-

Bacteria,” and in particular of a form described by De Vries

430 DR. S. F. HARMER ON THE

as Crenothrix Kihniana (=C. polyspora Cohn). A scientific

Commission was appointed to investigate the matter; and the

memoir cited was the result. The Bacteria in question, which

grow associated in long filaments, flourish in the presence of iron

in solution in the water. Under these conditions, and when pro-

vided with the requisite organic matter, they deposit the iron, in

an insoluble form, in the sheath which forms the outer part of

the filament. Where they are present in sufficient numbers

they then flake off the inside of the pipes in masses which pass

into the circulation and are delivered with the water into the

domestic supply, where they cause great annoyance. This result

may become a serious calamity by rendering the water extremely

unpleasant for domestic use, and completely unusable for many

industrial purposes (De Vries, $0, p. 9). The remains of animals

and plants which have died in the pipes are said to provide

the organic matter which is required by the Iron-Bacteria. The

growth of these organisms is commonly associated with the

formation of a hard deposit on the inner sides of the pipes;

and this may give rise to serious trouble by reducing the flow

of water.

Tam indebted to my colleague Mr. A. Gepp, of the Botanical

Department of the British Museum, for calling my attention to

some of the special points of interest in connexion with Tron-

Bacteria. A good general account of the subject may be found

in De Vries’ memoir and in the works of Lafar (98, 04), while

an extensive bibliography is given on pp. 90-92 of one of

Kemna’s memoirs (05). One of the special points of interest in

this connexion is the theory, first brought forward by Winogradsky,

that the Iron-Bacteria take up iron in solution in the ferrous state

and obtain their energy by oxidizing it to the ferric state, which

appears in the form of iron-rust deposited in the sheaths of the

filaments. The theory has been criticised by some of the more

recent workers, as by Molisch, who claims to have kept Tvon-

Bacteria for several generations in iron-free solutions. It has

been suggested that the deposit of iron-rust in the sheath of

/renothri« and other forms is of the same general nature as the

appearance of silica in the cell-wall of a Diatom: and that it has

not the physiological significance which was attributed to it by

Winogradsky. But whatever the nature of the process may be,

the peculiar property possessed by these Bacteria of depositing

iron in the ferric state is a matter of the utmost practical

importance to Water Engineers.

The removal of the pipe-fauna, or the establishment of condi-

tions under which a fauna of this kind cannot effect an entrance,

appears to be of great importance in preventing the growth of

Tron-Bacteria. As De Vries pointed out, these organisms require

organic matter, which is supplied to them in abundance, in un-

filtered water, by the disintegration of the constituents of the

pipe-fauna. An instructive case, which appears to be of this

nature, is discussed by Kemna in his account of the troubles

POLYZOA OF WATERWORKS. 431

which have been experienced at Liverpool. In order to increase

the supply of this city a dam was constructed at Vyrny, in

Wales. Filters were installed at Oswestry, thirty kilometres

from the dam ; and the water was then carried for about twice

that distance to Liverpool. In this latter part of the system no

difficulties have been experienced ; but in the part which carries

the water from Lake Vyrny to the filters, constant trouble has

been caused by the formation of a mucilaginous layer which

seriously reduces the calibre of the pipes. In the deposit thus

formed Kemna states (05, p. 78) that he has found Crenothrix,

one of the Iron-Bacteria.

Although De Vries was principally concerned with the study

of Crenothrix, he did not restrict himself to the consideration

of the flora of the system. He points out (pp. 34, 35), as the

result of his observations, that the fauna in the Rotterdam

water-pipes is closely similar to that which had been found by

Kraepelin at Hamburg. Among the organisms thus recorded,

Cordylophora, Paludicella, and two forms of “ Pliumatella”’ [one

of which may have been F’redericella| were important con-

stituents. Here, as at Hamburg, deficient filtration was the

real cause of the trouble. In parts of the system where the

conditions were favourable, owing to faulty construction of the

filters, some of the animals occurred in astonishing profusion.

I may refer specially to certain Crustacea (Asellus and Gam-

marus), which were found in almost incredible numbers. This

is brought out sufficiently clearly by the observation recorded on

p- 65 of the memoir cited, that the bottom of one of the filters

wes found to be coated by a continuous layer, of almost the

thickness of a finger, of the feces of Aselluws, while the animals

themselves occurred in ‘‘ countless millions ” !

-De Vries’ memoir gives references to other cases of water-

works which have at various times suffered seriously from the

occurrence of Iron-Bacteria or an unduly rich fauna, or both

combined. Among these [ may mention the “ Berlin water-

calamity,” in 1878 onwards, which was completely cured by the

introduction of sand-filtration in 1883 (pp. 9, 14, 15, 37);

Philadelphia (p. 35); and Lille (p. 37). Further information

with regard to cases which have occurred in America is given by

Whipple (10), who mentions Boston, where Polyzoa, especially

Fredericella and Plumatella, were the most important constituents

of the fauna. Other cases recorded are Brooklyn (p. 169) and

Henderson, N.C. (p. 123). In addition to the genera of Polyzoa

which have already been mentioned as occurring in drinking-

water, Whipple gives Pectinatella and Cristatella.

There is some reason to believe that the Corporation of Man-

chester has suffered from troubles of a similar nature to those

experienced at Hamburg and Rotterdam. Hickson (04, p. 675)

has stated that, two years before the date of his Address, the

mains had become partially blocked by the growth of an organism

which was described as a “ moss,” but which Hickson had the

43% DR. S. F. HARMER ON THE

opportunity of inspecting and thus satisfied himself that it was

really composed of Polyzoa. 700 tons of this “moss” are said

to have been removed from the mains by an expensive and

laborious process ; but the subject was not investigated biologi-

cally, and I have been unable to ascertain anything more with

regard to the subsequent history of this case.

At Paris (cf. Kemna, 05, p. 26) the Molluscan fauna of the

water-pipes was investigated by Locard, who recorded 44 species,

belonging to 13 genera.

At Ypres, in Belgium (Kemna, 05, p. 63), a quantity of shells,

with masses of Polyzoa and innumerable Infusoria, could easily be

obtained by opening a hydrant in the streets.

Mr. 8S. C. Chapman, M.Inst.C.E., the Water Engineer to the

Borough of Torquay, to whose kindness I am indebted for much

information on the subject of waterworks, is preparing a paper in

which he proposes to give an account of the troubles which have

been experienced in the district supplied by-the system under his

own charge, and of the result of the enquiries which he has made

of the Engineers of other waterworks, at home and abroad. ‘This

paper is expected to appear in the ‘Transactions of the Institu-

tion of Water Engineers.’ I believe Lam right in saying that it

will contain valuable information tending to prove the correct-

ness of the view, which is held by nearly everyone who has studied

the subject, that the troubles which are caused by the occurrence

of a rich pipe-fauna can be got rid of by the introduction of an

efficient system of filtration.

Enough has been said to show that the occurrence of conditions

favourable to the growth of Polyzoa and other organisms in the

pipes of a water-supply is not a matter to be treated lightly. A

study of the literature shows, moreover, that the Biology of

waterworks is a subject of great complexity and of very special

scientific interest. An excellent general account of the problems

involved has been given by Dr. A. Kemna (99, 05), who speaks

with special authority since, in addition to his scientifie qualifica-

tions, he has had the advantage of being the Manager of an

important Water-Company which supplies the town of Antwerp.

As the subject is one which has not attracted much attention in

Zoological literature, | may perhaps be excused for adding a few

remarks, based principally on Kemna’s account (05), on the

subject of sand-filtration, which 1s so closely connected with the

practical side of the question.

The introduction of sand-filters was due to James Simpson,

Engineer to the Chelsea Company, who gave evidence on the

subject before a Royal Commission on the Water-Supply of

London in 1827-1828, and before a Committee of the House

of Commons in 1828. Kemna comments (p. 109) on the fact

that although Simpson’s views on the theory of sand-filtration

were not altogether sound, he nevertheless succeeded at his

first attempt in introducing a system which has not been

materially improved on since.

POLYZOA OF WATERWORKS 433

In this system the water is introduced into shallow filtering-

beds, which are of large superficial area and have a bottom of

sand through which the water passes before it is admitted to the

pipes. The efficiency of the filter does not depend on a mechanical

straining action of the sand, but is a biological process, princi-

pally confined to the surface-layer of the sand. In this layer

grows an abundant flora, consisting of Alge and Bacteria; and

the beneficial action is the result mainly of the presence of

Diatoms. By means of the slimy or gelatinous investment

formed by these organisms, the intervals between the sand-

grains are filled up, thus increasing the effectiveness of the

filter as a strainer; while the surface-tilm of living organisms has

a very marked effect in reducing the number of Bacteria, and

probably affects the character of the water in other ways. As

the filter continues in action, the number of organisms in its

floor increases and the surface-film grows in thickness in a corre-

sponding proportion. The filtering surface also tends to become

choked by the accumulation of detritus which has been strained

off, until from these causes it ceases to transmit the water so fast

as is necessary. The filter then has to be put out of action and

the surface-film must be removed. The film has to be allowed to

re-form for two or three days before the filter is in a condition

to be used once more. .

The organisms which are of importance in the filtering process

undergo seasonal variations. Thus at one time of year Kemna

finds that one species of Diatom is dominant, while at another

time the effectiveness of the layer depends on some other species

or group of species. The filter thus needs constant observation,

since it may readily get out of working order. An interesting

cease is recorded by Kemna (99, p. 48), who found that Dipterous

larve (Chironomus) may have an important practical effect. In

the observation alluded to, the surface-film was inhabited by

numerous larve, which were living in tubes constructed out

of sand-grains. So long as the larve inhabited their tubes the

filter worked well. But when they left the surface-film in order

to undergo their metamorphosis, their empty tubes formed a

number of perforations which were naturally not closed by

Diatoms, ete., and the filter then ceased to work properly.

Information that the insects were undergoing their meta-

morphosis was given by the appearance of Swallows circling

rouud the filters in order to feed on the emerging Diptera. On

noticing this indication the filter was put out of use and given

three days’ rest, during which the holes left by the emergence

of the Chironomus became closed by a new growth of the

surface-film.

The practical importance of sand-filtration is well brought out

by the facts which are cited by Kemna in the two memoirs

already referred to. One of the most instructive cases is the

history of the Hamburg Waterworks during the years which

preceded Kraepelin’s investigation of the pipe-fauna. In 1842

434 DR. S. F. HARMER ON THE

a disastrous fire destroyed a large part of the city of Hamburg,

including the buildings which contained the machinery of several

distributions of water. The Municipality thereupon took ovei —

the supply of water; and it is satisfactory to learn that the

‘English Engineer, Lindley, who is described as the author of the

pr oject, insisted on having an installation of sand-filters. Partly

owing to financial Aieniniog! and partly in consequence of the

objections raised by the manufacturers and vendors of domestic

filters, Lindley’s recommendation was not followed for many

years. At last, however, the difficulties were overcome, and the

construction of filters, which were to be completed in 1894, had

commenced in 1890. Shortly after this date the water taken

from the Elbe became contaminated by a temporary encampment

of Russian emigrants, and cholera broke out in August 1892

before the completion of the new works. This epidemic, which

claimed 10,000 victims, was clearly traced to the defect in the

water-supply, since the neighbouring town of Altona, which took

its water from the same source but filtered it before using it,

remained unaffected. The cases which actually occurred in

Altona were traced to the use of Hamburg water ; while blocks

of houses in Hamburg which were supplied from Altona escaped

infection. Mr. 8. C. Chapman informs me that he has heard

from Professor Kraepelin that the introduction of filtration,

“through extensive gravel-beds,” was almost immediately suc-

cessful at Hamburg; the pipe-fauna dying from starvation in

the course of a few months after the installation of the filters.

It might naturally be anticipated that the conditions in the

reservoirs and filter-beds (in the layer of water above the sand)

would be favourable for the growth of numerous freshwater

organisms. ‘These are the sources from which the pipes may

become colonised, if not protected by filters; and it is thus of

interest to notice that the facts are in accordance with expectation.

A case of this kind is recorded by Hickson (04, p. 675), where

great trouble was caused in the supply of the town of Burnley

by the occurrence of enormous numbers of Limnea peregra

in the Hecknest reservoir. An even more striking instance is

described by Kemna (99, pp. 47, 48) from his own experience

in the Antwerp reservoirs, in 1896. In this case the trouble

was caused by the extraordinary multiplication of Cladocera,

which were so numerous that it was necessary to strain all the

water passing to the filters through screens of wire-gauze. So

great was the number of these small Crustacea that it was found

necessary to keep six men constantly employed, night and day,

in changing the strainers. The quantity of Crustacea thus

removed was estimated by Kemna at ten tons at least.

Another striking case, due, however, to defective construction

of the filters, is tho one recorded by De Vries (90, pp. 56 e¢ seq.),

where enormous numbers of Gammarus and Asellus occurred in

the filter-beds and other parts of the system.

It need not be a matter for surprise if a particular system should

be found to have been free from trouble due to Polyzoa and other

POLYZOA OF WATERWORKS, 435

organisms for a long period, and then to have become suddenly

infected. It is at least theoretically possible that Polyzoa are

absent in certain waterworks merely because no statoblast or

embryo happens to have found access to the system; even though

there may be nothing to prevent these organisms from flourishing

if they succeed in effecting an entrance. The means of distribu-

tion of freshwater organisms have frequently been discussed.

Darwin gives important information on the subject in ‘The

Origin of Species’, Ch. xiii :—with particular reference to dis-

persal by water-birds and insects. Other facts of the same

nature are recorded by H. Wallis Kew, in ‘The Dispersal of

Shells’ (Internat. Sci. Series, 1893). A short general discussion

of the subject was contributed by O. Zacharias to the ‘ Biologisches

Centralblatt,’ ix. Nos. 3 and 4, April 1889. Reference may also

be made to B. M. Puschkarew’s recent paper, ‘‘ Ueber die Ver-

breitung der Siisswasserprotozoen durch die Luft” (Arch. f. Pro-

tistenkunde, xxvii. Heft 3, 1913, p. 323).

It can hardly be doubted that the method by which Polyzoa

most commonly colonize a new place is by means of their stato-

blasts, which from their habit of floating at the surface are

extremely likely to adhere to some part of the lees, beak, or

feathers of a wading or swimming bird, or to some part of a

water-insect. De Guerne (88), struck by the occurrence in the

Azores of a lacustrine fauna almost entirely composed of European

types, was led to examine the material adhering to the feet of

Ducks; and on one occasion (p. 297) he discovered a half stato-

blast of a Plumatella on the web of the foot of a Wild Duck.

Similar positive information with regard to the dispersal of fresh -

water Polyzoa has been published by Garbini, in a paper which I

have only been able to consult by means of the notices given in

Vol. ii. (p. 195) of the ‘ Zoologisches Centralblatt’, and (by Matz-

dorff) in the “Jahresbericht wber die Bryozoen fiir 1897 und

1898” in the ‘Archiv fiir Naturgeschichte, lxi. Jahrg., ii. Bd.

Heft 3, p. 181. It appears from the latter notice that stato-

blasts of a Plumatella were found attached to the beak of a

Heron (“ Reiher”’).

IJ. Occurrence of Polyzoa in British Waterworks.

I turn now to the consideration of the cases which have come

under my own observation in this country. I am able to record

five cases, in very different parts of Great Britain, in all of which

serious trouble has been experienced. I commence by giving

some account of the occurrence of the organisms, reserving the

question of the determination of the specimens to Part IIT. of

this paper. The cases are as follows :—

(1) Torquay AnD Newron Apzot.

Towards the end of September 1910, I received samples of a

Plumatella (P. emarginata, var. muscosa) from Mr. Samuel

Proc. Zoou. Soc.—1913, No. XXX, 30

436 DR. S. F. HARMER ON THE

C. Chapman, M.Inst.C.E., Water Engineer to the Borough of

Torquay. I am greatly indebted to Mr. Chapman for the

trouble he has taken in giving me the fullest information with

regard to this particular occurrence, and in calling my attention

to similar instances in other places.

The Torquay case is one of the usual kind, in which no efficient

system of filtration bad been in operation. In spite of this fact

no trouble had been given by the growth of Polyzoa or other

organisms, in sufficient quantity to give rise to anxiety, during

fifty years. The apparently sudden invasion may perhaps have

been due to the fact that the Polyzoa in question had not

previously effected an entrance into the system. The Pluwmatella

was first observed in pipes which had been in use a little more

than two years. The pipes were at the bottom of a steep hill, on

a trunk-main which supplied a large part of Newton Abbot, and

where the flow of water was constant and of considerable velocity.

The Polyzoon was soon traced in every direction all over the

system, both at. Torquay and at Newton Abbot. It quickly

began to give rise to serious inconvenience, by becoming detached

from the walls of the pipes and by being carried to the ball-taps

and the strainers of meters, which became blocked and were thus

thrown out of work. The trouble became specially acute imme-

diately after a frost in the early part of 1912. This is entirely

in accordance with expectation ; the Plwmatella which had been

growing on the walls of the pipes having no doubt broken up

during the cold weather, its branches having then passed into the

flow of water and so having given rise to the choked taps and

meters which were reported in large numbers. Jn one case men-

tioned to me by Mr. Chapman, eleven houses in one block of

buildings were without water owing to this cause. It need

hardly be remarked that the well-known habit of dying at the

end of the year gives these freshwater Polyzoa special advantages

in distributing themselves over a wide area of a system. It is

only necessary for statoblasts or fragments of branches containing

them to remain attached to some irregularity on the inside of a

pipe to make it practically certain that some of them will have

the opportunity of germinating in the next period of warm

weather.

No complete examination of the pipe-fauna has been made in

the Torquay Waterworks, but a few other animals have been

observed in them. One of the most conspicuous of these is a

Freshwater Sponge, which has been found growing on the insides

of some of the pipes, and has been determined by Mr. R. Kirk-

patrick as Hphydatia fluviatilis. In some of the samples there

were large quantities of a Dipterous larva, of a bright red colour.

These have been determined by Mr. F. W. Edwards as a species

of Chironomus. The majority of these larve, and perhaps nearly

all of them, could hardly have succeeded in completing their

metamorphosis; and it seems probable that, as in the case of the

Dipterous larve recorded by Kraepelin (85, p. 11), the trans-

POLYZOA OF WATERWORKS. 435

formation into the imago, or at least the emergence of the adult

form into the air, could not take place except by some chance

combination of circumstances which would hardly occur very

often. It may be noted that Kraepelin, in commenting (p. 11)

on the absence of Chironomus-larve from the underground pipes

of the Hamburg system, points out that, according to his obser-

vations, these larvee are the most dangerous enemies of freshwater

Polyzoa, whose delicate branches they tear to pieces. This might

have an important bearing on the distiibution of the Polyzoa,

since, even in the most active period of their growth, branches

containing statoblasts would be liberated by the Chironomus-

larve, and some of them would come to rest in places where a

new growth of the Polyzoa would be started.

In at least one of the samples, considerable numbers of a small

freshwater Oligochete were found; but [ am not able to deter-

mine it from the preserved specimens in my possession.

From the descriptions which have been given to me by

Mr. Chapman, there can be no doubt of the occurrence in the

Torquay system of small Gasteropods (perhaps Dithynia or

Limnea, and almost certainly Planerbis), and of Asellus, Gam-

marus, Cladocera, and Copepoda. 1 have not had the opportunity

of examining specimens of any of these animals.

I have no certain evidence of the occurrence of [ron-Bacteria,

but the appearances suggest that these occurred. In some places

the pipes were found to be coated with a thick black layer of

“‘ neaty substance,” cemented by iron-compounds. My colleague

Mr. A. Gepp, who has kindly examined one of the samples, tells

me that its condition is not such as to make the demonstration

of Jron-Bacteria certain, but from his experience of other similar

cases, he has little doubt but that these had really been present.

It may be remarked that the Torquay authorities have now

instituted a filtration-system ; and, judging from the experience

of other water companies, it may be presumed that the trouble

from which they have suffered will soon be a thing of the past.

(2) Lorp Powis’s StycHEe Esrare (at Lydbury, North Shrop-

shire, near Market Drayton).

In May 1912 I received information, through Dr. Augustus

Voelcker, of the Royal Agricultural Society, of the occurrence of

a Polyzoon which had been giving serious trouble near Market

Drayton. I later received specimens, in the fresh condition,

through the kindness of Mr. hk. H. Newill, Lord Powis’s agent.

The specimens 1n question were well-grown examples of Pluma-

tella fungosa, var. coralloides. In this case a feed-tank received

its water from a small open pool in which the Polyzoon was

proved to be growing. ‘The water passed to the tank through a

strainer, vahrcr did no more than filter off the coarse detritus.

From the tank the water passed down an incline, in an under-

ground pipe, to a ram-chamber situated about thirty-three yards

30*

438 DR. S. F. HARMER ON THE

away. The ram isa pump which raises spring-water from another

source for use in the house and other buildings. From infor-

mation supplied by Mr. Newill, the Plumatella is found in the

open pool (from which the sample examined was obtained), in

the copper strainer at the beginning of the pipe from the pool to

the tank, in the same pipe, in the feed-tank, in the pipe leading

from the tank to the ram, and in the ram-house. ‘The middle

parts of the pipes were said to be less affected than the parts

near the tank and the ram. Considerable trouble had been

experienced in 1911; and it is expressly pomted out that the

Polyzoon ceases to grow about August. In May 1912, when I

received the sample, the Polyzoon was in vigorous growth. ‘The

nature of the trouble was of the same kind as at Torquay, and

consisted in the choking of the pipes and in the blocking of the

ram to such an extent that it ceased to work. Unsuccessful

attempts had been made to check the growth by the use of copper

sulphate. With the Polyzoon were found multitudes of bright

red “‘ worms,” of a greenish hue when young and dark red when

mature. They are described as being marked by dark bands, as

growing to a length of one-half to three-quarters of an inch, and

of having the habit of changing their skins, “ which float on the

water.” From these indications there is a high probability that

the animals in question were Dipterous larve, probably of some

species of Chironomus, as in the case of the Diptera found at

Torquay.

Mr. Newill informs me that sand-filters will be installed in the

spring of the present year.

(3) Bartey,

The first intimation of trouble at Batley was received from the

Director of the Royal Botanic Gardens, Kew ; but I later obtained

samples from Mr. J. C. Barrowclough, the Waterworks Engineer

at Batley, who kindly gave me information on the subject. The

Polyzoa sent belonged to Fredericella sultana, and they had been

responsible for trouble of the usual kind—-namely, blocking of

the meters, strainers, etc. In the sample examined the meter

had been fixed on Aug. 19, 1912, and was removed on Oct. 16

completely choked by branches of the Mredericella. The supply

of these waterworks is obtained from a moorland gathering

ground on the Pennine Chain, “having large impounding

reservoirs, but no form of filters.” The trouble had been known

to exist for a long time, but it had been ascribed to peat and

other deposits. An examination of the trunk-main had recently

been made on account of its reduced capacity through incrus-

tation. The large quantity of iron-compounds found in a dry

sample of this inerustation points to the presence of Ivon-

Bacteria, though no certain evidence of their presence was dis-

covered. It is stated by Mr. Barrowclough that the autumn is

the time when special trouble is experienced. As in the other

POLYZOA OF WATERWORKS. 439

cases, this is no doubt due to the fact that that time of year

is the period at which the tubes of Polyzoa naturally disin-

tegrate, with the result that branches are set free and choke

the meters.

(4)

At the end of October 1912 I received, through Mr. 8. C.

Chapman, a sample from an English waterworks installation, the

locality of which I am not permitted to mention. It consisted,

in the main, of a mass of tubes of Paludicella articulata. The

polypides had already disappeared owing to the lateness of the

season; but the sample was a particularly beautiful specimen,

covered with a profusion of hibernacula or winter-buds, which

were developed on every branch. Among the branches of the

Paludicella there were disintegrated fragments of Fredericella

sultana. Mr. Chapman informs me that serious trouble had been

experienced for some time at the locality in question, although

the nature of the organisms causing it had not been recognized.

(5) ABERDEEN.

At the end of March, in the present year, I received a sample,

very similar to the last one, from the Aberdeen Waterworks,

again through the kindness of Mr. Chapman. This consisted

principally of a tangled mass of Paludicella articulata, composed

of empty zoccia bearing numerous hibernacula. The sample

also included a few pieces of Spongilla lacustris (determined by

Mr. R. Kirkpatrick), crowded with gemmules, a few young speci-

mens of Limnea peregra (determined by Mr. G. C. Robson), and

some Dipterous larve. ‘These were referred by Mr. F. W.

Edwards to a species of Chironomus, which was, however, not

the same as the species found at Torquay. There were also a

number of tubes, formed of cemented mud, which appear to

have belonged to the Chironomus-larvee.

The following case may also be mentioned, because, although

not illustrating the occurrence of Polyzoa in waterworks, it has

an obvious bearing on the importance of filtration :—

(6) Hampron-on-T'HAMES.

On May 22, 1912, ‘The Daily Mail’ published a short account

of the occurrence of enormous numbers of ‘ freshwater mussels ”

in a 36-inch water main at Hampton-on-Thames. It was stated

that 90 tons of these animals had been removed from a length of

a quarter of a mile of this main, the diameter of which had been

reduced by them from 36 inches to 9 inches. The “ mussels ”

were examined by Mr. G. C. Robson, who found them to belong

to Dreissensia polymorpha, a Lamellibranch which has been

recorded in the pipes of the Hamburg water-supply and elsewhere.

Mr. H. B. W. Stent, A.M.Inst.C.E., the Resident Engineer of

440 DR. S. F. HARMER ON THE

the Metropolitan Water Board, has kindly given me some addi-

tional information with regard to this case. Without certifying

the exactness of the above recorded computation of the weight

of the Molluses removed, he admits the existence of large

numbers of these animals in the main. He emphasizes the fact,

however, that no trouble of this kind has been experienced in

the water which has been admitted to the pipes after filtration.

The Molluses are found only in the mains which are used for

conveying water from the Thames to the storage reservoirs, or

from the reservoirs to the filter-beds. The importance of filteri ing

the water before admitting it to the pipes is admirably br ought

out by this case, since the Molluscs in question have been unable

to get past the barrier of the filter-beds.

The consideration of the above cases is sufficient to show that

the risks which might be caused by the entry of animal life into

the pipes of a water-system are not confined to any particular

part of the country. The cases which I have examined come

from Aberdeen in the north to Torquay in the south-west. It

is obvious that if the water is unfiltered and thus contains micro-

scopic organisms in sufficient quantity, the chance introduction

of a few Polyzoon statoblasts or embryos might be followed by a

growth in the pipes sufticient to interfere very seriously with the

circulation, and ultimately to choke the taps. It is important

to notice, as has been so well shown by Kraepelin, that the

Polyzoa play a leading part in promoting the growth of animal

life in the pipes. They are specially adapted by the disposition of

their ciliated tentacles to catch the diatomsand other microscopic

organisms on which they habitually feed, or, as Kemna* has

expressed it, they are ‘‘ micr: ophagous.” They serve in their turn

as the food of larger animals ; and, unless they prepared the food

in this way, it is hardly likely that animals such as Eels would be

capable of flourishing in the pipes of a water-system. The great

quantity of Polyzoa found at Hamburg, Torquay, and elsewhere

justifies the conclusion that in any discussion of this question

the occurrence of Polyzoa is a fact of cardinal importance.

Sponges are perhaps the organisms which rank next in im-

portance to the Polyzoa in this respect.

The practical and theoretical interest of this subject has

received very little attention in this country. Professor S. J.

Hickson, in his Presidential Addiess to Section D of the South-

port Meeting of the British Association (04, p. 675), has, how-

ever, used words to which I can completely subscribe. He states

“that our ignorance of the general balance of animal and vege-

table life in the large reservoirs is profound, and that a systematic

inquiry conducted by competent persons would most certainly

lead to knowledge which would be of great scientific importance,

and in the long run remunerative to the community.”

* Kemna, A., “ Octaenemus, Une Ascidie mégophage,” Ann. Soc. roy. Zool. et

Malacol. de Belgique, xli. 1906, p. 57

POLYZOA OF WATERWORKS. 44]

It is a curious coincidence that, at the same Meeting of the

British Association at which Professor Hickson’s address was

delivered, the Presidential Address of Mr. Charles Hawksley to

Section G (Engineering) dealt specially with water-supply and

the construction of waterworks. In this address it is stated

(Report, p. 767) that “most waters obtained from gathering

grounds are much improved by filtration”; and the mode of

construction of sand-filters is indicated. But there is no refer-

ence to the importance of the filter as a means of depriving the

water of the microscopic organisms which would otherwise supply

nourishment to Polyzoa and other ‘“ microphagous” animals

capable of flourishing in the pipes if they are supplied with

appropriate food im sufficient quantity.

I am indebted to Mr. 8S. C. Chapman for having put at my

disposal the result of an extensive enquiry which he has made as

to the occurrence of Polyzoa and other animals in the waterworks

of this country. As he proposes to publish the result of his

enquiries J will not attempt to anticipate him; but he returns

show that organisms of the kind indicated occur more frequently

than has generally been supposed, and that the occurrence of

trouble due to their presence can usually be traced to the want

of an efficient system of filtration.

III. Species of Polyzoa found in British Waterworks.

The species of Polyzoa which I have had the opportunity of

examining may be referred to the following four species :—

(1) Paludicella articulata Khrb.

(2) Fredericella sultana Blamenbach.

(3) Plumatella fungosa Pall., var. coralloides Allman.

(4) Plumatella emarginata Allman, var. muscesa, Kraepelin.

I subjoin a few notes on the synonymy of the species and on

the specimens which form the subject of this paper.

(1) PatupiceLia arTicuLATA Khrb. (PI. LXIT. figs. 1-10.)

Alcyonella articulata Ehrenberg, ‘‘Symbole Physice,” “ Ani-

malia Evertebrata,” Dec. 1, ‘‘ Phytozoa Polypi,” 1831, fol. a.

Paludicella Ehrenbergii Dumortier & Van Beneden, ‘“ Hist.

Nat. des Pol. Comp. d’eau douce,” 11. Partie, Nouv. Mém. Acad.

Bruxelles, xvi. 1843, p. 38.

Although Allman, in his classical monograph (56) on the

Freshwater Polyzoa, accepted the specific name ehrenbergi—and

has been followed in this respect by many other writers—the

reasons given by Dumortier and Van Beneden for rejecting

Ehrenberg’s name are not in accordance with modern principles

of zoological nomenclature. They state as their motive for intro-

ducing a new name that “ articulata” refers to a generic character,

and therefore cannot be used as a specific name. The invalidity

449, DR. S. F. HARMER ON THE

of this argument has not escaped the notice of some other

writers.

The first specimens examined were received from a waterworks

system (No. 4, above), which does not permit itself to be men-

tioned by name, at the end of October 1912. They were remark-

able for the extraordinary freedom with which hibernacula were

developed. In correlation with the lateness of the season, the

polypides had degenerated, and the zocecia were represented

merely by their empty ectocyst. Nearly every zocecium, through-

out the whole of the material, bears one or two hibernacula im

the place of the ordimary lateral daughter-zoecia; and, if it

happens to be at the end of a branch, a terminal hibernaculum

as well. These hibernacula (Pl. LXII. figs. 1-10) are, almost

without exception, of an elongated fusiform shape, although

varying much in length and, to a jess extent, in breadth. They

thus differ strikingly from most of the figures of these structures

which have previously been published.

The hibernacula were originally described, under the name of

‘“hybernacles,” by Dumortier and Van Beneden, in the memoir

cited above (p. 51, pl.i. fig. 1”; pl. ii. figs. 24— 35). Although

shown as replacing levherell bale, and ahhenefione in the same

position as in the specimens which have come under my notice,

they are described and figured as short bodies, not more than

about one-fifth of the length of the zoecia. They are pointed at

their free end, and were said to be always strongly compressed.

In colour they were greyish black.

Kraepelin, who states (87, p. 76) that the hibernacula do not

appear to have been found by any observer since Van Beneden,

describes and figures them (pl. iv. fig. 117) as swollen and ellip=

soid or irregular, and states that iley are yellowish in colour,

and that they have a considerable amount of calcareous matter

in their cuticle. Several of the specimens figured by Kraepelin

resemble the hibernacula described by Van Beneden in their

form, but one or two of them are fusiform, although not so elon-

gated as most of the hibernacula which have come under my own

observation, Some of them remain attached to the substratum

when the zocecia, which have meanwhile lost their contents, dis-

integrate and break up. In the second part of his monograph

(92, p. 61) Kraepelin refers to the irregular form generally

possessed by the hibernacula. He co the occurrence in

them of an elongated polypide-bud (pl. v. fig. 163), which is

formed early in their development, in re ee for germination

in the ensuing spring; and he mentions the yolk- like material

by which the polypide-bud is surrounded. He was unable to

discover the mode of formation of the hibernacula.

Levinsen (94, p. 85, pl. viii. figs 24-26) gives figures of the

hibernacula which are much like those of Kraepelin.

Wesenberg-Lund ($6, pp. 321, 363, xxiv; pl. iv, figs. 44, 45)

found the hibernacula in large numbers i in October. They were

all of the same form and colour as in Kvraepelin’s description,

POLYZOA OF WATERWORKS. 445

and were attached to the substratum, none being discovered on

the erect shoots. Their mode of development was not ascertained.

Chirica (04, pp. 7, 8, sep.) does not describe the development

of the hibernacula, but records one or two interesting observa-

tions. He obtained them, not only at the beginning of the

winter, but also attached to branches of dead zocecia, floating at

the surface of the water in April, and he regards this as the

means by which the species is distributed in the spring. He

adds that colonies which have resulted from the germination of

hibernacula develop ovaries and testes at once (cf. also Kraepelin,

87, p. 86).

The hibernacula of the material examined by myself are, as a

rule, very different in form from those which have been described

by Dumortier and Van Beneden, Kraepelin, Levinsen, and

Wesenberg-Lund. Although varying much in their proportions,

the great majority have an elongated fusiform shape, as shown

in the figures on Pl. LXII. It does not, however, seem to me

necessary to conclude that there is any specific difference between

the form examined by me and those of which hibernacula have

previously been described. Although Paludicella is a light-

shunning organism (Allman, 56, pp. ae rhis 5), these specimens,

obtained from the interior of the pipes of a waterworks system,

must have been growing in absolute darkness, and under condi-

tions which were otherwise different from the normal habitat of

the species. It would thus not be surprising if the hibernacula

were found to show some differences from those growing in a

normal environment. The conclusion that the specimens under

consideration should be referred to P. articulata is confirmed by

the fact that a single branch observed bore two hibernacula (one

of them shown in fig. 1) which have the form described by

Dumortier and Van Beneden.

It remains to be seen, however, whether the hibernacula

usually developed by Paludicella in this country are typically

spindle-shaped or not. Mr. C. F. Rousselet has kindly allowed

me to examine specimens of hibernacula, collected in Norfolk

by Mr. H. E. Hurrell, in his own collection. These hibernacula

are all of the general forin described below, although they are on

the average rather less elongated than the specimens which have

come under my own observation. Some of them have germinated,

in the manner described by Dumortier and Van Beneden and

by Kraepelin. The distal end of the hibernaculum splits into

two valves, in order to allow of the escape of the tissues of the

young zocecium which grows out of the hibernaculum. It is of

some interest to notice, in connexion with what is said below,

that the stall of the hibernaculum remains unsplit during the

process of germination.

Mr. Rousselet informs me that he has never seen hibernacula

of any other form than the spindle-shape here described.

The hibernacula observed by me are white in colour, although

some of them show signs of becoming darker. The material was

444 DR. S. F. HARMER ON THE

obtained towards the end of October, and therefore soon after the

hibernacula had been formed. It is not improbable that they

would all have become darker as the season progressed. All were

growing on erect branches of the colony, and most of them were

quite free, although some show the habit which has been described

by other observers of attaching themselves to foreign objects ;

in this case zocecia, or even other hibernacula, of the Paludicella

itself, or zocecia of the Fredericella with which the species was

associated. Allusion has already been made to the fact that one,

two, or even three hibernacula are borne on nearly every zocecium.

They vary much in length and to a less extent in breadth.

Leaving out of account the two hibernacula of which one is

shown in fig. 1, they are nearly all spindle-shaped, although

occasionally showing some irregularity of form, and vary in

length from 608 pw (fig. 4) to about 1968. They are usually

pointed at their free end, the broadest part being as a rule beyond

the middle of their length. ‘The proximal stalk-like portion is

generally slender, but varies in breadth from about 72 p to 96 p.

Measured at their widest part, the width varies from 144 p to

160. They are usually developed in the position occupied by

either the lateral or the terminal daughter-zocecia ; but, as shown

in fig. 7, the hibernaculum may originate more proximally than

the position normally occupied by a lateral bud; and the bases

of the lateral hibernacula are not always opposite one another

(fig. 8). Most of the hibernacula contain an elongated polypide-

bud, in agreement with Kraepelin’s statement (92), of the form

which also occurs in the development of the polypide-buds of

ordinary zoecia. J have not observed any calcareous matter in

their wall.

It has often been suggested that the statoblasts of Phylolacto-

lemata may have been evolved from structures like the hiber-

nacula of Paludicella (cf. Kraepelin, 87, pp. 163, 167; 92, p. 62).

But while the mode of origin of the statoblasts, from the funi-

culus, is well understood in its general outlines, I have not been

able to discover any account of the development of the hiber-

naeula, The material at present under discussion enables me to

make a contribution to this subject, although the facts could have

been interpreted with more certainty if the specimens had been

collected slightly earlier in the year. There is, however, enough

evidence to show that the hibernaculum is not a modified terminal

yoecium, as is assumed by Braem (90, p. 112) in criticizing

Kraepelin’s opinion, but is something which is left when the

distal part of a zocecium has been thrown off. The evidence of

figs. 2, 3, 9, 10 seems to be unmistakable on this point. In these

cases the hibernaculum is well formed, with its thick cuticle and

inclusions of yolk-like material. But although its stalk appears

to be constituted, in some cases at least, by the proximal part of

the zocecium, the distal end of the hibernaculum lies freely in the

cavity of the zocecium, which is prolonged in its normal form

beyond it. The interpretation which naturally suggests itself is

POLYZOA OF WATERWORKS. 445

that the formation of the hibernaculum is inaugurated by a con-

traction of the endocyst or body-wall from the original ectocyst

of the zocecium, in the distal part at least, and by the deposition

of a new cuticle over that part of the hibernaculum which lies

freely inside the old ectocyst. In some cases, as in fig. 10,

the hibernaculum has a wall of its own which is distinct

throughout from the original ectocyst. In the specimen shown

in fig. 2 a single zoccium has developed two hibernacula, the

first of which is attached to the parent-zocecium in the position

of a terminal bud, while the second, or distal hibernaculum, lies

freely in the cavity and has no base of attachment. It can

hardly be doubted that the hibernacula represented in figs. 3, 9,

and 10 would have assumed the appearance shown by most of the

hibernacula in this material by the loss of the empty caps formed

by the zocecia within which they have severally been developed.

IT vegard it, therefore, as probable that the hibernaculum owes

its terminal position to the loss of the distal part of its own

zocecium, which, as shown in fig. 10, is not necessarily the

terminal individual of a branch.

There are, however, certain appearances, shown in figs. 6-9,

that are in need of explanation. In these cases most of the

ectocyst of the zocecium which precedes the definitive hiber-

naculum has split into two valves, in the manner characteristic

of a germinating hibernaculum. Like the other zocecia of this

material, these zowcia are destitute of cellular contents ; the only

living tissues left in the colonies being those which constitute the

hibernacula. It is important to consider whether the bivalve

arrangement indicates that a process of germination has actually

taken place. In view of the close resemblance between these

zoecia and a hibernaculum germinating after the end of the

winter and arrived at the stage of having completed the formation

of the first regenerated zocecium, it seems legitimate to assume

that germination of a hiber naculum has Gcourred in spite of the

fact that the material was collected in the autumn. It would

appear that a new zocecium has been completely formed, since

the remains of its tubular orifice are present. But the develop-

ment of a new colony has stopped short with the formation of a

single zocecium, and a second process of hibernaculum-develop-

ment has taken place, resulting in the definitive winter-buds.

The living material of the regenerated zocecium has passed into

the hibernaculum, or hibernacula, which have been formed from

it, though there is no evidence to show whether the transference

of the material took place en masse or by the more gradual process

by which a bud is ordinarily supplied with its living contents.

It is perhaps not impossible that no polypide was formed by the

regenerated zocecium, but that on the splitting of the hiber-

naculum which preceded it the contents of that structure passed

bodily into the newly-formed hibernaculum. Some such trans-

ference of cellular material may be indicated by fig. 10, where

each of the hibernacula is seen to traverse the septum which

44.6 DR. S. F. HARMER ON THE

occurs between two zowcia. In fig. 8 the definitive hibernaculum

may be proved by measurement to be of exactly the same length

as the hibernaculum which is indicated by the split valves in the

wall of the preceding zoccium. The material of the first hiber-

naculum would thas! just fill the cavity of the second one. In

fig. 9, however, the second hibernaculum is distinctly smaller

than the first one; while in fig. 7, what is assumed to have been

the first hibernaculum has had to provide the material for two

new hibernacula. Further observations on material collected at

a more favourable season will be requir ed to settle these points.

The great majority of the zowcia in a Paludicella-colony give

rise to a pair of lateral buds, situated opposite one another and in

a position which has a definite relation to the orifice of the parent-

zoecium. It may be significant that in all the cases shown in

figs. 6-9 there is some abnormality in this respect. In figs. 6

and 9 only one of the lateral zoccia is indicated; while in fies, 7

and 8, where two are present, they are not opposite one another,

and in fig. 7 one of them is much nearer the proximal end of the

parent- Zo0ecitim than is normally the case. I have observed

similar irregularities in other zocecia of the same kind which have

not been figured; and these cases perhaps support the view that

ger mination of the hibernacula first formed has taken place, since

it is well known that in young Polyzoon colonies the zocecia first

developed frequently fail to give rise to the full number of buds

which are formed by the zocecia in an actively growing branch.

If the explanation above suggested is correct, the germination

of the first set of hibernacula may perhaps hese heen. induced by

the occurrence of warm weather succeeding a cold period, which

induced the formation of the first set. But the explanation i in

question is not the only one which can be imagined; and it is

not inconceivable that as the time of hibernaculum- formation

approaches, some of the zocecia make an abortive attempt to

develop hibernacula, and that the bivalve arrangement seen in

figs. 6-9 implies an unsuccessful effort of this nature.

In the majority of the zowcia of this material the definitive

hibernacula have been produced at the first attempt, and the

zoecia Which respectively precede them show no appearance of a

bivalved arrangement. But in all cases the zocecia are empty of

cellular contents, and there has obviously been a centrifugal

passage of the living material of the colonies into the hibernacula,

which are finally left as the outermost members of the series.

It remains to be considered whether these observations have

any bearing on the question of the homology, if any corres-

pondence really exists, between the hibernacula of Paludicella

and the statoblasts of Phylactolemata. It has frequently been

maintained that statoblasts have been evolved from structures

more or less resembling the hibernacula of Paludicella, which on

this view is regarded as related to the ancestral form from which

the Phylactolemata have been derived. This argument has been

specially developed by Kraepelin (87, pp. 163, 167; 92, p. 62),

POLYZOA OF WATERWORKS, 447

who based his conclusions on the general similarity of the hiber-

nacula to statoblasts, particularly those of Hredericella in which

no annulus is present. He did not, however, bring forward any

evidence to show how the hibernacula are developed. Braem

(90, p. 112) has disputed Kraepelin’s conclusions, basing his

objection largely on the assumption that the hibernacula are

complete, specially modified, terminal individuals of the colony.

On this assumption there would be a considerable difference

between them and the statoblasts, which are essentially buds

produced internally within the body-cavity of a zoccium.

Braem’s objection loses some of its force if his assumption as to

their mode of origin can be shown to be incorrect. I think that

the evidence here brought forward is sufficient to render the

conclusion that the hibernaculum is typically an end-bud of the

colony very doubtful. Although the proximal part of its ecto-

cyst is commonly a modification of the corresponding part of the

ectocyst of the parent-zocecinm, cases like those shown in figs. 2

and 10 appear to prove that the hibernaculum may be formed as

an internal bud-like structure, with a complete cuticle of its

own; and there is indeed no very essential difference between

the proximal hibernaculum shown in fig. 10 and the statoblast

ot a Fredericella. I do not, of course, overlook the fact that the

statoblasts of Phylactolemata are developed in a peculiar way

from the funiculus of a polypide. But Paludicella is, in any case,

only distantly related to the Phylactolemata; and the difference

between the winter-buds of the two forms is not, apparently, so

great as has hitherto been assumed by most observers who have

considered the question.

When one considers the facility with which freshwater

organisms, from Protozoa upwards, develop some special cyst-

like coat which enables them to survive the winter or periods of

drought, one must no doubt be cautious in assuming that the

homology suggested is a probable one. But my observations

seem to show that the hibernaculum of Paludicella, like the

statoblasts of Phylactolemata, is a special form of winter-bud

which is developed inside the parent zocecium out of a mass of

living tissue which becomes surrounded by a chitinous coat ; and

that thus the two kinds of winter-buds are morphologically com-

parable. In view of this consideration it does not appear to me

impossible that the hibernaculum and the statoblast represent

two different conditions which have been evolved from a common

starting-point.

The specimens of this species received from Aberdeen (No. 5,

above) agree closely with those just described. Numerous hiber-

nacula of an elongated type are present, although they are

perhaps more variable in length than in the other sample.

Some of them are even longer than the longest described above.

Paludicella articulata las previously been recorded as an

inhabitant of the pipes of waterworks, both at Hamburg

(Kraepelin, 85, p. 6) and at Rotterdam (De Vries, 90, p. 25).

448 DR. S. F. HARMER ON THE

(2) FReDERICELLA suLTANA Blumenbach. (PI. LXIII. figs. 11—

14.)

It is unnecessary for me to discuss the synonymy of this

Species, since there is a general agreement with regard to it

among practically all recent writers. Excellent accounts of the

species are given by Allman (56), Kraepelin (87), and many

others. It was recorded by Kraepelin (85, p. 6) as one of the

commonest forms found in the Hamburg Waterworks system ;

and I suspect that one of the species of “ Pluwmatella” recorded

by De Vries (90) from the Rotterdam Waterworks may have been

really Mredericella.

I have myself found /. sultana in two English waterworks ;

namely the system (No. 4) in which the Paludicella above described

occurred and at Batley. In both cases the material was collected

in October, and the polypides had already degenerated, leaving

empty tubes containing statoblasts. ‘The tubes show the septa

which have been described by Allman (56, p. 112), Kvaepelin (87,

p- 100), and others, at the point where a tube diverges from the

main stem. The statoblasts are developed in both localities with

greater profusion than is sometimes the case in this species.

Kraepelin has commented (pp. 103, 104) on the variability of the

statoblasts of /. sultana. He states that it is hardly possible to

find two alike in the same colony; and that they may be reni-

form, almost quadrangular, or even nearly circular. The average

length is given by him as 430 p, and the average breadth as 300 p.

The length varies between 380 and 570; and the breadth

between 210 p and 370 p.

Figs. 11-14, taken trom the Batley material, show that my

own results are in close agreement with Kraepelin’s. The varia-

tion in form is sufficiently obvious from these figures. The

average length of nine statoblasts from Batley was 444 py, a

measurement almost identical with Kraepelin’s. The extreme

measurements were 416 and 480. The average breadth of

the same nine specimens was 242 »—somewhat less than Kraepe-

lin’s figure,—while the extremes were 200 and 272. Some

were more than twice as long as broad (fig. 14); while the

statoblast shown in fig. 11 is almost circular.

Kraepelin states that the funiculus of Yredericella generally

gives rise to one or two statoblasts, rarely to three. In my own

material, while many of the zocecia contain a single statoblast,

and others contain either two or three, one of the tubes in the

Batley material contains four statoblasts, which certainly belonged

to one polypide. In several instances it can be seen that the

statoblasts are closely adherent to one wall of the tube. In cases

where two tubes are connate, the statoblasts are adherent to the

common wall which separates the two tubes. In the Batley

material the statoblasts are almost black in colour, and a large

proportion of them, although still contained in the zocecia, have

split into their two valves, between which can be seen a mass of

living tissue. This isa clear sign that the statoblasts are ready

POLYZOA OF WATERWORKS, 449

to germinate. Itis well known that the statoblasts in this species

ordinarily germinate without leaving the tubes in which they

have been produced. It need hardly be pointed out that the

state of the winter-buds, here as in the Paludicella described

above, is such as to give every opportunity for the dispersal of

the species throughout the system into which it has effected an

entrance. In the Batley case at least (and probably in the other)

the branches examined were not taken from the places where

they had been growing in the pipes, but had become free by the

disintegration of the colonies at the approach of winter, and had

accumulated in the meters and taps to such an extent as to

block them.

(3) PLUMATELLA FUNGOSA Pallas, var. coraLLormpes Ailman.

(Pl. LXIIT. figs. 22—25.)

I follow Braem (90, p. 5) in regarding Plumatella coralloides of

Allman as a form of P. fungosa. This seems to be indicated by

Allman’s fig. 4 (pl. vii.) of the floating statoblasts. It should,

however, be noted that Annandale, in his recent account of the

Indian species (11, pp. 217, 218), gives P. coralloides as a synonym

of P. fruticosa.

In dealing with representatives of the genus Plumatella one

is confronted with numerous difficulties; since there is no

general agreement with regard to the species which ought to be

recognised. It is worth remembering that Pluwmatella has three

different kinds of reproductive bodies, namely, the embryos

produced by the development of the eggs, the floating statoblasts,

provided with an “annulus” of gas-containing cavities, and the

sessile statoblasts. We are in almost complete ignorance with

regard to the question whether the form of the colony depends

to any extent on the particular kind of reproductive body from

which it has been developed. As Braem has pointed out (90,

p. 8; 11, p. 9), experimental evidence on this subject is urgently

needed ; and it is probable that a well-conceived series of breeding

experiments carried out with statoblasts and embryos of Plwmatella

might resolve some of the difficulties which at present exist.

Braem (90, p. 6) has recorded the important observation that in

cases Where the remains of the statoblast from which a colony

has germinated can be detected at the proximal end of the

colony, the form of that statoblast can sometimes be proved to

be identical with those which are being produced in the tubes of

the same colony. ‘There are reasons for believing, as has been

remarked above, in dealing with Paludicella, that ovaries and

testes are usually produced early in the summer from colonies

which have resulted from the germination of statoblasts or

hibernacula (cf. Kraepelin, 87, p. 85). But it has not been

sufficientiy ascertained how far, if at all, these sexually mature

colonies differ from those produced by statoblasts; nor is it

known whether there are any differences between colonies

450 DR. &. Fe. HARMER ON THE

produced from floating statoblasts and those which have developed

from sessile statoblasts.

It is believed by most recent authorities that Alcyonella is not

generically separable from Plumatella. Alcyonella is character-

ised by its compact form, resulting from the arrangement of its

crowded, more or less parallel, tubes, more or less at right angles

to the substratum. It has been suggested that this habit of

growth is largely, if not entirely, the result of the simultaneous

germination of a number of statoblasts situated close together.

But I am not aware of the existence of any experimental

evidence to show whether a large colony of Alcyonella fungosa

can be produced by the germination of a single statoblast. Some

evidence that the completely parallel tubes of the typical /ungosa-

form owe their arrangement to want of room, is given by the fact

that the peripheral parts of a fwungosa-colony may have the form

of the var. coralloides, which is characterised by a looser habit of

growth.

The conclusion that “ Aleyonella” is merely a form of Pluma-

tclla seems to me to be well founded. The only generic distinction

which can be maintained is the habit of growth; and in this

character P. fungosa var. coralloides is intermediate between

“ Aleyonella” and Plumateila. WKraepelin (87) has gone so far

as to regard P. fumgosa as merely a form of P. repens, which is

termed by him, on insufficient nomenclatorial grounds, P. poly-

morpha. In this conclusion he was at one time criticised by

Braem ($0), who pointed out differences which led him to believe

that P. fungosa is specifically distinct from P. repens. Braem

has more recently (11, p. 9) retracted this view, and he now

regards P. repens and P. fungosa as varieties of one species.

The specimens which may be referred to the P. fungosa type

were sent on May 31, and were received alive, with their

polypides in full vigour. As has been pointed out above, they

were taken from the pool which supplies the system, and I have

not seen specimens from the pipes themselves, where great trouble

has been caused. Mr. R. H. Newill, to whose kindness I owe

the specimens and as much information as he was able to give,

has, however, assured me that the specimens which blocked the

pipes and ram resembled those which had been taken from the

pool supplying the system. There is thus a strong presumption

that the specimens examined belonged to the same species as

those which occurred in the pipes ; a conclusion which 1s rendered

all the more probable by the fact that this form has a very

characteristic habit of growth, which Mr. Newill cannot have

failed to notice.

IT have bad some difficulty in deciding whether the specimens

in question should be regarded as typical examples of P. fungosa

or should be referred to its variety coralloides (= Plumatella

coralloides Allman, cf. Allman, 56, p. 105). In the more central

parts of the masses the tubes have the fungosa-arrangement. In

the more peripheral parts of the material they may be less

POLYZOA OF WATERWORKS. 45]

intimately connected with one another, and are therefore like

the form which has been described as var. coralloides.

Braem (90, p. 4) has stated that the floating statoblasts of

P. fungosa (typical form) have an average length of 410 pu, and

an average breadth of 320 4; the corresponding measurements

for var. coralloides being respectively 380 « and 300. In my

own material the average length of eleven floating statoblasts, im

one series of measurements, was 371 4; and the average breadth

was 240 ». The extreme measurements were :—Length, 328 y and

416 w; breadth, 224 and 2644. In another series of measure-

ments, in which the statoblasts of the compactly arranged tubes

were distinguished from those of the peripheral, more coralloides-

like tubes, the average length was 388 w and the average breadth

rather more than 280 » * for the former; while for the latter the

length was 358 y and the breadth was 266 ». There is thus some

difference between the sizes of the statoblasts in different parts of

the same mass of tubes; the loosely arranged zoccia at the

periphery having somewhat smaller statoblasts than the more

centrally placed ones. But even the largest statoblasts agree

more closely in size with those of var. coralloides (from Braem’s

measurements) than with those of the typical fwngosa-form (as

given by the same author). I refer the specimens, therefore, to

var. coralloides ; and this conclusion is confirmed by the measure-

ments which I have made of a very typical fungosa-form from

a pond at Upper Norwood (Brit. Mus, 85.7.28.5-8), in which

the average length of the floating statoblasts was 431 yu, and the

average breadth 307 «: in close agreement with Braem’s figures.

The floating statoblasts of the material from Styche have the

form shown in fig. 22. They do not vary much in form, although

in some of them one of the sides is more convex than the other.

The annulus leaves a considerable part of the central capsule

exposed on both surfaces of the statoblast. The average ratio of

length to breadth (11 cases) is 1°543: 1,

The sessile statoblasts are considerably larger than the floating

statoblasts. On the attached surface (fig. 23) the central capsule

gives off an irregular peripheral ring of cementing substance,

which forms a vertical ridge by which the statoblast is fixed to

the wall of the tube in which it has been produced. On the free

surface (fig. 25), a saucer-like shape is produced by the develop-

ment of a thin marginal collar. This represents the annulus of

the floating statoblasts, as is shown by its structure.

In some statoblasts the gas-cells are represented by only a small

proportion of the number found ina floating statoblast ; and these

occur in isolated groups, of irregular and obviously inconstant

form (fig. 25). In other eases, as in fig. 24, the annulus forms a

complete band, which is almost as well developed as in the floating

statoblasts.

* The uncertainty about the breadth is due to the fact that the statoblasts

measured were not resting accurately on one of their flatter surfaces.

Proc. Zoon. Soc.—1913, No. XX XI. 31

452 DR. S. F. HARMER ON THE

The sessile statoblasts are very variable in shape and size

(figs. 23-25), but they are all larger than the floating statoblasts

of the same specimens. The largest one found measured 640 pu

by 432 p, giving a ratio of 1:-481:1. <A shorter and broader

one was 560» by 480, with a ratio of 1:166:1. The average

of seven measurements was 554 p by 434, with a ratio of

PATS) Bu

(4) PLumMareLLA EmARGINATA Allman, var. MuscosA Kraepelin.

(Pl. LXITI. figs. 15-20.)

In his well-known work on the German Freshwater Polyzoa,

Kraepelin (87) refers the forms of Plumatelle which have come

under his observation to three species. These are described

respectively as P. princeps, P. polymorpha, and P. punctata.

For the last, with which it is not necessary for me to deal, he

has adopted Hancock’s name. ‘The first two were renamed by

him, on the ground (pp. 118, 119) that the older names, from

which a choice might have been made, have been used in so

many different senses by previous authors that it would only

have introduced additional confusion to have made use of any of

them for the speciesas characterised by him. I do not think it

necessary to follow Kraepelin in this conclusion.

P. princeps and P. polymorpha were distinguished prmcipally

by the form of the floating statoblasts. In P. princeps these are

relatively long and narrow, and the ratio of length to breadth is

said to vary from 1:53: 1 to 2°79: 1, the typical ratio bemg given

as about 18:1. The species thus characterised includes P. emar-

ginata Allman and P. fruticosa Allman.

In P. polymorpha the floating statoblasts are broader in pro-

portion to their length, and the ratio of length to breadth is

described as varying from 1:1: 1 to 1-42:1; the average being

about 1°25: 1. Under this name Kraepelin includes P. repens

and P. fungosa auctt.

The forms of Plumatella with relatively broad statoblasts are

represented in the material which I have received from English

waterworks only by a variety of P. fungosa, which, following

Braem’s earlier opinion, I have treated above as a species distinct

from P. repens. Those with elongated statoblasts are represented

by the material from Torquay, and it thus becomes necessary

to discuss the question of the name which should be used in

describing them.

Braem (90, pp. 9, 10) has brought forward arguments to show

that P. fruticosa and P. emarginata are distinct species ; and he

points out the following differences between them :—

(1) P. fruticosa has relatively slender tubes, which grow into an

erect form; while P. emarginata has broader tubes, which are

more adherent to the substratum. (2) The floating statoblasts of

P. fruticosa are more than twice as long as broad (average ratio :

POLYZOA OF WATERWORKS. 453

length to breadth, 2°497: 1 *), while the annulus does not cover

so much of the central capsule as in the other species. In

P. emarginata, on the other hand, the floating statoblasts are

less than twice as long as broad (average ratio: length to breadth,

1868: 1), while the annulus covers nearly the whole of the

“upper” side, leaving only a small portion of the central capsule

uncovered. (3) In P. fruticosa the sessile statoblasts have a broad

vestigial annulus, and are more than twice as long as broad (ratio,

2-461: 1), while those of P. emarginata resemble the sessile stato-

blasts of P. repens, and have no annulus, while they are less than

twice as long as broad (ratio, 1°312: 1).

Through the courtesy of the authorities of the Hamburg

Museum, I have had the opportunity of examining specimens of

both var. muscosa Kraepelin and var. spongiosa Kraepelin, from

the original material described by that author. I have figured

a floating statoblast (fig. 18) and a sessile statoblast (fig. 20) of

var. muscosa, from the Hamburg material, for comparison with

the Torquay specimens. The measurements which I have made

of the statoblasts from Hamburg agree closely with those given

by Kraepelin, and are as follows :-—

FLOATING STATOBLASTS. SESSILE STATOBLASTS.

Length. | Breadth. ANTOIEY | Length. | Breadth. eweraze

P i ratio, L: B. i it ratio, L: B.

Var. muscosa ...| 482-496 | 240-280 1:817:1 | 384-592 | 272-352 1°456 : 1

Var. spongiosa...) 448-480| 256 1833:1 | 416-496 | 320-384] 1:278:1

Very few floating statoblasts are present in the spongiosa-

material, in which there are a number of sexually produced

embryos.

If Braem’s criticism of Kraepelin’s results is correct, it

follows from these measurements that both varieties, muscosa

and spongiosa, ave to be regarded as forms of P. emarginata;

and in this I am following what I understand to be Braem’s

opinion.

The specimens sent from the Torquay Waterworks by Mr. S.

C. Chapman appear to me to be referable to P. emarginata,

as understood by Braem, although their floating statoblasts are

less elongated than in the Hamburg specimens. I find that these

statoblasts vary in length from 328, to 416 and in breadth

from 224 to 264, the average measurements (11 cases)

being :—Length, 371 1; breadth, 240; ratio, L:B, 1-543:1.

* The ratio is calculated from the measurements given by Braem.

21s

A454 DR. S. F. HARMER ON THE

The sessile statoblasts have the following measurements :—

Length, 432 p-448 yp, average 4424; breadth, 320 p-368 u,

average 336; average ratio, L: B, 1-317: 1.

If Braem is right in separating P. emarginata and P. fruticosa,

the proportions of the sessile statoblasts would alone justify the

reference of the Torquay specimens to P. emarginata rather than

to P. fruticosa. Iam confirmed in the belief that they belong to

the former species by the fact that the annulus covers nearly the

whole of the central capsule on one surface of the statoblast

(figs. 15-17), as shown by Allman (56, pl. vii. fig. 7) in one of his

figures, and by Braem (90, pl. i. fig. 12), who has cailed special

attention to the importance of the character in question.

In Kraepelin’s series of measurements of the forms regarded

by him as belonging to P. princeps (87, pp. 112, 113) the average

ratio of length to breadth is givenas 1°8:1. But he admits a

considerable variation in this respect, the limits given being from

2°79: 1 to 1:53:1. It will be seen that my own measurement of

1:543:1 falls within these extremes, although it is much nearer

Kraepelin’s lower figure. The highest ratio of length to breadth

measured by me is 1°666: 1, and the lowest is 1:°333: 1. The latter

ratio would bring the statoblasts well within the limits given by

Kraepelin for his P. polymorpha, which includes P. repens auctt.

It thus becomes necessary to consider whether the Torquay speci-

mens should be referred to P. repens instead of to P. emarginata.

I have accordingly compared the Torquay statoblasts with the

variety of P. fungosa described above and with what I regard as

a typical form of P. repens, obtained at Cringleford, near Norwich.

It will be seen from the figures that the floating statoblasts of

the Torquay specimens (figs. 15-17) are distinctly longer. in pro-

portion to their breadth, than those of either P. repens (fig. 21)

or P. fungosa var. coralloides (fig. 22), and that the annulus

covers much more of the central capsule. It will be remarked,

moreover, that in the Torquay specimens the annulus extends

over the central capsule further on one side than on the other

side of the statoblast, in agreement with P. emarginata var.

muscosa (fig. 18, from a Hamburg specimen); while in the other

two forms the annulus is about equally developed on both sides.

The floating statoblasts of P. fungosa (fig. 22) are distinctly larger

than those of P. repens (fig. 21). I have not obtained sessile

statoblasts of P. repens for comparison. Kraepelin denies the

existence of this form of statoblast in the typical P. repens, but

Braem (90, p. 6) claims to have found them. In P. fungosa the

sessile statoblasts (figs. 23-25) are very large, and possess a rather

well-developed annulus; while they are smaller, and with a more

vestigial annulus, in the specimens of P. emarginata var. mus-

cosa, both from Hamburg (fig. 20) and from Torquay (fig. 19).

I fully admit that the floating statoblasts of the Torquay

specimens are short as compared with those of more typical

specimens of P, emarginata; but it appears to me that the result

POLYZOA OF WATERWORKS. 455

of the above comparison is to support the conclusion that the

Torquay material should be referred to that species, and I lay

special stress on the unequal development of the annulus on the

two sides of the statoblast. Kraepelin’s var. muscosa was

characterised by him asa luxuriantly branched form of Pluma-

tella in which the branches were not cemented together (in this

respect differing from his var. spongiosa *). The ‘Torquay speci-

mens seem to me to agree closely with those from Hamburg in

their mode of growth, and I have accordingly thought it best to

refer them to the same variety.

Literature cited.

1856. Auuman, G. J.—‘‘ A Monograph of the Freshwater Polyzoa.”

London, Ray Society.

1911. AnnanpALe, N.—“ Freshwater Sponges, Hydroids, and

Polyzoa,” in ‘The Fauna of British India.’ London

(Taylor & Francis).

1890. Brarm, F.—‘‘ Unt. wb. d. Bryozoen des siissen Wassers,”

Bibl. Zool., Bd. i. Heft vi.

1911. Brarm, F.—‘‘ Beitrige zur Kenntnis der Fauna Turkestans:”

vii. “ Bryozoen,” Trav. Soc. Imp. Nat. St.-Pétersbourg,

xlii. Lief. 2, Teil 1.

1904. Currica, C.—‘‘ Notes sur les Bryozoaires de Roumanie,”

Ann. Sci. de ’Université de Jassy, 111. pp. 4-14 (sep.,

pp- 1-11).

1888. p—E GuerRNe, J.—‘“‘ Sur la dissémination des organismes

eau douce par les Palmipédes,” C. r. hebd. des Séances

et Mémoires de la Soc. de Biol. (Paris), 8 ser., v.-Ann.

1888, p. 294.

1890. De Vries, Huco.—‘“ Die Pflanzen und Thiere in den

dunklen Riumen der Rotterdamer Wasserleitung,”

Bericht tb. d. Biolog. Untersuchungen d. Crenothrix-

Commission zu Rotterdam vom Jahre 1887.

1909. Harrmeyver, R.—“ Bryozoen, Moostierchen,” in A. Brauer,

‘ Die Stisswasserfauna Deutschlands,’ Heft 19, p. 49.

1904, Hawxkstey, C.—Presidential Address to Section G (Kngi-

neering), Southport Meeting, 1903, ‘ Report Brit. Assoc.’

pp. 764—, ‘“ Water-Supply.”

1904. Hickson, S. J.—Presidential Address to Section D

(Zoology), Southport Meeting, 1903, ‘Report Brit. Assoc.’

p. 675,

1899. Kemna, A.—“ Ta Biologie du Filtrage au Sable,” Bull.

Soc. Belge Géol. xiii. Ann. 1899, Mémoires, p. 34.

1905. Kemna, A.—‘“ La Biologie des eaux potables,” Ann. Soc.

roy. Zool, et Malacol. de Belgique, xxxix. (1904) p. 9.

* Hartmeyer (09, p.53) describes Kraepelin’s var. spongiosa as a distinct species,

under the name of Plumatella spongiosa.

lor)

DR. S. F. HARMER ON THE

1885. Krarpenin, K.—‘“ Die Fauna der Hamburger Wasser-

leitung,” Abh. Naturwiss. Ver. Hamburg, ix.

1887. Krarpenin, K.—‘“ Die Deutschen Siisswasser-Bryozoen,”

i. “‘ Anatom.-Syst. Teil,” Abh. Naturwiss. Ver. Hamburg,

x. Festschrift.

1892. Krarpetin, K.—“ Die Deutschen Siisswasser-Bryozoen,”

il. “ Kntwickelungsgeschichtlicher Teil,’ Abh. Naturwiss.

Ver. Hamburg, xii.

1898. Larar, F.—‘“ Technical Mycology,” vol. i. London,

Charles Griffin & Co., Ch. xxxiv. pp. 355-362.

1904. Larar, F.—“ Handbuch der Technischen Mycologie,”

A Lief. pp. 193—.

1894, Levinsen, G. M. R.— Zoologica Danica,” 9 Hefte,

‘* Mosdyr.”

1896. Wesenperc-Lunp, C.—‘“ Biologiske Studier over Fersk-

vandsbryozoer,” Vid. Meddel. naturh. Forening Kjoben-

havn, 1896, p. 249.

1910. WuiprLe, G. C.—“ The Microscopy of Drinking-water,”

2nd ed., New York (John Wiley & Sons) and London

(Chapman & Hall).

EXPLANATION OF THE PLATES.

Prate LXII.

Showing hibernacula of Paludicella articulata from the pipes of an English

waterworks-system.

The figures were drawn with a Zeiss A objective, and were then reduced two and a

half diameters. The scale represents hundredths of a millimetre.

Fig. 1. Zocecium bearing a hibernaculum of the type described by Dumortier and

Van Beneden. (Only two hibernacula of this type were found in the

whole of the material examined.)

Fig. 2. A zovecium, accidentally folded, which has developed two hibernacula,

while it bears two other hibernacula (bases shown) in the place of the

ordinary lateral buds. The preceding zovwcium also bears two lateral

hibernacula.

Fig. 3. Zocecium bearing three hibernacula. ‘The distal one is small, and is still

enveloped distally by the remains of the zocecium in which it has been

formed.

Fig. 4. The smallest hibernaculum found. It shows distally the remains of the

zocecium in which it was formed. The elongated polypide-bud is seen in

its interior.

Fig. 5. Zocecium with three hibernacula, each of which shows a polypide-bud.

big. 6. Basal view of a zocecium which shows two split hibernaculum-valves in its

ectocyst.

Fig. 7. A similar zocecium, in basal view, with a terminal and a lateral hiber-

naculum. The latter is developed nearer the proximal end of the

zocecium than is usually the case.

Fig. 8. Side view of a similar zocecium, showing the septa from which lateral

zocecia have broken off. A fully-formed hibernaculum is present distally.

Fig. 9. A similar zoecium, bearing a terminal hibernaculum, which is still enclosed,

distally, in the remains of the zocecium inside which it has been formed.

Fig. 10. A series of three zocecia, with the proximal end of a fourth zocecium.

The proximal zoccium has developed a hibernaculum which has passed

partly through the terminal septum. The third zocecium bears a terminal

hibernaculum, the proximal end of which partly traverses the corre-

sponding septum.

POLYZOA OF WATERWORKS. 457

Prate LXIILI.

Statoblasts of various Phylactoleemata (detailed structure of the annulus not repre-

sented).

The figures were drawn with a Zeiss C objective, and were then reduced two and a

half diameters. The scale represents hundredths of a millimetre.

Figs. 11-14. Fredericella sultana, showing the variable form and size of the stato-

blasts. Batley Waterworks.

Figs. 15-17. Plumatella emarginata var. muscosa; floating statoblasts. Torquay

Waterworks.

Fig. 15. “ Ventral” view ofa young statoblast. The central capsule is represented

by the deeper shading. The line concentric with the outline of the

central capsule, and next inside that line, indicates the extension of

the annulus on the “ventral” side. The inner circle represents the

edge of the annulus on the “dorsal” side, as seen through the central

capsule.

Fig. 16. “ Ventral” view of an old statoblast. The outline of the central

capsule is obscured by the annulus, and is not seen.

Fig. 17. “ Dorsal” view of an old statoblast. The annulus covers nearly the

whole of this surface of the central capsule.

Fig. 18. Plumatella emarginata var. muscosa. Hamburg Waterworks (from

Professor Kraepelin’s material). The annulus covers most of the central

capsule on this surface of the statoblast.

Fig. 19. Plumatella emarginata var. muscosa. Torquay Waterworks. Sessile

statoblast, showing the vestigial annulus; from the unattached surtace.

Vig. 20. Plumatella emarginata vay. muscosa. Hamburg Waterworks (from

Professor Kvaepelin’s material). Sessile statoblast ; from the attached

surface.

Fig. 21. Plumatella repens. Floating statoblast; from a pond, communicating

with the River Yare, at Cringleford, near Norwich.

Figs. 22-25. Plumatella fungosa var. coralloides. Styche Waterworks (from the

peol supplying the system).

Fig. 22. Floating statoblast.

Tig. 23. Sessile statoblast, from the attached surface; showing the cement-

substance by which it is attached to the inner wall of the zocecium.

Fig. 24. Sessile statoblast, with well-developed aanulus, the details ef which

are represented in a small area on the right side of the figure; from

the unattached surface.

Fig. 25. Sessile statoblast, with vestigial annulus, which shows remains of the

gas-containing cavities (only represented at the upper and left parts

of the figure); from the unattached surface.

458 MR. A. W. WA'TERS ON

34, The Marine Fauna of British Hast Africa and Zanzibar,

from Collections made by Cyril Crossland, M.A., B.Sc.,

F.Z.S., in the Years 1901-1902. Bryozoa *—Cheilo-

stomata. By ArtHur Wm. Watemrs, F.L.S8., F.G.S8.f

[Received March 18, 1918: Read April 22, 1913.]

(Plates LXIV.-LXXIII.¢ and Text-figures 79-82.)

INDEX.

Systematic : Page

JEYPAGHID WROOLGy Bs Wo cacect sao soosse cuapsaedscnanapccassoawacanasocso UBL)

(SOMMAORLTOFOOUBOOEISUSy S61 covanvoogadsova cbecooceoauncdaenn concen 2 KokS)

Sig QUIDGOVAR KIOEENG, coagne sooedooad ec degoandeee aoe cebbeacoeanadodepe conceal “Ans

LOAM OPOR, QOMIEEDSUS,; So Mo roaaco oppasocgenvocsoocooaaccocuseconee FAOY/

SISUPIOGHPUR CXOROSONCHPOADEOS, S06 Ws cuodeooodbeaossoodeon coogoonsoouncod | ZAOS)

Sh CAUPROGT AH DEG. SiDo Ts scoossananneancandos oebeaddocsbadtapaccasocunese “ffO)

Scrupocellaria wasinensis, SP. MN. ...0....0ceceeceeneeereceersereee ADO

Vittaticella elegans, var. zanzibariensis, NOV. ......... 22.06 485

Cellaria gracilis, var. tessellata, NOV. .........1..cseseeeeeereese ADS

Chapasinensisespamcres tte esas secace ics cock e aes eee eae So LOD

Osthimosia zanzibariensis, SP. MN. ...... 2-1 sce cee cne eee reeeeesee» 008

SULT LIU GIS D ca Meee eee ho fe se sbi sae Ghana ie aes SEER eee OEE EE are OL

TOGO RRUCEL UREISTIGIOSUS S05 Wo epacas sedaddocabcovssdegcoeagoaeaacosceasa | OIL)

IZQG RHE, QPUGECHOSIS,, SDs Wo sasodatorapesoscoocdacopasouasenncovececes HUIS}

IE, GOLEM WE CAGE Gy TXOe). “ sa3Boc bon ao0c00cn0 605000 son cob HEnocHoe OpIlfs)

Rhynchozoon profundum, var. laminatwin, NOV. ......-..000+. 523

AdeonellopisKenosslam de SD vnlls-c-barcee-pecer cece meee eee eeeeeee 531

Senupocelllarid soiree naecee scares danse ec eee ea eee aes

Catenicelllidias ita asecoe-cesciecs Sos bias. aeewctlene ee uae eee eee CEE ee LOD

(SMICTPOO. connie ones 66 Baa aaa EE RERBe aca adacpisa ncocdocancburinucroenes ior

[SADC SOPQNIG “sat aot aonoce ssa nanese reeeateceneMacdCcoccaddakcoacdsanodeams . taNS)

Calle pon genre acorns) soe aicisasvisins nels stee COs uet epee Soke eaee ee eeae OOO

LEGG C1100 HOR as EL ears dak iols Gold SE He af BERT RS REE OR LO.

TEC PRU Gani aes erie es ces oulspacd i deldee GRESHAM ORE EE RE SE OLD

IW ABKOVAONKOLEEh” “Guocuamoodssuesneooaeemadedeaacaeebocnoae cddonaasscuaccaneo. GPO)

ARCH XOIACD. Groton na scec Oca one aaee a aeRRER Paar ace eomE anne sespeaneannodaconT ar

The collection made by Mr. Cyril Crossland in the neighbour-

hood of Zanzibar contains 76 species or varieties of Cheilostomata,

and all are from shallow water, in fact with the exception of two

are from 10 fathoms or under; so that, for a purely shallow-water

collection, it is a very large one.

Points of Special Interest.

(1) In Stirparia the first zooecium of a tuft has the character

of a primary zocecium (p. 470).

* [In view of the difference of opinion as to whether this Phylum should be

called Bryozoa or Polyzoa (see Proc. Linn. Soc. 1911, p. 61) I have not interfered

with the preference of the author.—KprTor. |

+ Communicated by Cyriu Crosstanp, M.A., B.Sc., F.Z.S.

t+ For explanation of the Plates, see p. 532.

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BRYOZOA FROM ZANZIBAR,

BRYOZOA FROM ZANZIBAR. 459

(2) The nature of the articulation can be used in grouping the

Serupocellaride, the other characters being the ovicells, vibracula

(especially the number at a bifurcation), and the band, perhaps a

spermotheca at the distal end.

(3) The difference in form and position of the ovaria in Bugula

and Serupocellaria gives a generic character, and the same is the

case in other genera (p. 476). The form of the ovaria is likely

to give us considerable assistance in classification, but as the

ovaria pass through various stages of development, time and

caution are requisite to make it fully available.

(4) In Membranipora armata Hasw. there are two mature

polypides in most zocecia, and on one side of the zocecium there is

a long chambered aviculari ium, whereas on the other side there is

a similar long chamber containing a secreting gland (p. 488).

(5) In some species of Schizoporella a bar curving towards the

sinus is seen across the operculum. This is the commencement

of the lower wall of another zocecial layer, and in many cases the

distal walls are seen to extend over the operculum (p. 504).

(6) In Diplodidymia complicata Rss. a small ovum starts im

a small sac hanging down from the opercular region. Both grow

large, and the larva ultimately fills up a large portion of the

zocecium (p. 491).

Some of the species in this collection were also found in

Crossland’s Red Sea collections, and my report thereon is quoted

with a shortened reference *.

This communication may be considered as part of a series

dealing with tropical forms, of which three papers on the Red

Sea etc. have been published ; and I have a small collection from

Cape Verde, also collected by Crossland.

From the locality Wasin about 60 species were procured, and

from Ras Osowamembe about 30 in all.

Since the examination of this Zanzibar collection was com-

menced, Levinsen has published his ‘ Morphological and Systematic

Studies on the Cheilostomous Bryozoa,’ and we have for some time

been anxious to see his ideas on classification developed. He has

given the particulars of much valuable detail examination of the

calcareous parts of the Bryozoa, and doubtless many of the cha-

racters he deals with will receive increased attention, and our

knowledge is much advanced by the wealth of observation.

Unquestionably much of his altered classification will be

adopted ; but if any of us thought that all our troubles would

be ended as soon as Levinsen published his results, we find that

this is far from being the case. We have to examine each new

grouping, and see whether, when other characters are examined,

they uphold suggested alterations; also, do other species of the

groups fall into place? I already see where some modifications

will be required, and no doubt other workers who have carefully

* Journ. Linn. Soc., Zool. vol. xxxi, pp. 123-181, 231-256.

-460 MR. A. W. WATERS ON

examined Levinsen’s great work are coming to similar con-

clusions, and in this way our starting from Levinsen’s standpoint,

and using it as a stepping-stone, should ultimately advance the

correctness of the classification very much. At present the

position remains very difficult, since for a large part we are not

sure what will be accepted.

I do not want to be misunderstood as giving an adverse

criticism, and so would add that the task was too great to be

final, as Levinsen must have fully felt; and we must now be

alive to the fact that, in the future, classifications will to a

large extent be based upon the structure of the soft parts, which

furnish a very large number of characters, some of which are of

great value, while others will not be found very useful. For

every external character there are many internal ones of

importance.

It must be emphasized that, as I have often said*, characters

of great value in one group or family are almost useless in the

next; and all attempts at fixing certain characters as being

of Al importance, others of secondary importance, and so on

down the scale lead to no result, but we must get together our

groups of species based upon as many characters as possible, and

gradually build from them larger divisions; and this process must

be slow, but it will be natural, whereas the attempt to work from

the larger divisions has led to false results.

As a case in point, Levinsen makes great use of the rosette-

plates, which I have found in certain cases to give most useful

results, in other families none at all; and in tabulating Levinsen’s

results we find that the family character is often uni- or multi-

porous, with or without pore-chambers, that is to say, the

character in such cases is of no value for the higher group.

In those families in which we should have been most glad of

help, namely, in Membraniporide, Cribrilinide, Microporide,

Escharide, Smittinide, it is pore-chambers or uni- or multi-

porous rosette-plates, and in eight other families uni- or multi-

porous rosette-plates ; also in genera we find the same range, so

that, while Levinsen’s work in this direction is very valuable,

care is required lest we attach undue importance to the rosette-

plates or any one character.

Where the opercular aperture is not on a level with the frontal

surface of the zocecium there is often a shelf upon which the

distal end of the operculum rests, and Levinsen has shown that

this is a character of some value; but here, again, we must not

expect too much. The value of most of the characters used by

Levinsen, even for the main divisions, are still on trial.

Levinsen has followed 7 Norman in using names given by the

* Page 71, Levinsen expresses these facts as follows:—‘ The same structural

feature in different systematic divisions can have a very different systematic

importance, so that characters which are constant in one genus or family, in

other corresponding divisions are not always constant even within the species.”

+ Since I wrote the above, Canu has sharply criticised Norman’s suggested

alterations of generic names (Rey. Crit. de Paléozool. vol. xvii. p. 49, 1918).

BRYOZOA FROM ZANZIBAR. 461

earlier authors in a sense different from that applied to them

for generations ; and not only that, but generic names now well

established are shifted to be used instead of other well-established

names. Cellaria is to be replaced by Cellularia, a name that has

been used in numerous senses during the lifetime of most of us.

There is no rule obliging us to revert to old names which were

made for genera without any adequate and recognisable deserip-

tion; but even if a rule were being broken, we should continue

to do so in order to stay the appalling confusion caused by this

desire to keep alive a name in its doubtfully supposed original

sense. Cellaria still means for me what it has meant all my

scientific life. I do not accept Lepralia as meaning JMembrani-

porella, and this Levinsen also refuses to do and gives an amended

description of Membraniporella; but I am not prepared to accept

Aspidelectra, and should place melontha under Membraniporella.

Nor is the variously used Hscharoides a satisfactory name, seeing

that it was only given for a subgenus of Cellepora based upon

characters seen to be useless. Discopora is another genus that

should have been dropped, as it was quite insufficiently described

at first ; im fact, the name was given by Lamarck to a Cheilostome

and by Fleming to a Cyclostome, so that it has been employed for

all kinds of unrelated things, as Cellepora, Haloporella, Membrani-

pora, Mucronella, Smittina, Palmicellaria, Diastopora, Lichenopora,

Tubulipora, etc., and has heen var iously used by paleontologists.

It has not been proposed to retain Discopora on account of the

definition, which now tells us nothing, but from what is supposed

to be the first-mentioned species of an incongruous group.

Such changes back to discarded genera add much to the diffi-

culties of those who are closely following all that is written; and

are not these premature and puzzling changes of names keeping

back new workers from entering the thinning ranks? Until we

are sure of the characters, and have enough material to test the

relationships, we must often gather information round species

and genera the names of which we know will ultimately be

changed or disappear.

I have again to thank Mr. Kirkpatrick for allowing me to

make frequent comparison with the British Museum col-

lections. The Plates were mostly drawn before the appearance

of Levinsen’s work, otherwise they would have been slightly

differently arranged. Miss Thornely’s paper, ‘“‘ The Marine

Polyzoa of the Indian ee from H.M.S. Sealark,” Trans.

Linn. Soc. vol. xv. pp. 137-157, has also appeared during the

preparation of my aes It materially increases our knowledge

of the distribution of tropical Bryozoa from moderate depths.

Other groups collected by Mr. Crossland from Zanzibar have

already been described by specialists in the Proceedings of this

Society.

462 MR. A. W. WATERS ON

Table of Distribution from West to Hast.

Aetearangwina cal per seeteseasee ners: 208

» truncata Landsb. .... | 465

Brettia tropica, sp. n. ..................| 405

Synnotum aviculare Pieper ............) 465

3 pembaensis, Sp. N............. 465

is contorta, sp. N................| 466

| Hucratea chelata Li. ..........0.06..++-..| 466

Beania spinigera MacG. ...............) 467

» mirabilis Johnst. ...............,467

Bicellaria chuakensis, sp... ...+..-.. 467

Stirparia exilis MacG. ..................| 468

2” zanzibariensis, Sp. Nn. ...... 469

3 dendrograpta, sp. 0. 470

Bugula neritina var. minima Waters 471

ie robusta MacG. Mates ata AL

Scrupocellaria ferox Raskeeie oe 476

5 cervicornis Busk 477

3 macandrei Busk ...... 477

5 pilosa Aud. ............ 478

wasinensis, Sp. 1. 479

Canda ‘retiformis, Pourtales ......... 479

Catenaria lafontii Aud. ............... 481

diaphana Busk ............... 482

Vittaticella elegans Busk ..| 484

‘ 5 var. zanzibari-

ensis, NOV....... 485

Membranipora savartii Aud. .........) 486

3 armata Hasw. . 486

a catenularia Jameson 488

Farcimia oculata Busk ............... 489

Diplodidymia complicata Rss.......... 490

Chlidonia cordiert Aud. .. 4.92

Cellaria gracilis var. tessellata nov. | 495

if WASINENSIS, SP. ND. oe... ees ee 495

Thairopora mamillaris Lamx..........)497

Steganoporella magnilabris Busk ...|498

Cribrilina radiata Moll. ...............)501

Hippothoa distans MacG................ 501

7 divaricata Lamx. 501

Schizeporella unicornis Johust. 501

ia pertusa TOBE sooosooae 502

* nivea Busk ... 502

montferrandt Aud. ...|506

Gemellipora protusa Yhornely ...... 506

Trypostega venusta Norm. ....| 506

Arthropoma cecilii Aud. ...............|508

Osthimosia zanzibariensis, sp. 0. 508

Lagenipora rota MacG. sscencee|| G10

Haswellia australiensis Hasw. ...... 511

Tubucellaria cereoides var. chua-

kensis Waters ...... | 512

Susiformis VOrb. ......)512

3 zanzibariensis Waters 512

| Atlantic.

+++: +4:

++ 444:

+4:

| British.

Mediterranean.

+ ++

tte:

_(elstanle

<P ap 8

| Indian Ocean,

South.

n.=North.

« .

Sl 6

ei <a

Blel/S\eia

SlE18) 8/2

3/6/25) e/¢

D jlo es |e

[75] |+| N. Pacific

sPauat \-+

as oF ||

-F|...1...| California.

Slices + :

A 4| ( Pacific,

| yr* Barmah.

SEalleaa or

Ge +

ae +|

SSP 14

i albeoe|

ae leo) ae

leeaveeeilictal le Loyalty Isl.

+. | Loyalty Isl.

4) 4+)4+) 4+

J+ }...[ 2]... | 4]

|] +

Hel +|...|+

+ |...) 4F] oe

alive ...|.../+] New Guinea.

alte te

enlleae \ar|

+ +/+)+)4+) Brazil.

+)...) 4]... +

Nita tal

+|+

ocolfaso|| ar |e 4

|...|-.-|F|...|4] Bacific.

aria |

+i) +

In seal Ec +| {ane i

eran “ U S.

ies les auritius

50/4

sar

je. [+ IF |

ial

* This was not mentioned in my Red Sea paper, but I have a piece from a sounding-line

BRYOZOA FROM ZANZIBAR. 463

Table of Distribution from West to East (continued).

: al Lal el

= \\ASunish cul a ald

/ | espa ISH ad

| Vidar S53) |e! |

beige eels ey

5 /-a 5) o |= at es

|g |s2| 2 | @ la ll ilslsis/sin

faolSie2] S |S Sa ales) 2) ele

| @ |els Sy 2 |S |@) oO) S| ce!

(RIA) A |S Oia a &

Smittina trispinosa var. protecta Th. 513 + + | +N. j+

on 3 var. spathulata | | | |

¥ | |

WIEKAC% sanccoeee NSIS ois) soe | saa AB AP iio laellecs ae |

55 iRayadteteh NITES” 56 406 e6ccee case oll (Weasel fact |

3 GIs cececndessabcadnbsoooseoccass5oae| (AI! ihe | | |

| Lepralia feegensis Busk ............... Joe eealiecel| | age che |) ar asfo [ap ifaco, f |fecclacs|| ene WE

A turrita Smitt ...............,516|-++|...| ... san. Warley Sh |

55 wasinensis, Sp. 0. ............/516| | | esi

5 cleidostoma var.inermis Ort. BILZ/ jleeell ben! woldee 44: oe ey le eetaes +|...| N. Pacific.

Petralia japonica Busk ...... ......../518]...]... mY de |) aPasto ar iiar ooo lari

. chuakensis, sp. N. ......-...../518) | | ae (a

a vultur var. armata, nov. ...|518 | Vegi al

? Escharoides occlusa Busk ............|519}...)... 568 ot +t Fi) |

Holoporella columnaris Busk ......... BPE Nosollone| . onc sco. |) Arlo |S llooo P|) 3°

3 aperta Hincks ............ Bye aileell) Mona + | +N. |+]......]-..]...] Cuba.

5 (MOPORGPOS 3006.55 oosn0aca6 }522/+)...) ... er || SPS llocclfaoo"a|ls05) Ar

Microporella ciliata Pallas .............523;)+ +) + del] aeixio. i|arllecs ap| sear

Rhynchozoon Re var. lami- alec

natum, nov. spanaco 50 6a| (Oe) | | |

Retepora hirsuta Busk ..........0...-.. N523 |. + + |

3 producta Busk ...............| 525 |...| SPs) |fasalfasch Fue) far

5 denticulata Busk ............ 526 |...| bee Baa leaalenetiaa: +)...| Amboina,Lifu,

55 Jjermanensis Waters .........,526)}. + ia | Sanduri Isl.

a tubulata Busk Phun dd Ws le 626 |... Re cee we | EN. IE]. + |

12 7 52 oa] | :

Bifaxaria vagans soe Seeaaese 527 |...|...| .-- soe || ap | Gintins SEN

Adeonella platalea Busk . 31/3) scolleooll 00 Beetllh sapoNe a ldeal lace. seu sea| ieee 3

Vesela || | ana South Pacific.

Adeonellopsis crosslandi, sp. n. 531 | awl

| | |

Hinceks records Lepralia striatula from Zanzibar.

AETEA ANGUINA Linneus. (Pl. LXIV. figs. 1 & 2.)

For synonyms see Miss Jelly’s Catalogue, and add :—

Aetea anguina Calvet, “ Bry. Mar. de Cette,” Trav. de l’Inst.

de Zool. de Univ. de Montpellier, ser. 2, mem. 11, p. 8 (1902);

Jullien & Calvet, Bry. prov. des Camp. de l’Hirondelle, p. 122

(1903); Thornely, ‘Ceylon Pearl-Oyster Fisheries,” vol. iv. Suppl.

Rep. xxvi. Polyzoa, p. 108 (1905) ; Robertson, ‘‘ Non-Incr. Bry.,”

Univ. Calif. Pub.. Zool. vol. ii. No. 5, p. 244, pl. iv. figs. 1-4

(1905); Norman, “ Polyzoa of Madeira,” Journ. Linn. Soc., Zool.

vol. xxx. p. 283 (1909); Levinsen, Morph. & Syst. Studies on Cheil.

Bry. p. 93 (1909); Canu, “ Bry. Helv. de /Egypte,” Mém. de l'Inst.

EKeyptien, vol. vi. p. 190 (1912); Osburn, R. C., ‘“‘ The Bry. of the

Wood's Hole Region,” Bull. Fisheries Bureau, vol. xxx. p. 220,

pl. xxi. figs. 14, 14 @ (1912).

In the specimens from Wasin I have seen ovicells in the same

position as those of A. recta, namely, at the top of the tubular

projection at the back. This dorsal ovicell has now been seen

4.64. MR. A. W. WATERS ON

in A. recta from Rapallo* and Naples in the Mediterranean, and

in the ‘ Belgica’ Antarctic material. Miss Robertson describes

ovicells on the front, some distance down, but was in doubt as to

whether the species was a true A. anguina. Whether she really

had another species before her or not must be left uncertain, but

certainly the front position as drawn by her is a very strange and

unexpected one.

Miss Robertson has confirmed what I wrote about the ovaria

occurring in the creeping part, and this seems to be the usual

place ; however, in the present specimens I do not find that the

polypide extends far into this part, as it sometimes does in A. recta

and as Miss Robertson describes and figures in A. angwina.

Smitt, Waters, Jullien, and Robertson have all shown that the

polypide etc. does not entirely live in the tubular prolongation ;

but Jullien, although appreciating the fact, called this part the

peristome or peristomia. Surely the peristome is something

beyond the operculum and is the part where the polypide is only

to be seen when extended; so that this term applied to Aetea is

most unfortunate and misleading. This tubular prolongation

has been called the neck, and the terminal portion the spoon, but

no satisfactory name has been given to the creeping portion, which

is only a part of the zoecium. There are 12 tentacles.

The diaphragm does not make an infold when retracted, like

most of the Cheilostomata; and the appearance of sete, which

has frequently been alluded to, must be caused by a partial

extrusion of the diaphragm. In some respects Aeteaw approaches

the Ctenostome Cylindreciwm, but no Ctenostome has an external

ovicell.

I do not altogether understand what Levinsen f says regarding

the ovicells of this species ; for though the wall of the ovicell is so

thin that the embryo can readily be seen, yet decalcified pre-

parations and sections have been studied. If Levinsen means to

suggest that the sacs containing the ova and embryos are only

accidentally at the termination and might adhere in any position,

then this is not the case, as ] have now seen a large number, perhaps

hundreds, always in exactly the same position, and see no reason

why we should not speak of them as ovicells. One section shows

the zocecial wall bulging out and the ovum partly in this portion,

which is the commencement of the ovicell £.

Loc. Arctic; Atlantic; Mediterranean; Gulf of Manaar;

Zanzibar (Hincks); 8. Africa; Australia; New Zealand, Tas-

mania; Pacific ¢ (Robertson); Tristan da Cunha (‘ Challenger’).

Wasin, Brit. E. Africa, 10 fath. (500 $), collected by Crossland.

Fossil. Upper Tert. Italy (Veviani), Helvetian of Egypt (Canv).

* “ Bryozoa from Rapallo,” Journ. Linn. Soc., Zool. vol. xxvi. p. 5, pl. i. figs. 1-5

1896).

+ Morph. & Syst. Studies on the Cheil. Bry. p. 93 (1909).

{ Since the above was written, Prof. R. C. Osburn has confirmed the existence of

ovicells in the position described, having found numerous such ovicells in specimens

from Fish Hawk Station: see “ Bryozoa of the Wood’s Hole Region,” Bull. of the

Bureau of Fisheries, vol. xxx. Document No. 760, p. 220 (1912).

§ These and similar numbers are Crossland’s registration numbers.

BRYOZOA FROM ZANZIBAR. 465

ABTEA TRUNCATA Landsborough. (Pl. LXIV. fig. 3.)

There are only small fragments from Prison Island, Zanzibar

Channel. The creeping tube or stolon is dotted in just the same

way as the erect tube.

Loc. Arctic; British; Danish; Madeira, Naples, Rapallo,

Cette. Prison Island, Zanzibar Channel (505), 8 fath., collected

by Crossland.

Fossil. Helvetian, Egypt (Canz).

BRETTIA TROPICA, Sp.n. (Pl. UXIV. figs. 4, 5.)

There are only the zocecia figured, and it will be seen that the

species is closely allied to 5. australis Busk, but differs in the

shape of the area, which in B. iropica is about the length of

a zoecium. There are on the dorsal surface the two light disks *

on each side as in B. longa Waters, but in &. tropica the distal

ones are very minute with the proximal one much larger. The

lower zocecium arises from a calcareous knob, from which stolons

spread out. The zocecium is calcareous, and most of the species

of Lrettia are more or less calcareous, but with such small

fragments we cannot know much about its relationships.

I found species of Grettia in the Arctic and one in the Antarctic,

and though the differences are very small it does not seem that the

present form can be placed with any of those already described.

Brettia may have avicularia, but then it has been called Con "0 -

porella Hincks.

Loc. Wasin, Brit. East Africa, 10 fath. (501), collected by

Crossland.

SYNNOTUM AVICULARE Pieper.

Waters, Rep. Sudanese Red Sea, p.-129.

Loe. (additional). Wasin, Brit. EK. Africa, 10 fath, (501); Ras

Osowamembe, 10 fath. (504); Meweni Bay, 6 fath. (510).

SYNNOTUM PEMBAENSIS, sp.n. (Pl. LXIV. figs. 12-15.)

Zoarium with a spreading ramifying stolon, from which several

erect stems arise, just as in Stirparia ete. The stem is about the

same sizeas the creeping stolon, and issmooth for about the length

of two or three zocecia, then there is a short zocecium followed by

the pairs of zocecia.

The zocecia are shorter and stouter than those of Synnotwm

aviculare P. At one side.at the distal end there is a round

pedunculate avicularium, but no sessile avicularium as in JS.

aviculare. The first zocecium of each branch is uniserial,

is also the case in S. aviculare, Notamia bursaria L., Dimetopia,

and Calwellia, whereas in Gemellaria loricata L. there is a pair

of zocecia at each fresh bifurcation. The pairs of zoccia turn

alternately slightly to right and left, and there are radicles from

between the zocecia just as in Synnotwm aviculare. There seem

* There are similar disks in Catenaria and Vittaticella.

466 MR. A. W. WATERS ON

to be several pores in the large distal rosette-plate. There are

10 tentacles.

Loc. Wasin, Brit. E. Africa, 10 fath. (501); Chuaka, Zanzibar,

2 fath. (508); Chaki-Chaki Bay, Pemba Island, near Zanzibar

(517), collected by Crossland.

SYNNOTUM ConTORTA, sp.n. (Pl. LXIV. figs. 16-18.)

Synnotum aviculare Robertson, ‘* Non-Incrusting Bryozoa,”

Univ. of Calif. Publ., Zool. vol. 11. p. 286, pl. xiv. figs. 84, 85

(1905).

The zoarium is coiled up, especially at the end. The branches

of the zoarium dichotomise, and consist of pairs of zocecia back to

back directed alternately in the opposite directions at right

angles.

The zocecia are wide, subtruncate at the top, diminishing regu-

larly to the base, with a sessile avicularium at one or both sides

near the distal end, and there are a few large rounded, pedun-

culated avicularia replacing one of the sessile avicularia, but none

of these are found in the older zocecia; the area is large, occupying

more than two-thirds of the front, and the calcareous layer is much

more solid than in any other of the Gemellaridze examined. The

first zocecia at the bifurcations are single, whereas the next ones

are double, being back to back. There are long radicles from the

side of the zocecia. In the lateral wall there are two rosette-plates

near the distal end.

There are 11 tentacles.

We know Gemellaria loricata L. without any avicularia, S.

pembaensis nov. with terminal pedunculated avicularia, the

present species with sessile avicularia and a few short thick

pedunculated avicularia, and S. aviculare also with both sessile

and peduneulated avicularia. ‘he presence or absence of avi-

cularia is constantly turning out an unsatisfactory generic

character, and I have never felt quite satisfied that a new genus

was required for Synnotwn.

Miss Robertson’s figure shows the zocecia more attenuated

below, but it certainly seeras that this is the species she

described.

Loc. Chuaka, Zanzibar, 2 fath. (508); Chaki-Chaki, Pemba

Bay, near Zanzibar, low water (517); Wasin, Brit. East Africa,

10 fath. (501).

HucrAtTEA CHELATA Linn.

See Miss Jelly’s Catalogue, and add :—

Scruparia chelata WKirchenpauer, Bericht ther die Unter-

suchungs-Fahrt der Pommerania, ‘‘ Bryozoa,” p. 181 (1875).

Eucratea chelata Levinsen, Zool. Danica, p. 42, pl. 1. figs. 8-9 ;

Calvet, Bry. de Cette, p. 12; Robertson, ‘“ Non-Incrusting

Bryozoa,” Univ. of Calif. Publ., Zool. vol. ii. p. 248, pl. v. figs. 7-9

(1905); Barrois, Emb, des Bry. p. 194, pl. xv. figs. 10-12 (1877) ;

BRYOZOA FROM ZANZIBAR. A67

Osburn, “ Bry. of Wood’s Hole Region” Bull. Bur. Fish. vol. xxx.

p. 221, pl. xxi. fig. 15 (1912); Nordgaard, “‘ Die Bry. des West.

Norwegens,” Die Meeresfauna von Bergen, p. 76.

A tew zocecia were seen from Wasin.

Loe. As far north as the Lofoten Islands; Atlantic; British ;

Mediterranean ; California; Australia; S. Africa (4. W. I. coll.).

Wasin, Brit. East Africa, 10 fath. (501), collected by Crossland.

BEANIA SPINIGERA MacGillivray.

Diachoris spirigera MacG, Trans. Roy. Soc. Vict. vol. - iii.

p: 165, pl. uu. fig. 12 (1859); Prod. Zool. Vict. dec. v. p. 32,

pl. xvi. fig. 3; Waters, Ann. Mag. Nat. Hist. ser. 5, vol. xx.

p. 94 (1887).

There are some small specimens from Wasin which correspond

in most particulars, though the avicularia ave materially smaller

than the type, more like those of B. intermedia Hincks. There

are three terminal spines and usually 5-6 delicate lateral spines.

Loe. Victoria (Australia); New South Wales. Wasin, Brit.

Hast Africa, 10 fath. (501), collected by Crossland.

BEANIA MIRABILIS Johnston.

For synonyms see Miss Jelly’s Catalogue, and add :—

Beania mirabilis Hincks, Ann. Mag. Nat. Hist. ser. 3,

vol. vill. p. 36 (1862); op. cit. ser. 5, vol. xiii. p..357 (1884) ;

op. cit., ser. 5, vol. xix. p. 215 (1887); Waters, Journ. Linn, Soc.

Zool. vol. xxvi. p. 17, pl. u. fig. 1 (1896); Jullien & Calvet,

‘ Bry. prov. des Camp. de l’Hirondelle,’ p. 38 (1903) ; Thornely,

Ceylon Pearl-Oyster Fisheries, vol. iv. Suppl. Rep. xxvi. p. 109

(1905): Robertson, “ Non-Incrust. Chil. Bry.” Univ. of Calif.

Publ., Zool. vol. ii. p. 276, pl. xii. figs. 63, 64, & fig, in text (1905).

Loc. Northern; British; French coasts; Atlantic (Jull. &

Calv.); Mediterranean ; Ceylon (7/.); Burmah (H.); Australia ;

Pacific coast of N. America (/od.). Meweni Bay, Zanzibar (510),

collected by Crossland.

Beania has been considered to belong to the Flustride by

Busk, and to Bicellaride by Levinsen, but the large embryo

found by me in &. magellanica* seems to indicate the proba-

bility of the genus standing elsewhere. There are 20-26

tentacles in Seania, whereas in Bicellaride and its allies there

are usually fewer, 12-18. &. magellanica B. has 23-26; B. hirtis-

sima Hell., 20-30; Bb. hyadesi Jull., 20; B. quadricornuta H.,

23 (W.); B. spinigera MacG., 20; 6. mirabilis Johnst., 20.

BICELLARIA CHUAKENSIS, sp.n. (PI. LX VIII. figs. 7, 8.)

The zoarium arises froin a long, erect primary with radicles ;

it frequently anastomoses, forming a colony about 7-8 mm. high.

The primary zoccium has an elongate area with nine short

* Ann. & Mag. Nat. Hist, ser. 8, vol. ix. p. 493 (1912).

Proc. Zoot. Soc.—1913, No. XXXII. 32

468 MR. A. W. WATERS ON

spines, the next zoccium has seven spines, while the younger

zocecia have usually three spines, two outer ones and one inner,

though there may be occasionally two or four spines instead of

three. The area is less than half the length of a zocecium, and

the pedunculate avicularia are placed at about half the height

of the area, whereas in B. ciliata they are much below it. The

avicularia are of moderate size and similar in shape to those of

B. ciliata; however, avicularia do not occur on any of the lower

zocecia, so that, counting from the primary, no avicularia will be

found before about the 18th zocecium.

The pedunculate ovicell, directed laterally as in B. ciliata, is

situated on the inner side near the distal end.

Loc. Taken in tow-net, Chuaka Bay, Zanzibar (515); Wasin,

Brit. E. Africa, 10 fath. (500), on Steganoporella magnilabris ;

Chuaka, 2-3 fath. (612), collected by Crossland.

Srrrparta EXILIs MacGillivray. (Pl. LXVI. figs. 1-3.)

Stirparia ewilis MacG. “ Desc. of New or Little-known Polyzoa,”

pt. xiii. Proc. Roy. Soc. Vict. n.s. vol. 11. p. 107, pl. iv. figs. 1-16

(1890).

In the Wasin specimens the lower part of the stem is buried

in sponge and cannot be completely examined ; the upper part is

annulated for a short space; the rest is smooth, unjomted, and

without any strengthening rods, but at irregular intervals there

are contractions, or sometimes two or three together, with rosette-

plates across the stem in places. Although these stems differ

from those of the other two species found, yet they are divided

up by these contractions into lengths often about equal to those

of S. dendrograpta; however, the growth is much simpler, so

that, perhaps, S. ewilis may ultimately have to be placed in

another genus. Fresh branches are given off at right angles

to the main stem and start from an expanded disk (fig. 3).

The tuft is 5-6 mm. long, and the zocecia face to the outside

of the tuft. There are about 12 tentacles. The area is about

two-thirds of the length of a zoccium ; there are three spines,

or, in parts two; and only a very few, almost globular, avicularia

have been seen (about two in each tuft), and these are short with

apparently a wide mandible rounded at the end. The avicularia

are attached just below the area and there are no ovicells, only

the commencement of one. MacGillivray found neither avicularia

nor ovicells.

The opercular opening is low down. At a bifurcation one

zocecium extends up a short distance on the opposite side of the

bifureation (fig. 1), and this is well shown in the British Museum

specimens of S. exilis MacG., from Port Phillip Heads.

The mounted specimen from the ‘ Challenger,’ named S. glabra,

is S. annulata Mapl., though there are mounts of a stem of a

species which are like those of 8. glabra and S. dendrograpta.

Busk’s figure, however, seems to show S. glabra H., so that we

have a puzzle; but we may be right in coucluding that S. glabra

BRYOZOA FROM ZANZIBAR. 469

and S. annulata were both obtained from the ‘ Challenger’ Station,

off Bahia. In the British Museum ‘Challenger’ specimen there

are six spines to the primary of the tuft, not a number as figured ;

there are no avicularia or ovicells, but there is a central spine in

the same position as the avicularium in S. glabra H.

Loc. Port Phillip Heads (J/acG".). Wasin, Brit. East Africa,

20 fath. (522), collected by Crossland.

STIRPARIA ZANZIBARIENSIS, sp. n. (Pl. LXVIII. figs. 1, 2;

Pl]. LXIX. fig. 14.)

The stem throws out branches which may bear a tuft 6—7 mm.

long, and the internodes of the stem are approximately equal.

The zocecia are alternate and turn partly away from each other,

that is, the central line of the branch is raised so that the zocecia

slope laterally downwards. The zocecium is much wider at the

distal end than below, and the area is rather more than half

the length of a zocecium. The first zocecium of a tuft has 9-11

long spines, usually six on one side and four on the other

(or dorsal side) with the central spine long; then the second,

third, and sometimes fourth zocecium have several spines, whereas

the normal zocecia have usually only one stout spine at the upper

inner angle, though occasionally there is also one at the outer

angle.

There is sometimes an avicularium to the second zocecium, and

this and the subsequent avicularia, which are long and narrow,

are situated close to the base of the zocecium.

There are about 14-15 tentacles.

On the lower part of the stem or stolon the radicles are some-

times replaced by capsules, similar to those described in my paper,

“ Bryozoa from Rapallo,” * and they may be filled with bright

yellow homogeneous contents. Levinsent refers to similar

capsules as occurring in Bugila caliculata Lev.

The ovicells are pedunculate, and there is a calcareous cover

over a part only, not exceeding the half of a globe, so that the

embryo is thereby but slightly protected, sometimes not at all.

The calcareous wall of the ovicell is made of plates deposited

from centres and looking like the shell of a turtle.

The ovaria are central immediately proximal to the cecum,

and the testes fill up the proximal part of the zowecium. No

ovarium in my sections has more than one ovarian cell.

The zoarial growth is similar to that of S. dendrograpta, sp. n.,

with the long nodes as depicted in Pl. LXVI. fig. 4; but the

tufts are longer, and both the stout spinous processes and the

long avicularia are distinguishing characters. There is no line of

chitinous thickening as in S. dendrograpta.

Loc. West Australia, some imperfect specimens in my col-

lection. Chuaka, Zanzibar, 3 fath. (506), collected by Crossland,

* Journ. Linn. Soc., Zool. vol. xxvi. p. 19, woodcut fig. 6 (1896).

+ Morph. & Syst. Studies on Cheil. Bry. p. 102.

32*

ATO MR. A. W. WATERS ON

STIRPARIA DENDROGRAPTA, Sp. n. (Pl. LXVI. figs. 4-9.)

The stems grow from spreading stolons, and at frequent

intervals branches occur which may bear tufts of zocecia or may

produce other branches, and both the original stem and the

branches are divided into segments approximately equal, though

an internode below a tuft is frequently shorter than the others

From the base of most of the internodes there is a radicle or a

pair with frequently a cervicorn grapnel at the end. The colony

may grow to at least 50 mm. long, and the tufts 3-5 mm. long

originate from a zocwcium entirely different from the later zoecia,

having more or less the character of a primary zoccium. When

first described the stem of Stirparia was considered to be the

equivalent of radicles, but this is not the case.

The first zocecium (figs. 4, 6) has the area a little more than

half the length of the zocecium and is surrounded by eight very

long spines, often attaining about four times the length of a

zocecium. ‘The spines of the first zooecium of a tuft are, however,

not bilateral, but are five on one side and three on the other, the

smaller number being on the side from which the next zocecium

grows. The next zocecium has a somewhat similar area, with

about five spines, and the avicularium is near the base of the area,

while the following zocecium approximates to the later zocecia in

having the avicularium somewhat higher than in zoocium no, 2,

though still low down. In subsequent zomwcia they are placed

still higher, their normal position in the older zoecia being at

the distal end on the outside corner. Typically they may be

terminal in the fourth pair of zocecia, or they may continue

lateral until the eighth, and after the appearance of a terminal

one subsequent zocecia are also generally at the corner of the

distal end.

The avicularia are short with a distinct beak.

The branches of the tufts do not form a complete cup as in

S. exilis MacG. and S. zanzibariensis, sp. n., and the zoccia are on

the inside of the cup, whereas in the others they are on the

outside. The branches of the tuft dichotomise, and the spread-

out fan-shaped tuft is 3-5 mm. long, having often ten pairs in

succession.

The zocwcia are alternate and diagonal with the area a little

more than half the length of a zoocium, and the full number of

spines is three long ones at the distal edge, though many of the

lower zowcia may have one and the younger zoecia two spines ;

nor do the same number occur on both sides of a branch, the

zocecia on the outer zoarial side having more spines than those

on the inner; with three on the outer, there is often only one on

the inner side.

The ring-shaped oblique chitinous thickening, to which

Levinsen * refers as occurring in Bicellaria ciliata L., is often

* Morph. & Syst. Studies on Cheil. Bry. p. 101. Levinsen puts B. caliculata

Ley. under Bugula, but his Pl. ui. fig. 1 shows the character of Bicellaria in

having the long tubular proximal part.

BRYOZOA FROM ZANZIBAR. 471

quite distinct on the dorsal surface without being seen on the

anterior surface; in other cases a mark is seen all round

(fig. 4). When put into Kau de Javelle the zowcium often

breaks off at this line, and it is seen that this proximal tubular

part is connected at the base with the lower zocecium through a

rosette-plate. A rosette-plate higher up connects with the next

younger zocecium (fig. 7).

The ovicell is lateral and pedunculate, and there are about

12 tentacles.

Radicles may occur in abundance on the lower zowcia and not

merely from the normal position at the proximal end of the node.

Immediately below the cecum there is a small globular body

which, as it grows, is seen to be the ovarium, but in no case has

more than one ovarian cell been seen. The same thing occurs in

various Cellularide.

In many respects S. dendrograpta resembles S. glabra Hincks,

but that species has the stem internodes long and short alter-

nately and also there is no avicularium at the distal end in

S. glabra. From S. exilis MacG., it differs in the internodes

being approximately equal and in the different character of the

avicularia. 9. eailis has 6-7 spines on the primary zocecium.

The graptolite, Dendrograptus serpens Hopkinson *, has similar

colonies growing on a stout stalk, and the subcolonies and branches

are about the same size as those of the Stirparia ; and some sub-

colonies I collected in Llandrindod Wells of D. ser “pens H., or a

closely allied species, have the branching quite similar to this

Stirparia and might have been an impress of it, but as competent

authorities have found it to be a graptolite, it slows how identical

the growth may be in widely different classes. The name is

given on account of the superficial likeness.

Loc. Chuaka, Zanzibar, 2 fath. (508), collected by Crossland.

There is a specimen of S. dendrograpta from Port Phillip in

the British Museum.

BUGULA NERITINA var. MINIMA Waters.

Waters, “ Rep. Sudanese Red Sea,” p. 136, pl. xi. figs. 4-7,

for syn., and add Thornely, “ Mar. Polyzoa of the Indian Ocean,”

Trans. Linn. Soc., Zool. vol. xv. p. 141.

Loc. N. S. Wales (W.); Red Sea (W.); Cargados Reef,

Providence, 50-78 fath. (7h.). Prison Island, Zanzibar Channel,

8 fath. (505); Ras Osowamembe, Zanzibar Channel, 10—20 fath.

(514); Chuaka, Zanzibar, tow-net (515), collected by Crossland.

Buevza rosusta MacGillivray. (Pl. LXIX. figs. ae aa.

Bugula robusta MacG. Trans. R. Soc. Vict. vol. ix. p. 129

(1868) ; Prod. Zool. Vict. dec. viii. p. 29, pl. Ixxviil. fig. “1 (1883).

Bugula capensis Busk, MSS.

* Hopkinson & C. Lapworth, “On the Graptolites of the Arenig and Llandeilo

Rocks of St. Davids,’ Quart. Journ. Geol. Soc. vol. xxxi. p. 665, pl. xxxvii. fig. 3

(1875).

472 MR. A. W. WATERS ON

There is a specimen from Wasin which has the zoecia a trifle

smaller than the South African and Australian specimens, and

the avicularia are somewhat smaller; however, in a specimen in

my collection, determined by Busk a B. capensis *, there is one

small avicularium, while the rest are large. ‘There is no real

spinous process at the outer angle, but neither do I find more

than a projection in any of my specimens from other localities.

The distal rosette-plates are all close to the basal wall and very

small, so that it is difficult to distinguish them. In some other

Bugule they are similarly situated, though in others they are

spread over the wall. There are no ovicells in this specimen,

but the species has lateral ovicells like those of B. neritina L.,

and the brown colour suggests its belonging to the neritina group.

The primary zoecium is very long and narrow, followed by a

second long one, then a zoceciuin about the ordinary length, after

which the growth is biserial. The first two zocecia remind us of

the segments of the stem of various Stirparie.

Bugula, asa rule, shows no articulation, but B. reticulata B.

is distinctly articulated—that is, at the bifurcation there are

distinct thick articular chitinous tubes.

Other tropical species of Bugula are:-—B. dentata Lamx.,

B. mirabilis B., B. versicolor B., B. reticulata var. unicornis B.,

B. gracilis B., B. neritina var. rubra Thornely, B. neritina var.

tenuata Th., B. neritina var. ramosa Th. These last two in

many particulars resemble B. reticulata Busk. Most of the

tropical species have a very wide distribution.

Loc. Victoria (McG); South Africa (as capensis); Port

Hlizabeth, 8. Africa (A. W. W. coll.). Wasin, Brit. EH. Africa

(501), collected by Crossland.

SCRUPOCELLARIDS.

Levinsen 7 does not consider that Menipea can be divided up,

as If jroneeedl by the character of the jointing, and on p. 133

gives his account of the articulation of “all Bryozoa that oceur

in jointed colonies,” but his account is not exhaustive. Dealing

now with much more material than on the previous occasion,

my suggestion is more fully tested.

The jointing in the Scrupocellaridz varies considerably, giving

useful specific characters, and it is most important that we should

trace it from its simplest to its most complicated condition.

As I have shown, in most articulated Bryozoa the branches

are at first continuous, and the last two or three bifurcations

may show no sign of rupture, which only takes place after

the chitinous tubes have been formed; and in most articulated

Bryozoa the chitinous tube is formed within the calcareous wall,

though in some it may be formed merely within the membranous

* See my remarks, Journ. Linn. Soc., Zool. vol. xxxi. p. 137 (1909).

+ Morph. & Syst. Studies on the Cheil. Bryozoa, p. 183 (1909).

t{ Journ. Linn. Soc., Zool. vol. xxvi. p. 2 (1896).

BRYOZOA FROM ZANZIBAR. 473

wall. Usually, in Scrupocellaride, the inner zocecium of the new

pair is jointed close to the proximal end (Pl. LX VIII. fig. 14),

but the position of the articulation of the outer zocecium varies

considerably, having the part below the articulation much larger

than the similar portion in the inner zowcium. There are three

exceptions to this rule—WMenipea cirrata Hill. & Sol., MM. smitti*

Norm., MW. flagellifera Busk—all three of which are without a

scutum f.

A.—Beginning with what seems to me the simplest form of

articulation, namely, that found in Canda retiformis Pourt.

(Pl. LXIX. fig. 6), though both genus and species have previously

been described as non-articulated in consequence of the chitinous

tube being often entirely covered by the calcareous wall, in which

as yet there is no rupture, so that decalcification is necessary in

order to study the articulation. As mentioned on p. 480, in the

younger branches no articulation is found, but in the older ones

there is a chitinous tube on the inside of one of the two branches

and a similar tube on the opposite side in the next branch, and

so on alternately (fig. 6).

B.—From the simple form of Canda, we pass on to that of

Scrupocellaria with two chitinous tubes, but with the outer

zocecium havine the articular tube near the middle of the

zocecium (Pl. LXITX. fig. 8). As an example, S. jolloisii Aud.

C.—In the following group the chitinous tube in the outer

zowcium 1s very much lower than in B, but is not, however, close

to the proximal end of the zocecium. The articular tubes are

here narrower, more distinct, and often separated as in Menipea

patagonica Busk (Pl. LXIX. fig. 11). As examples, I/. ternata

Ell. & Sol., Bugulopsis peachii Busk (Pl. LXIX. fig. 10), J. occt-

dentalis Trask, MW. portert MacG.

D.—We pass next to a group in which both articular chitinous

tubes are close to the proximal end of the zowcia. I spoke of

the proximal ends of these new zocecia as small chambers, but

now consider that it would be better to compare them with the

“basis rami” (Harmer) of Crisia, and think the designation may

be used here, remembering that they are really the beginnings of

new zocecia. As examples, Menipea buskit W. Th. (Pl. LXIX.

fig. 12), MW. erystallina Gray, M. cervicornis MacG., MM. occi-

dentalis Trask, M. cirrata Ell. & Sol.

* Waters, “ Bryozoa from Franz Josef Land,” Journ. Linn. Soc., Zool. vol. xxviii.

pl. vii. fig. 8 (1900).

+ As I have always noticed the articulation when opportunities have presented

themselves, several have been carefully figured. In my paper on “ Bryozoa from

Rapallo,” Journ. Linn. Soc., Zool. vol. xxvi. pl. i. the position of the polypides in

the zocecia of Scrupocellaria inermis is SONY figs. 11, 12, and op. cit. vol. xxviii.,

the joints of Menipea gracilis, pl. vi. fig. 1 2, Scrupocellaria scabra, fig. 14, and

S. smittii Norm., fig. 8, are shown. In Te last the articulation only occurs

beyond the distal end of the outer zocium. In the present paper the articulation

is shown in Pl. LX VILLI. fig. 14.

A474 MR. A. W. WATERS ON

E.— Lastly, Menipea cyathus Wy. Thomp. has only one chitinous

tube (Pl. LXIX. fig. 13). In Scrupocellaride there is, in some

species near the distal end, a long body which may be folded

back as in S. ferow Aud. (PL. LXVIII. fig. 14), or it may be

very long, extending to the proximal end oF the zoceclum as 1n

M. fla gells fera Busk! There is much to suggest that this functions

as a testis, and that it should be compared with the organ in

Flusira abyssicola Sars, and Cribrilina figularis Johust.

The classificatory groups may now be considered :—

1. Canna. Articulation simple as A, p. 473; ovicell smooth,

imperforate; vibracula have the sete serrate. Two vibracula at

a bifurcation. Levinsen considers that the ovicell is enclosed in

the widened proximal half of the avicularium, whereas I should

say that the avicularium is on the ovicell.

CapereA. Articulation internal tubes, ovicell imperforate,

vibracula with smooth sete.

3. ScRUPOCELLARIA (div. 1). Articulation as B, p. 473 ; ovicell

smooth, imperforate, usually with two vibracula at a bifurcation ;

sete smooth. This includes S. delilii Aud., S. scruposa L.,

S. scabra Van Ben., S. cervicornis Busk, S. macandrei Busk,

S. ornithorhynchus W. Th., S. serupea var. dongolensis Waters.

(S. scabra has sometimes one vibraculum at a bifurcation.)

A, ScrupoceLLARia (div. 2). Articulation as B, p. 473; ovicell

perforated, usually one vibraculum at a bifurcation, sete smooth.

This includes S. bertholettii Aud., S. jolloisii Aud., S. mansueta

Waters, S. reptans L., S. ferox Busk, S. obtecta Haswell, and

probably S. portert MacG., and S. occidentalis Trask.

5. Bueutopsis. Articulation as C, p. 473: ovicell imperforate,

no vibracula. Example, B. peachii Busk. I follow Levinsen in

using this generic name for the present, but have not had the

opportunity of fully studying the genus.

6. Menipra (div. 1). Articulation as D, p. 473; ovicell truly

endozoccial, showing no external difference, no vibracula;

avicularia sometimes suboral and sometimes lateral, short inter-

nodes. ‘This includes J. cirrata Ell. & Sol., M/. crystallina Gray,

M. cervicornis MacG.

7. Munrpza (div. 2). Articulation as H, p. 474. Two zoeecia

in an internode, scutum directed downwards from the distal end

of the zocecium *. Possibly a new genus will have to be made for

this. This includes J. cyathus.

FLABELLARIS. Levinsen leaves in MZenipea species that cannot

be referred to any other of the genera, and puts under it what I

* Specimens of Menipea fuegensis Busk and WM. aculeata Busk have cases of one

zocecium of a new braneh growing from the distal end of the terminal zocecium and

also one from the side of the lower zocecium.

BRYOZOA FROM ZANZIBAR. 475

placed in a new genus Flabellaris*, and I then showed that

Craspedozoum MacGillivray must be united with one of the types

of Menipea Lamx., namely W/. flahellum Hill. & Sol., but it seems

better to keep the name JJenipea for the first species mentioned

by Lamouroux, namely I. civrata Lamx. However, this group

of flabellaris does not seem to belong to the Scrupocellaride

at all, but to the Membraniporide, having a Membraniporidian

oviceli much like that of JZ. lineata L., MW. craticula Alder,

M. unicornis Flem. The species included are /. flabellata Ell. &

Sol., #. (M.) cuspidata Busk (Pl. LXIX. fig. 9), /. trisertata MacG.

(specimens i in my collection have ovicells), /. (‘‘ Craspedozoum ”)

reborata Hincks, F. (C.) ligulatwm Mee F. multiseriata Busk.

fF. roborata and F. ligulatum when broken through have at the

articulation interior tubes like the radicles.

The ovaria of Flabellaris roborata are distal with many ovarian

cells, and one or more grow to a considerable size. Jullien

considered that Menipea must be merged in Scrupocellaria in

consequence of having found one vibraculum on Menipea clausa

Jull.=Serupocellaria marsupiata Busk, and this conclusion he

considered was upheld by the fact that some colonies of S. scabra

Van Ben. have no vibracula, while others have a few or some-

times many. This seemed quite reasonable, and since then the

idea has received further support, as Levinsen has found one

vibraculum on WM. ternata, and, also, he found vibracula on

Menipea benemunita Busk of the ‘ Challenger,’ for which Levinsen

proposed the genus Caberiella. Also, the form and position of

the radicle chamber in Scrupocellaria serrata Waters} suggests

that a recent ancestor had vibracula.

Although these cases prove that the presence of vibracula does

not give a Sharp divisional line between what has been under-

stood as Serupocellaria and Menipea, yet all the species could

scarcely remain in one genus, and separation can be made on

other grounds. The presence of avicularia gives but very limited

assistance in classification, although there are characters in the

avicularia which are very useful; so it is, therefore, not sur-

prising to find that in some species vibracula may be found as

an exception.

Most of what have been called Menipea have an anterior

avicularium immediately below the area or slightly to one side,

though there are some species without any, as S. inermis, nor

are lateral avicularia universal.

Levinsen § considered that Caberea and his Caberiella had the

avicularium divided into two chambers, whereas I was unable to

find two, for while there is a prolongation of the vibracular

chamber, this only seems to be for the groove in which the

* Waters, “On Membraniporide,’ Journ. Linn. Soe., Zool. vol. xxyi. p. 672,

pl. xlvini. figs. 10, 11; xlix. figs. 7-10 (1898).

+ ‘Cap Horn,’ p. 69 (1888), and Bull. Soc. Zool. de France, p. 507 (1882).

~ Report on Red Sea Bryozoa, Journ. Linn. Soc., Zool. vol. xxxi. p. 133, pl. x.

figs. 11-14 (1909).

§ Morph. and Syst. Studies on the Cheil. Bryozoa, p. 134.

476 MR. A. W. WATERS ON

vibracular seta les. Exactly the same thing, though not so

marked, is seen in Scrupocellaria, for in several species the

vibraculum extends beyond the median line, as in S. macandret

Busk, S. incurvata Waters, ete.

As I told Dr. Levinsen that I did not find two chambers,

he kindly sent me some vibracula, skilfully separated, which,

however, only confirmed what I had seen in my own specimens.

In the Scrupocellaride, so far as I have seen, the ovaria are

large, situated near the distal end, and contain several ova which

are developed into large ova before they pass into the ovicell.

On the other hand, in Bugula, Bicellaria, etc., the ovaria are, at

the proximal end, usually very near to the base of the ceecum ;

they are very small with usually two small ova, and when still

extremely small, an ovum passes into the ovicell. It is very

interesting to nal these generic differences in the ovaria, and

undoubtedly the form, size, and position of the ovaria will be

found to furnish useful characters i in many species of Bryozoa.

The direction of evolution of the Scrupocellaridee seems to be

indicated in the articulation, and a comparison of the changes in

this family may help us to understand the Catenicellide better.

SCRUPOCELLARIA FEROX Busk, (Pl. LXVIII. figs. 11-15;

Pl. LXIX. figs. 7, 20.)

Scrupocellaria feroc Busk, B. M. Cat. Mar. Polyzoa, p. 25,

pl. xxu. figs. 1) 2; & 5.

The avicularia vary considerably in size, being largest just

below a bifurcation, and smallest or wanting in the younger

zocecia, The avicularian chamber has the lateral projection to

which I referred and figured in S. mansweta Waters*, from the

Red Sea, and the long dorsal opening of the vibracular chamber

in the older zocecia has a calcareous band across dividing it in

two (see fig. 14). There is one vibraculum to a bifurcation some-

what directed towards the front, as in S. cyclostoma Busk, and

the radiele, which is hooked at the end, is not ringed as in

S. cyclostoma, but there are only a few complete radicles in the

specimens. No ovicells occur on the Zanzibar specimens.

There are about 24 tentacles.

The rosette-plate into the vibracular chamber is at the base of

the chamber and has many pores; as this rosette-plate is not

always very distinctly marked off these pores might be looked

upon as several plates. Hach zocecium has its own lateral wall,

so that when prepared in Hau de Javelle they may separate.

Stained preparations show a band near the distal end bent

back upon itself (fig.-126). The contents are granular, with

hollow places at intervals, and probably the function is the same

as in the bodies? I mentioned in Bugula bicornis Busk ¢, and they

* Journ. Linn. Soc., Zool. vol. xxxi. p. 134, pl. x. fig. 15 (1909).

+ Those on page 474 are compared with somewhat similar structures in

M. flagellifera B., Flustra abyssicola Sars, and Cribrilina figularis Johnst.

~ Résult. du Voyage 8.Y. Belgica, “ Bryozoa,” p. 21, pl. 1. fig. 4 (1904).

BRYOZOA FROM ZANZIBAR. ATT

are probably testes, or connected with the testes, and in all the

specimens prepared these bands are found in all the zocecia, but

there are no traces of ovaria, so it may be that the male organs

are on one colony and the female on another,

In one mount with about 130 zocecia all contain fully developed

polypides, showing that degeneration does not always take place

at such short intervals as has been often stated. There are,

however, large buds developing by the side of the active

polypides.

Loc. Louisiade Archipelago, Bass’s Straits (B.). Prison Island,

Zanzibar Channel (505), 8 fath.; Ras Osowamembe, Zanzibar

Channel, 10 fath. (504 & 514); Wasin, Brit. EK. Africa, 10 fath.

(507), collected by Crossland.

Other species of tropical Scrupocellaria, not however found in

the Crossland collections, are S. delilii Aud. ; S. ornithorhynchus

B.; S. ciliata Aud. ; S. annectans MacG.; S. diadema B.; S. minuta

Kirkp.; S. clypeata Hasw.; S. obtecta Hasw.

SCRUPOCELLARIA CERVICORNIS Busk. (Pl. LXIX. figs. 3, 4.)

Waters, ‘‘ Rep. Sudanese Red Sea,” p. 166.

Loc. add Ras Osowamembe, 10 fath. (504); Wasin, Brit. Kast

Africa, 20 fath. (522), collected by Crossland.

SCRUPOCELLARIA MACANDREI Busk. (Pl. LX VIII. figs. 5, 6.)

Scrupocellaria macandrei Busk, B. M. Cat. Mar. Polyzoa, p. 24,

pl. xxiv. figs. 1-3 ; and add to Miss Jelly’s synonyms :—Haswell,

W.A., “ Polyzoa from the Queensland Coast,” Proc. Linn. Soc.

N. S. Wales, vol. v. p. 37; Philipps, “ Rep. on the Polyzoa,”

Willey’s Zool. Results, pt. iv. p. 442 (1899); Calvet, ‘ Exp. Scient.

du Travailleur et du Talisman,’ p. 375 (1907); Thornely, ‘‘ Mar.

Polyzoa of the Indian Ocean,” Trans. Linn. Soc., Zool. vol. xv.

p-. 140 (1912).

This belongs to the S. scrwpea group. There are three outer

spines and one inner near the peduncle of the scutum. The

outer spines are often very long. The groove of the vibraculum

is continued beyond the vibracular chamber, passing the median

line of the zocecium, in this respect somewhat resembling S. i-

curvata Waters. There are two vibracula at a bifurcation. The

ovicell is smooth, placed somewhat diagonally, and has a clear

space on the front. The vibracular seta is about the length of a

zocecium, smooth but flat in the middle, so that perhaps we may

call it sickle-shaped. The oral aperture is placed diagonally

back, as in Caberea darwin Busk.

Loc. Cape Verde Island, 1070-1150 fath. (B.) & 110-180 met.

(Calv.) & Crossland Expedition ; St. Paul’s Rocks, N. Atlantic

(B.); Coast of Spain (B.) ; Adriatic (Heller) ; Lifu (Phil.) ; Queens-

land (Haswell); Providence, 50-78 fath., Amirante, 29 fath.,

Farquhar Reef, Cargardos, 30 fath., and Seychelles, 34 fath.

478 MR. A. W. WATERS ON

(Ind. Ocean, Thornely). Prison Island, Zanzibar Channel (505),

8 fath., coll. by Crossland.

SCRUPOCELLARIA PILOSA Savigny & Audouin (now Busk).

(Pl. LX VITI. figs. 3, 4.)

Crisia pilose Aud. Description de ’Egypte, Hist. nat. p. 241;

Savigny’s pl. xi. figs. 1,—1,.

Cellularia spatulata dOrb. Pal. Frang., Terr. Cret. p. 50

(1850-52).

Some dried specimens from Wasin, B. E. Africa, seem to be

the species figured by Savigny. The zocecia are narrow, pro-

ducing a wavy appearance, as figured by Savigny. The opesium

has a very narrow border, and the scutum, which varies in shape,

is small and does not nearly cover the aperture. Both the distal

and proximal ends of the seutum are rounded, with the distal end

the larger, and the scutum is without cervicorn marks. There

are two or three oral spines on the outer side and one or two on

the inner, the outer ones, especially the lower one, which is

stouter, are sometimes long, though in the specimens they are

mostly broken off. The vibraculum is small, partly free at the

outside of the zoarium, with the groove extending slightly beyond

the vibracular chamber, but not as much so as in S. macandrei B.

The vibracular sete are smooth and rather longer than a zocecium.

The zoarium at a bifurcation has a medium spine and one vibra-

culum. The lateral avicularium is placed somewhat diagonally,

instead of standing straight out. The radicles are large and are

serrate near the ends.

There are no ovicells on the specimens.

Busk in the ‘Challenger’ Report, p. 24, describes a species

from the southern hemisphere as S. pilosa Sav., but this seems

a doubtful determination, for the shape of the scutum is different

and the vibracular chamber is very wide and large.

Busk speaks of it as the species of Audouin, whereas Audouin

ealls it the species of Lamouroux, and supported the identity on

Lamouroux having presented some of the type to M. Bory de

St. Vincent which was compared. However, it was never the

species of Lamouroux but of Pallas, and while his description

would tally with this species, 1t would equally well cover a large

number of other Scrupocellarie. D’Orbigny gave the name

spatulata to Savigny’s figure, and in the same way he named

many of Savigny’s other figures, although already named by

Audouin, Savigny having given recognisable figures, we may

suppose that Pallas and Lamouroux were dealing with the same

thing, although this can never be known with certainty.

This is much like Scrupocellaria pusilla Smitt, which, however,

has cervicorn markings on the scutum.

The various species now considered might well be ranged round

S. scruped as varieties.

Loc. ¢ Mediterranean (Pallas, etc.). Wasin, Brit. Hast Africa,

10 fath. (500), collected by Crossland.

BRYOZOA FROM ZANZIBAR. 479

SCRUPOCELLARIA WASINENSIS, sp. n. (Pl. LX VIII. figs. 9, 10;

P}, LXIX. tigs. 17-19.)

Zoarium usually with 5 or 7 zocecia im an internode. Zocecia

wide, with the round area occupying more than half the length

of the zoccium:; at the distal end there are three exterior spines

and one interior; the anterior avicularia are medium-sized,

raised, tubular, with a narrow triangular mandible; the lateral

avicularia are very small; the vibracular, together with the radi-

cular chamber, is about half the length of a zocecium, the delicate

vibracular setee are smooth and shghtly longer than a zocecium ;

the vibracular chamber is separated near the base from the

radicular chamber, and the distal end is contracted. There is

one vibraculum at a bifurcation. A few large lateral avicularia

have been seen which are divided at the end, and the mandible

is forked like that of S. serrata Waters *.

The globular ovicell has numerous pores.

The radicles sometimes pass from one branch to another, as in

Caberea retiformis Sim.

There are about 16 tentacles.

The ovaries containing many ova are at the distal end, while

testes in the same zocecia are at the proximal end. The ovaria

in Scrupocellaria contain several ova, but the ovaria in this

species are peculiar, as there are at first a number of large homo-

geneous cells of which usually only one or two show any nucleus

(PL LXIX. fig. 19). The ovaria at this stage show no follicular

wall, but ata later stage, when the yolk-mass of an ovum has

become very large, then the follicular wall is distinct.

In this species, as in other Scrupocellarie, a large ovum passes

into the ovicell, whereas in Bugula the ovaria instead of being

distal are proximal, and a small ovum in a very early stage passes

into the ovicellt. The ovarian cells, which are frequently far

apart, are often surrounded by a protoplasmic network which

passes up to the rosette-plate of the next zocecium (figs. 18, 19).

This species is of about the same size as S. mansueta Waters, but

has four distal spines and a much smaller vibraculum, with sete

more delicate, and about half to one-third the length of the latter

form. The large lateral avicularia are known in S. varians

Hincks, S. serrata Waters, and S. obtecta Haswell, but apparently

are only the ordinary avicularia more developed.

Loc. Wasin, Brit. Hast Africa, 20 fath. (522), collected by

Crossland.

CANDA RETIFORMIS Pourtales. (Pl. LXIX. figs. 1, 2, 6.)

Canda retiformis Pourtales, Bull. Mus. Comp. Zool. Harvard

Coll., I., No. 6, p. 110 (1867); Philipps, “Rep. on Polyzoa,” Willey,

* “ Bryozoa of Sudanese Red Sea,” Journ. Linn. Soc., Zool. vol. xxxi. p. 133, pl. x.

fig. 11.

+ In Bugula there are two small ova in each ovarium, occasionally three, or even

four, small ones; however, in B. murrayana Johnst., now called Dendrobeania

by Levinsen, the ovarium is distal, and the ova in the ovarium grow to a large size,

so that the material differences in the ovaria would alone suggest that murrayang

does not belong to the same genus as B. avicularis, ete.

480 MR. A. W. WATERS ON

Zool. Results, pt. iv. p. 441, pl. xl. fig. 1 (1899); Thornely,

“Mar. Polyzoa of the Indian Ocean,” Trans. Linn. Soc., Zool.

vol. xv. p. 141 (1912).

Caberea retiformis Smitt, “ Floridan Broyzoa,” pt. i. p. 16, pl. v.

figs. 43-46 (1872); Thornely, ‘‘ Ceylon Pearl-Oyster Fisheries,”

vol. iv. Suppl. Rep. xxvi. p. 109 (1905).

2 Canda fossilis Waters, Q. Journ. Geol. Soe. vol. xxxvii. p. 322,

pl. xvi. figs. 51, 52 (1881); MacG. “Tert. Polyzoa of Victoria,”

p- 25, pl. ili. figs. 12-14, Trams. R. Soc. Vict. vol. iv. (1895).

The specimens from Zanzibar have the scutum very narrow,

ointed distally, and rounded proximally, as described by Miss

Philipps, and I follow her in considering them to be the form

described by Smitt, and although allied to C. arachnoides they

seem quite distinct. It occurs in the Atlantic, the Indian Ocean,

and the Loyalty Islands, without any connecting links being

known. Caberea crassimarginata B., of the ‘Challenger,’ and

Scrupocellaria clypeata Haswell, seem to belong to this group.

No anterior avicularium has been described, but sometimes

there is one attached to the inner side of the zocecium, and then

it is usually just below a bifurcation. The avicularian chamber

is wide, and the mandible is triangular. There are two vibracula

at a bifurcation.’

Both Smitt and Levinsen have considered this species as having

no articulation, and as the chitinous tube is often entirely covered

by the calcareous wall in which there is as yet no rupture, it

requires decalcification to study the articulation, which is peculiar

and seems to differ from that of any other species examined.

In the younger branches there is no sign of articulation, but in

the older ones there is one chitinous tube on the inside of one of

the two branches, but not of both, and a point to be noticed is

that when the chitinous tube is on one side, say to the right, it

is in the next branch on the left, in the one following on the

right, and so on alternately, though in a few cases in the older

parts of the colony I have seen a chitinous tube to each branch.

In (. arachnoides Lamx., as I have shown*, there are two

chitinous tubes, one on the inside of each branch. The articula-

tion of CO. tenwis MacG. is somewhat similar.

When decalcified, a chitinous tube is shown in the peduncle of

the scutum, just as is seen in the base of many spines.

Near the base of the vibracular seta there is a projecting

delicate free arch at right angles to the axis of the seta, and in

other species of the Scrupocellaride there seems to be a similar

structure, though not so pronounced.

One of the cross radicle tubes usually connects the two new

branches at a very short distance from the bifurcation, and this

may partly account for rupture at the articulation so seldom

taking place (see fig. 1).

* Ann. Mag. Nat. Hist. ser. 5, vol. xx. p. 89, pl. iv. fig. 7.

BRYOZOA FROM ZANZIBAR. 481

There are about 16 tentacles.

The vibraculum of Canda retifornis is of the same character as

that of Serupocellaria, in which genus some of the vibracular

chambers extend beyond the median line of the zoarium, though

not quite as much so as in the two species of Canda; also the

vibraculum of Caberea has the same general character.

I have several times maintained* that there are material dif-

ferences between avicularia and vibracula, but that the length of

the mandible or seta is of no importance in indicating which they

are. The distinctions are in the chamber and the basal part of

the chitinous organs. While the mandibles of avicularia are

symmetrical and have the closing muscles attached by Tone or

two long tendons, the base of asymmetrical seta of the vibracula

is very complicated with a large number of curiously shaped pro-

tuberances, to some of which the muscles are attached by a

fascia, but without any long tendon, so that instead of there

being two main muscles, there are more attached by a short band

to various parts of the base of the seta. The vibracular base is

very small, so that it is difficult to follow this complicated

mechanism ; the reason for this complication is found in the seta

being movable in all directions, whereas the mandibles of the

avicularia only move in one. The mandibles all have a straight

proximal edge, but this is not the case in the seta; further, the

avicularian mandible works from this straight base either against

the calcareous bar, or, in case this is not complete, then from two

teeth. The universal movement could not take place with a

eross-bar, and none has been found, nor must we ever expect to

find one in vibracula. The vibracular chambers of this group

are different from those of Cupularias, ete., in which the vibracular

chamber takes the place of a zocecium.

Loc. Florida, 68 & 270 fath.; Loyalty Islands (Ph.); Galle,

deep water (7%.); Amirante, 23-29 fath., Saya de Malha, 55 fath.,

Seychelles, 39 fath., Cargados, 30 fath. (Thornely). Wasin, Brit.

East Africa, 10 fath. (501); Prison Island, Zanzibar Channel

(505), 8fath., Ras Osowamembe, Zanzibar Channel, 10 fath. (504),

collected by Crossland.

CATENARIA LAFoNTIT Audouin.

Waters, ‘“ Rep. Sudanese Red Sea,” p. 131.

Near the distal end there are small ovaria with two ovarian

cells.

Loe. Wasin, Brit. Kast Africa, 10 fath. (501); Prison Island,

8 fath.; Chuaka, 2 fath. (508), 3 fath. (626); Meweni Bay, 6 fath.

(510); Zanzibar town, shore (527).

* Résultats du Voyage du S.Y. Belgica, “ Bryozoa,”’ p. 27. . Zool. Chall. Exp.

vol. lxxix. p. 22, pl. 1. fig. 12.

+ See page 529.

ft Very minute glands occur in the vibracula of Cupularia.

482 MR. A. W. WATERS ON

CATENARIA DIAPHANA Busk. (Pl. LXIV. figs. 6-11.)

Scruparia diaphana Busk, Q. J. Micr. Se. vol. vii. p. 281,

pl. xxxi. fig. 1 (1860).

Catenaria diaphana Busk, “ Polyzoa,” Zool. Chall. Exp. vol. x.

pt. xxx. p. 14, pl. 11. fig. 3 (1884).

Halysis diaphana Norman, “ Polyzoa from Madeira,” Journ.

Linn. Soc., Zool. vol. xxx. p. 296 (1909); Levinsen, Morph. &

Syst. Studies on Cheil. Bryozoa, p. 274 (1909).

The ovicell has not before been described, in fact Levinsen says

itis wanting. A growth of the outer calcareous wall of this recum-

bent ovicell projects forward from the distal end in the middle

and also from the sides, forming at first one elongate space which

is subsequently divided, making two vacant spaces or large pores.

The node with an ovicell or ovicells is never a single zocecium,

and there are often many zocecia with ovicells in a node (fig. 9) ;

six together have been counted many times. We have seen

many ovicelligerous zocecia in a node in Catenicellide. The node

may, however, be only one ovicelligerous zocecium followed by an

ordinary zowcium. Asa rule, from the distal end of the older

zoceclum a new zocecium arises in the median line, and there

may also be one growing laterally from very near the end, or

occasionally one on each side.

The front wall is but little raised and is perforated, the per-

forated part being bounded on each side by a raised ridge, and on

the dorsal surface there are two lines of pores. The parietal

muscles start under the longitudinal ridge. The operculum has

a dark mark in the proximal part (fig. 8), and there are 20-22

tentacles. There are radicles attached to the side of the zowcium,

with the attachment elongate in the direction of the long axis.

None of the ovaria seen in sections are surrounded by the

follicular cells occurring in nearly all species. The ovarium

(fig. 10), with many ovarian cells, is found near the basal wall,

a short distance below the ovicell. The ovum is seen in the

upper part of the zocecium, where there are strong muscles ready

to force it into the ovicell from below internally, and sometimes

there is an embryo or ovum in the ovicell as well as a large ovum

below in the zocecium. Levinsen * considers that in some species

of Bryozoa the ovum comes out of the zoecium and then enters

the ovicell, but until proof is brought forward we may hold a

position of doubt, as the contrary is known in so many cases.

All the characters given by Norman for his genus Halysis

apply to Catenaria lafontii Aud., and it is not clear that a new

genus 1s required. Levinsen makes the absence of the avicularia

and of ovicells a reason for separating this species from his genus

Savignyella=C, lafontii Aud., but in a large proportion of genera

there is sometimes an avicularium, sometimes none. The genus -

Catenaria was discussed in my Report on the Sudanese Bryozoa 7,

* Loc. cit. p 67.

+ Journ. Linn. Soe., Zool. vol. yxxi. pp. 130, 131 (1909).

BRYOZOA FROM ZANZIBAR. 483

Loc. Madeira (.); St. Paul’s Rocks, N. Atlantic (Challenger),

shallow water. Ras Osowamembe, Zanzibar Channel, 10 fath,

(504); Prison Island, Zanzibar, 8 fath. (505), collected by

Crossland.

CATENICELLIDA.

In the Catenicellide the ovicells are of much more use in

classification than has been generally recognised, but Levinsen

has ignored them in his synopsis of the genera, nor has he used

the shape of the opercula.

Based upon the form of the ovicells, there are two main

divisions :—

First, those in which the ovicell is a terminal gonecium, as

Scuticella Lev., Costaticella* Maplestone, Cribricella ¢ Lev., all

three of which have an operculum with a straight or but slightly

curved edge, and Calpidiwm, which has a sinus. In all these

genera the ovicelligerous zocecium has a much wider operculum

than the ordinary zocecium, but the proximal edge is straight,

although in both the goneecium of all of these, and the ordinary

zoceclum of the first three, there may be an apparent sinus in

the calcareous wall.

SECOND, those in which the ovicell occurs in a node with other

zoccia ; divided into

I. Species with the ovicell occurring between two zocecia in a

straight line and the operculum straight or nearly so on the

proximal edge; and here we have Vittaticellat Maplestone, in

which the ovicell is partly imbedded in the superior zocecium and

is surrounded by a beaded structure; also Catenicella delicatula

Wilson and Claviporella pulchra MacG. have the ovicell in the same

position but perforated all over. Perhaps a new genus is required

for these two, and C. wmbonata$ B. may have to be included.

IL. Species with the tuberculate imperforate ovicell at the end

of a mother zoccium of a biglobulus, namely, C. perforata B.,

C. taurina B., C. cornuta B., but very few ovicells have been seen

in this group, and perhaps it is a matter of secondary importance

whether they are on a biglobulus or a triglobulus.

IIL. Species with the ovicell belonging to the mother zocecium

of a triglobulus, including Pterocella Lev., which has a double area

to the ovicell and the ovicelligerous aperture different from those

of the ordinary zocecia ; Strongylopora || Maplestone, with a per-

forated ovicell and the operculum straight at the proximal edge

* “Wurther Desc. of the Tert. Polyzoa of Victoria,” Proc. Roy. Soc. Vict. vol. xii.

n.s. p. 9 (1899). Levinsen, in making the genus Costicella, evidently overlooked

the fact that Maplestone had already made a genus Costaticella, of which the type

was lineata, a species included by Levinsen in his Costicella.

+ As indicated in the * Zoological Record,’ the name Cribricella has already been

used by Canu for a fossil belonging to the family Adeonide.

{ The surface of none of the Vittaticella seem to be perforate, but smooth or

papillose, but some have wrongly been described as perforate.

§ Described as fusca by MacGillivray.

|| This Levinsen calls Hincksiella, but Maplestone has priority, as his gens was

described in 1899 (Proc. Roy. Soc. Vict. vol. xii. p. 4).

Proc. Zoou, Soc.—19138, No. XX XITT, 33

AR4 MR. A. W. WATERS ON

of both the ordinary zoecia and the ovicelligerous zoecia, although

the notch in the caleareous wall has been taken for an oral sinus ;

Claviporella Ley., with a perforated ovicell and triangular aperture

to both forms of zocecia. :

The characters relied upon by Levinsen are none of them now

mentioned, as my object is to show the importance of the ovicell

and of the operculum in elassification, though of course all

available characters must be used.

Levinsen (p. 254, pl. xiii.) mentions a closure: in Vittaticella

and other genera which he calls an “occlusion.” J have not seen

anything quite like Levinsen’s description, in which it is said to

start from three processes, which, of course. is the youngest stage,

but in a specimen of Catenicella cornuta 8. from Western Port,

Victoria, there is in the older zocwcia a calcareous closure which

is, however, under the operculum and quite Independent of it.

There are two round openings near the distal end, that is, one

at each side, and a rather larger one at the proximal end.

Tn a few very old zowcia these two distal pores eoalesce and the

proximal opening becomes much larger, as if it were being dis-

solved away. I have only been able to find this closure in

C. elegans, in var. zanzibariensis, nov., and in C. cornuta as men-

tioned, although I have looked through the Catenicellidee in my

collection and the British Museum, though Levinsen gives it as

a character of Vittaticella (Catenaria Lev.). However, it only

occurs in the older and empty zocecia, and now, knowing this,

T might on re-examination find some which had been overlooked.

On p. 505 it is suggested that certain closures of Melicerititide

should be compared.

VITTATICELLA ELEGANS Busk. (PI. LXV. figs. 1-7, 12.)

Catenicella elegans Busk, Brit. Mus. Cat. Mar. Polyzoa, p. 10,

pl. ix. (1852); Zool. Chall. Exp., Polyzoa, vol. x. pt. xxx. p. 12

(pars); Ortmann, “ Die Japanische Bry.,” Arch. fiir Naturgesch.

vol. i. p. 27 (1890); MacGillivray, Prod. Nat. Hist. Vict.

dec. ili. p. 23, pl. xxiv. fig. 10 (1895); Thornely, Ceylon Pearl-

Oyster Fisheries, Suppl. Rep. xxvi. p. 109 (1905).

Vittaticella elegans Maplestone, “ On a new name— Vittaticella—

for the Polyzoan genus Caloporella McG.,” Proc. Roy. Soc. Vict.

vol. xiii. n. s. p. 203 (1900).

Specimens from Zanzibar growing on seaweed seem to be this

species. There is sometimes on one side of the zocecitum a very

long, raised avicularium with the mandible directed distally

(figs. 1, 2). A similar avicularium occurs in a specimen in the

British Museum from Avafura Sea. On the dorsal surface there

are the two small dots as figured by me in Vittaticella conter Aud.

(also in Brettia), and the radicles are in the same position as in

V. contei. In one case a new zocecium springs from the anterior

surface of an older zocecium, and this I have seen in other species

of Vittaticella.

The operculum is larger than that of V. contec, and is more

BRYOZOA FROM ZANZIBAR, 485.

curved on the proximal edge; on the other hand, it is not so

large as that of V. buskei W. Th. These are all nearly related,

but the difference in the operculum and the number of tentacles,

besides other characters, indicate that they must be separated.

There are often many ovicelligerous zocecia in one node. In

one specimen there are two cases of a node having six ovicelli-

gerous zocecia in one continuous line, and in both cases a lateral

zocecium grows from the side of the unjointed node. In the

ovicelligerous zocecia the avicularia near the oral aperture are

directed forwards instead of laterally, as in the other zowcia.

In the Tertiary beds of Curdies Creek, Australia, there is a form

with long biserial nodes which I described as Catenicella inter-

nodia*, but for which MacGillivray has since made the genus

Ditaxipora ; also from the North Italian tertiaries I have described

two forms with long biserial nodes as C. septentrionalist and

C. continua, of which continua is. probably Vittaticella, but

septentrionalis will require a new genus, so far as we can see at

present.

Among recent forms no long nodes have previously been

described, but in the specimen from Madeira, which Norman

considered to be C. contei Aud., there are two zocecia with ovicells

following one another in an internode. Long ovicelligerous nodes

are also found in Catenaria diaphana B. (see p. 482).

The ovaria commence at the proximal end at one side, whereas

the testes are near the distal end to one side. There are ciliated

embryos in the ovicells, and below the ovicells there are several

fleshy bands or tubes by which, no doubt, material for growth is

transferred to the ovicell. '

Loc. Bass’s Strait, 47 fath., Banks’ Peninsula, Algoa Bay,

Port Dalrymple; Tasmania (b.), Victoria (MJacG'.); Sagamibai,

Japan (Ort.); Gulf of Manaar, on floating oyster-cages (Zh.);

Arafura Bay (Brit. Mus.). Prison Island, Zanzibar shore (503) ;

Ras Osowamembe, Zanzibar Channel, 10 fath. (504), collected by

Crossland.

VITTATICELLA ELEGANS, var. ZANZIBARIENSIS, nov. (PI. LXV.

figs. 8-11.)

Zoarium about 40 mm. high, with the branches curved over.

Zocecia elongate ovoid, surface smooth or slightly papillose ; large

lateral avicularia with a large pore at the base, in this respect

differing from the type. There are on the dorsal surface near

the distal end, and often resting on the large avicularia, two

minute oval avicularia or an oval raised tubular opening. The

radicles arising from the dorsal surface form a thick bundle.

No ovicells are known.

It is very similar to V. elegans B., but the larger beaked

_* “Foss. Chil. Bry. from S.W. Victoria,” Quart. Journ. Geol. Soc. vol. xxxvii,

p. 318, pl. xvi. figs. 78, 79 (1881).

+ “ North Italian Bryozoa,” Quart. Journ. Geol. Soe. vol. xlvii. p. 5, pl. i. figs. 1-8

(1891).

Bon

486 MR. A. W. WATERS ON

avicularium with the large pore at the base and the minute

dorsal avicularia suggest its being separated as a variety.

The structure ofl She vittee has not received much attention,

though Harmer* has alluded to it; but in this species it has

been possible to obtain some explanation. The vitte are sunken

perforated grooves in the calcareous wall. and along each groove

there is a cylindrical tube, and within this, from the pore-tubes

(the perforations just mentioned), organic cords spread out and

reach the upper tree surface at definite spots or pores (fig. 10).

It thus seems that the vittze should be compared with pore-

chambers of many Cheilostomata in so far as there is indirect

communication from the interior to the water-surface, through

the vittee.

Loc. Prison Island, Zanzibar Channel, 8 fath. (505); Wasin,

Brit. Kast Africa, 10 fath. (500), collected by Crossland. Algoa

Bay and Natal (Brit. Mus.).

MEMBRANIPORA SAVARTII Audouin. (Pl. LXXI. figs. 1-4.)

In my Report on the Bryozoa from the Red Sea (Journ. Linn.

Soc., Zool. vol. xxxi. p. 138), I refer to the astonishing amount

of anastomosing protoplasmic threads in a specimen from Zanzi-

bar, and as some from the Sudan are also very full, this seems to

be a specific character. It certainly seems strange to find such

an extraordinary quantity, for though in my collection there are

preparations of a large number of species showing the threads

exceedingly well, I have never seen anything approaching these,

and further study of the funicular cords is desirable.

These threads are very abundant in zocecia with active poly-

pides having digestion in full activity. In these threads are

included small granular patches, either round or filiform, and

where the polypides have degenerated or are degenerating there

are large masses of this granular substance also surrounded by

and connected with the protoplasma (fig. 2). In earlier stages

the protoplasmic threads are in some cases surrounding the

granular cord (fig. 4); in others there are only one or two plasmic

threads by the sides of the granular cord or mass.

The collecting together of these masses naturally suggests that

waste products. are thus brought bepeuney and afterwards got

rid of.

Loc. Zanzibar Channel from the under side of buoy (528);

Ras Osowamembe, Zanzibar Channel, 10 fath. (504); Prison

Island, Zanzibar Channel, 8 fath. (505), collected by Crossland.

?MEMBRANIPORA ARMATA Haswell (2072 Koschinsky). (PI. LX VII.

fig, 10, & Pl. LX.XI. figs. 5-10.)

Biflustra armata Haswell, ‘On some Polyzoa from the Queens-

land Coast,” Proc. Linn. Soc. N.S. Wales, vol. v. p. 38, pl. i.

fig. 7 (1880).

* “ Morph. Cheil.,” Quart. Journ. Micr. Sc. vol. xlvi. p. 306 (1902).

BRYOZOA FROM ZANZIBAR. A487

Membranipora panhoplites Ortmann, “ Die Japan. Bry.,” Arch,

Naturgesch. vol. i. p. 28, pl. 11. fig. 4 (1890).

Membranipora armata Waters, “On Membraniporide,” Journ.

Linn. Soe., Zool. vol. xxvi. p. 687, pl. xlvii. fig. 3 (1898).

The specimen from Wasin is in parts in the hemescharan stage,

in others in the bilaminate, and one in my collection from Port

Molle is also bilaminate. This specimen from Wasin starts from

a unilaminate incrusting layer, but in places the zoarium is

tubular.

The distal wall of the zocecium is slightly prominent, somewhat

reminding us of what Busk calls the penthouse projection in

Aspidostoma giganteum B. There are no ovicells in any of the

specimens examined, and on most of the zocecia an avicularium

occurs on one side only, at the distal end, directed proximally,

while on the other side, in a long chamber, there is a long gland

with distinct secreting cells (figs. 8, 9), but usually without any

lumen. These glands are irregular in shape, sometimes lobed,

and there may be two elongate lobes side by side. In all the

specimens seen there is a distinct calcareous bar or arch to the

avicularium, and there are two openings on the front of the avicu-

larium, though sometimes the lateral projections in the avicularium

do not meet, when there is, consequently, only one opening

(fig. 7). In the membrane covering the avicularian chamber

there is, under the mandible, a chitinous ring where the peculiar

body * ends (fig. 9, pb.), and there are other species of Cheilo-

stomata with a chitinous ring or other thickening. As we have

seen, there is at one side an extremely long avicularian chamber,

at the proximal end of which there are stout muscles (fig, 9 a)

attached to a very long tendon by which the mandible is closed ;

further up there are muscles also on both sides of the chamber,

but much more delicate than the last (fig, 9 6), aud attached to a

shorter tendon fastened to the base of the mandible with a median

attachment, whereas in some species of Cheilostomata this muscle

is attached at each side.

The chamber containing the glands (fig. 9, gc.) is also very long,

and may be close to the avicularian chamber of the zocecinm next

above or below, but no connection with the avicularia has been

found after careful examination of many microtome sections.

On the inside there ave pores like rosette-plates, and the proto-

plasmic threads from these are sometimes seen passing to the

gland, but no other internal opening has been found, Externally

there are three or four minute pores along the line of junction of

the zoecia, which, however, were only noticed when carefully

searching for openings, There are similar small pores over the

avicularian chamber. We have here another Bryozoan puzzle,

for the gland is not the same as the oral gland, and is contained

in a caleareous chamber without any openings except small pores.

* This peculiar body is in many Cheilostomata contained in a sheath, homologous

with the tentacular sheath, but in this species no sheath was found.

488 MR. A. W. WATERS ON

The operculum is membranous, with a diagonal band or sclerite

on each side to which the muscle for closing the operculum is

attached ; but also attached to the same sclerite there is a muscular

band fastened to the tentacular sheath. I have not noticed a

double attachment like this before, but examination may show

that it occurs in other species.

In a large proportion of zocecia there are two polypides of

about the same size, so that 20-30 zocecia with two polypides

may be seen adjoining one another, whereas other pieces may

show a much more limited number of double polypides. Although

budding polypides in the same zoccia as mature polypides are

known to us all, and have been described by Haddon*, Ostromoff 7,

and Harmer {, they are only in a limited number of zocecia ; nor

have I been able to see that they are the same as the two zoecia

described by Prouho § in Aleyonidium duplex P. The two tenta-

cular sheaths are side by side, and are attached to the operculum

and the neighbouring wall. No ovaria or ova have been seen,

and only in a very few cases were testes found occurring in round

masses near the lateral wall.

From the lateral walls there are bundles of muscles (6-12)

attached to the frontal membrane of the zoecia.

In all the lateral walls there are pores at fairly regular

intervals all over the walls, and besides there are in some cases

disks with numerous pores near the opercular wall. Further, in

a bilaminate piece of 17. armata, there are in several cases large

perforated disks on the basal walls, like those described in

Petralia for the radicle attachment, though, strangely, in the

unilaminate parts no distinct perforated disks have been found.

This form cannot remain under J/embranipora, though I am not

suggesting that it is Petralia, but call attention to various similar

oer dehors in forms placed far asunder.

There are about 30 tentacles, which is a larger number than

has been found in any true Membranipora. Membranipora nigrans

Hincks and WM. marginella H., with avicularia similarly placed

near the distal end, have also cur ious large vicarious avicularia.

Loc. Port Danion, Holborn Island, oy 0 fath. (Haswell); Port

Molle, Australia ; Sagamibai, Japan, 40 fath. (Ortmann). Wasin,

Brit. East Africa, 10 fath. (500); Zanzibar Channel, from under-

side of buoy (528), collected by Crossland.

MEMBRANIPORA CATENULARIA Jameson.

For synonyms see Miss Jelly’s Catalogue.

Although the Arctic W. monostachys Busk from Franz Josef

Land has many points of similarity, the operculum in J. catenw-

laria from Zanzibar is only about ‘half the width of that of the

* Quart. Journ. Micr. Sc. vol. xxiii. p. 520 (1883).

t Arch. Slaves de Biol. vol. 11. p. 341 (1886).

apace Excretor, y Processes in Marine Polyzoa,” Quart. Journ. Micr. Sc. p. 189 (1891).

( § “Cont. a Vhist. des Bry.,” Arch. de Zool. Expér. et Gen. 2nd ser. vol. x. p. 581

1892). :

BRYOZOA FROM ZANZIBAR. 489

former species, and does not reach to the border of the opesium,

so that they can be distinguished by this character. *

Loc. Widely distributed, but there is much uncertainty about

some of the determinations. Chuaka, Zanzibar, 3 fath. (506),

2 fath. (508), on the dorsal surface of iseareponclia aan is b.,

collected by Crossland.

Farcimia ocunata Busk. (Pl. LX VII. figs. 8, 9.)

For synonyms see :—

Farcimia oculata Waters, “Rep. on the Mar. Biol. of the

Sudanese Red Sea, Bryozoa,” Journ. Linn. Soc., Zool. vol. xxxi.

p. 167 (1909) ; and add Canu, “ Bry. des Terrains Tert. des Envy.

de Paris,” Ann. de Paléont. vol. ii. p. 20, pl. 11. figs. 36, 37 (1907);

pebsiye Helv. del Egypte,” Mem. 4, I Inst. Eeyptien, vol. Wie Dole

pl. x. figs. 16-19 (1912).

Nellia tenella Le svinsen, Morph. & Syst. Studies of the Cheil.

Bry. p. 120, pl. i. figs. 13 a—13 e (1909).

The growth is from a spreading stolon from which, at intervals,

sub-colonies grow (figs. 8, 9), commencing with short calcareous

nodes joined ‘by chitinous tubes. There are usually three or four

nodes in the stalk, though there may be only one. Quite simi-

larly sub-colonies on a stalk with internodes grow from delicate

stolons in Chlidonia cordiert Aud., Diplodidymia complicata

Reuss (Pl. LX VII. figs. 11, 12), Catenaria parasitica Busk *.

I figured it some years ago for Chlidonia cordieri Aud. 7, and

since then Calvet and Levinsen have dealt with the species, All

these species have a somewhat similar operculum, about the same

size, and the number of tentacles in all is 11-12. They are

probably more nearly related than we have imagined.

_- There are two very thick chitinous tubes connecting each new

branch.

The ovarium, with several ovarian cells, often in a row, is near

to the inner wall, about equidistant from the distal and proximal

ends. One ovum grows extremely large, and is pressed into all

kinds of shapes thr ough want of room. The ovum passes into a

sac near the basal “nal and the distal end, and before an ovum

has passed into it muscle-threads are seen radiating over this sac,

Levinsen has figured the ovicell, which is a ‘small cap-like

growth, and this I have seen in a Basin cases, but often in zocecia

with ovicelligerous zocecia no external difference is recognised.

There is, however, often a vertical division separating the ovi-

cellular wall from the rest of the zocecium,

The triangular mandible of the avicularium is found with

difticulty and is extremely minute, being about 0°006 mm., while

a large number of the mandibles, such, for example, as in Retepora

cellulosa, are 35 times as long; some, as for example in Lepralia

* The Honourable Mary Palk informs me that Catenaria lafontii Aud., grows on

a similar stalk.

+ Journ. Linn. Soe., Zool. vol. xxvi. pl. 1. fig. 8 (1898).

490 MR. A. W. WATERS ON

occlusa B., are 60 times as long. In many cases there seems to

be no mandible, only a disk, to which the peculiar body is attached,

and this peculiar body is relatively very large. Smitt refers to

the mandible of the Floridan specimens often being wanting,

“presenting the opening closed only by a membrane.”

There is no possibility of knowing what the Cellaria tenella of

Lamarck was, and it certainly may have been Cellaria*, so that,

as I have previously said, it is better to retain the name oculata.

The genus Vellia was not satisfactorily described, and therefore

various authors have adopted the genus Farcimia of Pourtales

and Smitt. Fleming had made a genus Farcimia which might

include Wellia, but as he made it for Cellaria, with C. fistulosa

as type, it was always a superfluous genus, and Smitt considered

it non-existent. Although Levinsen adopts WVellia, it seems

better to adhere to Farcimia, the name used in my recent papers,

and which has been used by most workers recently.

Loc. See my paper referred to, and add Wasin, Brit. East

Africa, 10 fath. (501); Prison Island, Zanzibar Channel, shore

(503), 10 fath. (505); Ras Osowamembe, Zanzibar Channel,

10 fath. (504); Meweni Bay, Zanzibar, 6 fath. (510); Chuaka,

Zanzibar, 3 fath. (526), collected by Crossland. Texas and

St. Thomas, W. Indies (Levinsen).

DIPLODIDYMIA COMPLICATA Reuss. (Pl. LXVITI. figs. 11-15, &

text-fig. 79.)

Diplodidymia complicata Reuss, ‘‘ Foss. Fauna der Oligoc. von

Gaas,” Sitzungsber. d. k. Ak. der Wissensch. Wien, math.-nat. Cl.

vol. lix. Abth. i. p. 469, pl. ii. figs. 6-9 (1869).

Micropora ratoniensis Waters, Ann. Mag. Nat. Hist. ser. 5,

vol, xx. p. 185, pl. iv. fig. 5 (1887).

Micropora articulata Waters, Quart. Journ. Geol. Soe. vol. xlvii.

p. 14, pl. u. figs. 5, 6 (1891). é

From Chuaka, Zanzibar, there are several pieces throwing

light on this form, which has only been seen in fragments

previously.

The zoarium has a stalk consisting of long barren internodes,

sometimes as many as eight, followed by long articulated inter-

nodes with the zoeecia placed diagonally on the four sides. The

contents of the barren internodes send out a branch to the

surface, just as we have seen in Chlidonia cordierti Aud. From

near the starting-point of each sub-colony a number of narrow

radicles radiate, and sometimes from one of these radicles a

fresh sub-colony grows, as is frequently the case in species with

creeping stolons.

The branches are dichotomous and articulated, having two

chitinous tubes forming each articulation. Occasionally there

are more than the four rows, but this will only be for a short

distance near the articulation, and in one piece there is a median

* Busk gives it with ?synonym of Cellaria gracilis Busk.

BRYOZOA FROM ZANZIBAR. 491

line on the dorsal surface with the zoccia on each side. The

front wall has a minute pore on each side below the oral aperture

(fig. 13), but these are only seen in Eau de Javelle preparations,

and in the same way the Jong slit-like pore on the one side is

scarcely seen so long as the membrane covers the walls. The

muscles passing through this slit are wide, and are attached to a

Text-fig. 79.

Diplodidymia complicata Reuss.

A. Showing small sa¢és hanging from the opercular region, and in each an ovum

grows (ov.). At y. there seems as yet no ovuin in the small sac; m., the

muscles of the opesium. X 50.

B. Section showing an ovum in the small sac. X 250.

C. Section in which the ovum has segmented and a blasluta is formed. X 250.

chitinous thickening or sclerite on the front membrane, and are

partly protected by a calcareous wall. There are two rosette-

plates in the entire lateral wall. On the avicularian chamber

there is a concave pit with a central perforation, no doubt indi-

cating that radicles can be given off from this pit, and thereby

492, MR. A. W. WATERS ON

the two pores described in the fossil Wieropora articulata Waters

are explained, the one being the avicularian and the other the

radicular opening. ‘The avicularium is by the side of the aperture,

and the mandible is triangular.

There are 11—12 tentacles.

The testes extend all down the lateral wall. A very few small

ovaria with one to three small ovarian cells were found.

A small ovum is found in a small sac hanging from the

opercular region (text-fig. 79). The sac and ovum grow until

ultimately the embryo nearly fills the zocecium, and now there

are small muscles from the opercular wall to the ovicell and

a strong lateral band. There is no external indication of any

ovicell nor of which cells contain an embryo. The embryo is

relatively large, with the couronne very large and distinct, and

the way in which the embryo grows in the pendant sac may

throw light upon the development of the ovum and embryo in

Adeonella, but no stage has been found with a sac hanging from

the opercular region in Adeonella.

The present species, as [ have previously indicated, is closely

allied to the fossil Cellularia diplodidymoides Meun. & Pergens

from the Chalk, and both belong to the same genus *, Canu has

described four fossil species from the Paris Basin, but, unfor-

tunately, he has not given figures as well as photographs. In

the work there are a number of magnificent photographs,

showing the characters most beautifully, but some of the Paris

authorities are making too hard and fast a line that everything

must be photographed. The specimens in question (pl. v. figs.

6-10) do not lend themselves to photography, and require either

figures or full description for elucidation.

These Zanzibar specimens are so different in the younger and

older joints that with fossils several species might be made from

one colony.

Canu 7 says that the genus Diplodidymia is the Poricellaria of

dOrbigny, but the description of this latter leaves recognition

impossible without direct comparison, and therefore Canu is quite

right in retaining the name Diplodidymia Reuss.

Loc. Off Katow, New Guinea, 7 fath.; Singapore (fide White-

legge, in lit.). Chuaka, Zanzibar, 3 fath. (506), collected by

Crossland.

Fossil. Gass, near Dax, 8. France, Oligocene; Montecchio

Maggiore, N. Italy, Bartonian.

CHLIDONIA CORDIERI Audouin. (Pl. LXV. figs. 15, 16.)

EHucratea cordiert Aud. Descrip. de ’Egypte, Hist. nat. p. 242,

2nd ed. p. 74; Savigny’s pl. xiii. fig. 3.

Chlidonia cordiert Waters, Journ. Linn. Soc., Zool. vol. xxvi.

* “ Bryozoaires du Systtme Montien,” Louvain, p. 8, pl. ii. fig. 3 (1886).

+ “ Bryozoaires des Terrains Tertiaires des Environs de Paris,” Ana. de Paléont.

t. liv. p. 39 (1907).

BRYOZOA FROM ZANZIBAR, 493

p. 18, pl. i. figs. 8, 9 (1896), which see for synonyms, and add:

Calvet, ‘Bry. Mar. de Cette,” Trav. de I’Inst. de Zool. de Univ.

de Montpellier, ser. 2, mem. 11, p. 13, pl. 1. figs. 1, 2 (1902) ;

‘“‘Bry. Mar. des Cotes de Corse,” op. cit. mem. 12, p. 6 (1902) ;

Levinsen, Morph. & Syst. Studies on the Cheil. Bry. p. 197,

pl. viii. figs. 6 a-6 y (1909).

From the front wall to the zocecial wall, through what has been

called a second chamber, but is the equivalent of a compensation

sac, there is a bundle of three, four, or more muscles (see fig. 15).

The attachment of these on the front has been mistaken for a

suboral pore, and in dried specimens there is frequently an

opening here.

No ova or ovaria have been found in any of my sections,

whereas some show an embryo about half filling the zocecium,

though no external difference has been noticed.

The operculum is interesting, as it has at each side a projection

or wing at right angles to the operculum (figs. 15, 16); also at

each distal corner there is a slight projection. The wing reminds

us somewhat of the thin membranous growth of many Mem-

branipore and some Micropore.

There are 9-11 tentacles.

Loc. Red Sea (Aud.); Naples, Trieste, Rapallo, Nice, Cette,

Corsica, Algiers, Tunis, Tyre, Calvados, Egypt, Victoria (Aus-

tralia), Cape York, New Zealand, Atlantic (fide Carus); Canaries

(dOrb.). Wasin, Brit. Hast Africa, 10 fath. (500), on calcareous

seaweed, collected by Crossland.

CELLARIA.

Levinsen, following Norman, uses Cellularia for what we

understand as Cellaria, but as these names have long been used

for widely distinct genera I must certainly, in the most definite

manner, refuse to use the name Cellularia for what we have for

many years understood as Cellaria.

Cellularia of Pallas was a simply ridiculous jumble of forms for

which a place had not been found elsewhere. The species men-

tioned by him are now placed in nine genera, one of which is the

Cyclostome Crisia, and as the description of the genus refers to

the ovicell of Crisia, perhaps the least objectionable thing would

have been to have retained the name Cellularia for Crisia. The

real difficulty, however, is that the name Cellulavia has been

retained for a quite different group, and to interchange and now

use a name long understood in a different sense would cause the

greatest confusion.

Solander employed the name Cellaria for a group approxi-

mately, but not absolutely, similar to the Cellularia of Pallas,

giving a definition also not quite the same, and it has been con-

sidered to be only a change of spelling, though curiously, Ellis &

Solander never indicate that they considered it was the same as

Cellularia of Pallas, nor throughout the description of the genus

494 MR. A. W. WATERS ON

do they ever refer to Pallas, although many of the species dealt

with had been mentioned by that author. This is certainly

difficult to understand, and perhaps would have been different

if Solander had lived to complete the work himself.

Hincks considered that the species in Solander’s genus had all

found places elsewhere, and that the genus of Solander had lapsed.

He therefore took Cellavia as the genus of Lamouroux, who under

Cellaria put Cellaria (as now understood) with the type C. sali-

cornia and also Tubucellaria. Lamouroux says of all genera none

seems to contain as widely distinct species as this, and that

it seems to have been formed to contain everything that could

not be placed under Flustra or Sertularia.

Instead of dropping Celluluria as hopeless, Busk unfortunately

retained it for a small division, and this has been accepted.

The first species mentioned by Pallas in his genus is tubu-

cellaria, which also is included in Lamouroux’s Cellaria, though

not as the type, which was Cellaria salicornia; and Stoliczka*, in

a long discussion of the subject, took the view that as tabucellaria

was first mentioned the genus must be called Cellaria, and

Cellaria, as now understood, must be separated as Salicornia.

We now see that it is unfortunate that Hincks should have

retained Cellaria, though it then appeared that this would not be

challenged, and it has been adopted generally, and no genus

seemed more firmly established. However, it is now clear that if

Hincks had continued to use Salicornaria there would have been

no possibility of the name of a now long recognised genus being

replaced by one used in most various ways, and now limited to

another small group.

Those who are at work upon the class know how often the

descriptions of the earlier authors are now meaningless, for the

characters then used are found to be useless; but this can hardly

be appreciated by those who have not had occasion to consult

such descriptions. It is as if some well known tree had genera-

tions ago received a name and a few lines of description which

would apply to a quarter of our phanerogams and some cryptogams.

I have previously shown that we are brought into a perfectly

ridiculous position by being asked to adopt such names at all

costs, when we often have no idea what they meant. It is not

science, and since Cellaria as modified by Hincks is well established

I shall still use it f.

* Foss. Bry. aus dem tert. Griinsand. der Orakei Bay bei Auckland, pp. 142-149

(1864).

+ Norman (“Polyzoa of Madeira,” Journ. Linn. Soc., Zool. vol. xxx. p. 293

(1909)) challenges the correctness of considering that Tubucellaria opuntioides

should have been considered the type of Cellularia, but I cannot agree with his

conclusions as to what I say being contrary to the British Association Rules of

Nomenclature. My edition is later than Norman’s, but apparently is only areprint,

and it says “ When they omit doing so” (7. e. fixing a type), “it may still in many

cases be correctly inferred that the first species mentioned on their list, if found

accurately to agree with their definition, was regarded by them as the type.” This

rule of course means that if the generic diagnosis is taken from some one species,

and that an error has been made in including the first, then common sense may be

BRYOZOA FROM ZANZIBAR. 495

CELLARIA GRACILIS, var. TESSELLATA, nov. (PI. LXVII. fig. 7.)

For synonyms of Cellaria gracilis see Miss Jelly’s Catalogue

and add :—

Meissner, ‘* Liste der von Herren Prof. Simon bei Amboina und

Thursday Island ges. Bry.,” Jena Denkschr. vol. vill. p. 730.

A specimen from Ras Osowamembe growing on Hydrozoa

seems to be a variety of C. gracilis. In this specimen the trabe-

cule mentioned by Busk enclose an area formed of large tessellated

divisions, about 20 (fig. 7); and curiously, another species, which 1

eall C. wasinensis, sp. n., has also a tessellated area, but the

avicularium of that species is triangular with an acute mandible,

and belongs to the C. tenwirostris group. These large tessellated

arens are unusual, and have not been found in various species of

Cellariain my collection ; nor in the British Museum ‘ Challenger’

collections are they found in C. bicornis B., C. dubia B., C. malvi-

nensis B., C. variabilis B., C. divaricata B., C. australis Hincks,

CU. rigida MacG. The significance both of the trabecule and of

the divisions is at present obscure.

The opercula are a trifle larger than the type from Holborn

Island, but the mandibles are the same.

Loc. Type: Cumberland Island ; Cape Capricorn ; Victoria,

8 fath.; Torres Straits; Holborn Island (Queensland); Katow,

New Guinea, 7 fath. Variety: Ras Osowamembe, Zanzibar

Channel, 10 fath. (504), collected by Crossland.

CELLARIA WASINENSIS, sp.n. (PI. LXVII. figs. 1-6.)

Zoarium about 25mm. high, with small branches dichotomously

jointed and connected by two or three stout straight chitinous

tubes and a “knot.” The zoarium swells out where the ovicells

occur, as is frequently the case in Cellaria, there being sometimes

two such swellings in an internode ; and it will be seen on fig. 6

how these are equidistant from the joint in the two branches.

The zowcia are rather elongate, hexagonal, or rhombie, with the

lateral walls of the zocecium much raised, surface finely granular ;

oral orifice wide, with two teeth on the distal edge, which is

slightly turned up, forming a lip.

The ovicellular aperture is large, round, with a plate from the

proximal border spreading out and partially closing the aperture.

used in saying the first is not the type, but surely it never meant that any individual

could subsequently pick and choose which was the type among those agreeing

equally well. However, in my work, to which Norman refers, I and other specialists

were under an obligation to follow the rules of the Zoological Congress, and the rule

in question is “other things being equal the name is to be preferred which stands

first in the book or article.” In anything I write now the same obligation does not

exist, though the rule entirely commends itself, and it does not seem to clash with

the rules previously mentioned, drawn up by Strickland. Both practically say that

when there are adequate reasons to indicate which was the type intended, that may

be considered the type although not first mentioned, but without good cause to the

Say the first is the type, and independently this is what workers are constantly

oing.

496 MR, A. W. WATERS ON

The oyvicellular opening of C. gracilis Busk, var., C. australis

MacG., C. rigida MacG., and C. hirsuta MacG., is similar; but in

the few cases where the ovicellular aperture 1s mentioned, it is

often only the incompleted opening in an early stage which is

described. ;

The avicularia are triangular with mandibles like those of

C. tenuirostris B. In the avicularium the calcareous sub-

mandibular wall rises up to the proximal edge of the mandible

with a notch on each side, perhaps for the muscles (fig. 5), so that

the submandibular part of the avicularium is almost entirely

closed; and this 1s an interesting point, for Levinsen considers

that the Melicerititide differ from the living Cheilostomata in

having the submandibular portion entirely calcified.

As I have mentioned *, there is, however, a specimen in the

Museum dhistoire naturelle in Paris from the Banecs des

Aiguilles, 8. Africa, which is probably the J/acropora cribrilifera

Maplestone t, fossil from Mitchell River, in which the large

vicarious avicularia have the submandibular part entirely

calcified. Maplestone mentions that three of the zocecia have a

‘“‘caleareous closure.” I have previously stated that there seem

to me to be some points of relationship between Cellaria and

Melicerititide.

Levinsen deals but very shortly with the avicularia of Cellaria,

and I am not quite sure that I follow what he means about the

avicularium of C. malvinensis. He, however, says that the sub-

mandibular er yptocyst, reaches up to the operculum in C. fistulosa.

I have not seen it rise as it does in C. wasinensis in any of my

specimens of C. fistulosa L., but there is a similar wall rising to

the base of the mandible in C. variabilis B., C. hirsuta MacG.,’

and C. gracilis B.; C. variabilis has two slits in the submandibular

part. More frequently there is an open rounded submandibular

space as in C. dennantt MacG.t, C. malvinensis B.t, C. australis

H., C. fistulosa, C. tenuirostris Busk, and C. wandeli Calvet. In

C. fistulosa this is not much more than a wide round sinus.

There are two species which have been taken for C. malvinensis

B.. The first, which I have from Wanganui, New Zealand, has a

fairly large submandibular space with a distinct ridge where the

proximal end of the mandible comes, in fact in a few cases this is

continued, forming a bridge across. The mandible soon contracts,

with the distal end lanceolate. The other form is slightly

smaller, from Baie Orange, S. Africa, and mentioned by Jullien as

C. malvinensis. It has the sides of the zoecia straight, the distal

end rounded, and the submandibular part of the avicularium rises

up to the mandible, having two diagonal slits. The mandible is

shorter than in the other species, sloping gradually to the apex.

Whenever this is figured it might be called C. jullient. The

* Résultats du Voyage du S.Y. Belgica, “ Bry oz0a," ’ p. 85 (1904).

+ ° Further Desc. of the Tert. Poly. ot Victoria,” Proc. Roy. Soc. Vict. vol. xiii.

n. s., p. 204, pl. xxiv. fig. 2 (1901).

4€ Rés. du Voy. du 8.Y. Belgica, pl. it. fig. 9a & pl. viii. figs. 4, 5.

BRYOZOA FROM ZANZIBAR. 497

forms from Curdies Creek which I considered to be malvinensis

have since been separated by MacGillivray as C. contigua, but the

fossils from Bairnsdale and New Zealand are C. malvinensis.

The operculum of C. wasinensis is nearly semicircular with

large hollows fitting on to the teeth, and is similar to the

operculum of C. gracilis. The trabeculee (figs. 3, 4) enclose in the

lower part large divisions, about 7 in number, and the divisional

walls are very thick, whereas in C. gracilis var. tessellata, nov.

(fig. 7), they are linear with numerous divisions.

There are about 13 tentacles. In Cellaria we find * the number

of tentacles is approximately the same throughout the genus, only

C. dennanti MacG., a species showing other differences, has 20.

Now Cellaria is a well marked genus hhaving opercula of a special

form, with a hollow cup fitting on to the teeth in the oral

aperture, and the ovicell has a characteristic chamber, and also a

characteristic ovicellular aperture.

Levinsen 7 states that a zocecium does not correspond with an

area, but with this I cannot agree, as I find the superficial

divisions approximately mark off the zocecia, even though they

may in parts extend somewhat under the divisional line, and this

is the case In many genera.

The ovarium usually has two ova, though there may be one or

three, and the ovum is large when it enters the ovicell.

Loc. Wasin, Brit. Hast Africa (501) (507), 10 fath.; Ras

Osowamembe, 10 fath. (504); Prison Island, Zanzibar Channel

(505), 8 fath.

Cellaria tenwirostris B., C. salicornioides Aud., C. magnifica B.,

O. malvinensis B., C. gracilis B., have been previously described

from tropical regions.

THAIROPORA MAMILLARIS Lamouroux.

Flustra mamillaris Lamouroux, ‘“ Polyp. corall. flexibles,” p. 110

(1816) and add to Miss Jelly’s synonyms :—

Thairopora mamillaris MacG. Prod. Zool. Vict. dec. xx. p. 351,

pl. 196, fig. 2 (1890).

Membranipora mamillaris Hincks, Ann. Mag. Nat. Hist. ser. 6,

wolevaltenp.. 9 (USO):

Thalamoporella mamillaris Levinsen, Morph. & Syst. Studies

on Cheil. Bry. p. 194, pl. via. figs. 5 a—5 e (1909).

There is a small piece from Chuaka, Zanzibar, which was de-

calcified when received. It was preserved in HgCl,, but no

doubt there was acid as well. The main poimts could, however,

be seen, and the opercula and mandibles were separated, and

although the mandibles are smaller than in the specimen received

from the Red Sea, to which reference is made below, yet they are

quite characteristic of 7. mamillaris. Levinsen has shown that

there are calcareous spicules in all the Thalamoporellide, affording

* Résultats du Voyage du S.Y. Belgica, “ Bryozoa,” p. 37 (1904).

7 Morph. & Syst. Studies on the Cheil. Bryozoa, p. 211.

498 MR. A. W. WATERS ON

good specific characters, and the way in which he has dealt with

them is one of the most important features of his book.

Loc. Victoria (MacG’.); South Australia (my coll.); a weed,

brought up on the sounding-line in the Red Sea, covered with this

species, was given to me by a passenger. We may take it that

there is every probability of its being from the Red Sea, though

we cannot record it as such without pointing out the possibility

of its having remained on the line from some previous locality.

Chuaka Bay, Zanzibar, 2 fath. (509), collected by Crossland.

SrEGANOPORELLA MAGNILABRIS Busk. (Pl. LX XIT. figs. 12-20.)

Membranipora magnilabris Busk, Brit. Mus. Cat. Mar. Polyzoa,

p. 62, pl. Ixv. fig. 4, in the explanation of the plate called

M. grandis.

Steganoporella magnilabris Harmer, “‘ Rev. of Gen. Stegano-

porella,” Quart. Journ. Mier. Sei. vol. xli. p. 279, pl. xii. fig. 10,

pi. xiii. figs. 31, 44-46, which see for synonyms. Add Thornely,

‘“‘ Mar. Poly. of the Indian Ocean,” Trans. Linn. Soc., Zool. vol. xv.

p. 145.

Specimens from Wasin agree in most respects with the descrip-

tion of this species, though the mandibles of the A zocecia having

no teeth are more like those of S. buskit Harm. However, a

comparison with the British Museum specimens has led to my

leaving them under magnilabris, though some workers might *

separate them asa variety. Thezoaria are bilaminate, irregularly

contorted, and evidently attain to a very considerable size, as

some pieces, evidently only fragments of larger ones, measure

two or three inches across.

There are very few B zoecia, in fact two mounts, each con-

taining about 300 zocecia, had no B zoecia, On examining all the

remaining material a few scattered B zocecia were found, and it is

interesting to find that from these zocecia, which are somewhat

larger than most A zowcia, two new zocecia usually arise, that

is the row here bifurcates, and to this but few exceptions were

found in all the material examined.

I called attention to this duplication from the B zocecia occurring

in some fossils, but Harmer ft has shown that it is by no means

universally the case. At the time I wrote very little material

was available for comparative study. It, however, is a fact that

in many species two zoccia usually, or at any rate frequently,

grow from the distal end of B the zoccia. In this species, when

the operculum is removed, it is often not possible to be sure which

are B zocecia, as there are large A zocecia, with a large shelf, in

all respects similar excepting the operculum, so that with fossils

there will often be uncertainty. In most other species there is a

greater difference between the A and B zoecia. The A opercula

vary considerably in size. In a specimen from (508) the B zowcia

are much more common than in those from (500).

* “Bryozoa from Aldinga,” Quart. Journ. Geol. Soe. vol. xli. p. 292.

+ Revision of the genus Steganoporella, p. 266.

BRYOZOA FROM ZANZIBAR, 499

According to Harmer the A zocecia are in most species of

Steganopor -ella more abundant than the B, and he says there are

no B in 8S. simplex H., and goes on to say that they are rare

in S. suleata H., S. lateralis H., and S. magnilabris B. In

S. truncata H. the ratio of A to B is 4-5 to 1; on the other hand,

in S. alveolata H., there are few A zoecia, and S. connexa H. has

only B zoecia.

As Harmer™* has shown, there is considerable difference in the

teeth of A opercula of magnilabris from various localities, and we

have opercula of this Wasin specimen without any, also the B

opercula may be with or without lateral teeth. The A opercula of

a ‘Challenger’ specimen (208-90°4.16.13) has the teeth so minute

that they would be overlooked with a low power, but are seen

with a quarter-inch objective.

The operculum of Steganoporella can scarcely be compared with

the opercula of other Bryozoa, as it closes the whole of the distal

chamber, and in a zoarium there is an amount of variation in size

of the opercula which is unknown in other genera of Bryozoa.

In a British Museum specimen of S. truncata Harm. from Port

Dalrymple, there are a few cases where the operculum is thrown

back, and then the whole of the opercular opening (that is over

the distal chamber) is covered by a membrane with a large round

opening, thus furnishing a most interesting form of closure.

The wall, dividing the two chambers, passes vertically from the

opercular wall to the basal wall, and in the middle there is a

round opening, sometimes with a slight tubular projection through

which the polypide passes. The polypide rests partly in the two

chambers, that is the tentacles are not entirely withdrawn into

the proximal chamber. The embryos develop at the base of the

distal chamber, but this chamber can by no means be spoken of

as an ovicell.

The Wasin specimens show some blind zoccia either entirely

closed or with a central opening, and the same occurs in a speci-

men of S. tubulosa H., where two zocecia are about the normal

size, one with a large round opening, the other with an oval one,

and in these the whole of the frontal wall is granular. On the

other hand, a British Museum specimen of S. solace H. has two

zocecia entirely closed by the perforated eryptocyst.

The S. magnilabris from Wasin has floating in the proximal

chamber many small oval bodies, surrounded by a membrane

containing diatoms and other detritus. These are the excre-

ment pellets, and they are evidently frequently ejected within

the zocecium.

Study of the growing ends is instructive. At first there is

an absolutely empty oblong zocecium covered with a plain mem-

brane, and in this either A or B opercula may be formed, but at

first the operculum has no basal sclerite (fig. 20), as this is

formed subsequently. Next the proximal part of the eryptocyst

* Toc. cié. p. 234.

Proc. Zoou. Soe.—1913, No. XXXIV. j4

500 MR. A. W. WATERS ON

is formed (fig. 20, z 2-2 3), then the tube or opening between the

two chambers (fig. 20, 24) which is at a much lower level than

the operculum, next the lip is formed, which at last reaches up to

the operculum. The muscles are formed after the operculum,

and the polypide not until after the tube and lip.

It has not been mentioned that the part between the main

sclerite of the operculum is covered by a membrane, thus enclosing

a space (fig. 17), and the tentacular sheath is fastened * between

the two muscular attachments of the operculum, having a tissue

across from one muscular attachment to the other (fig. 19,

from r-r). As mentioned, the large distal muscle is attached to

the operculum, but the large muscle just proximal to it in the

A zocecia is fastened by a tendon to the frontal membrane (fig. 19 d)

close up to the operculum, where there is a slight thickening or

sclerite. Harmer f speaks of this muscle as perhaps inserted into

the frontal membrane immediately adjacent to the basal sclerite.

Further back (proximally) there is on each side a diagonal

sclerite (fig. 14, sc.), to which the tendons of a smaller muscle are

attached, which draws down the frontal membrane {. These are

called depressor muscles by Harmer $}).

In the B zocecia the retractor muscles are much larger than in

A, and are in two groups instead of one, but in the present species

I am unable to find any very material difference between A and

B zocecia.

The embryos develop at the base of the distal chamber, but

this chamber cannot be spoken of as an ovicell, for the polypides

are, even when retracted, partly in this chamber. It seems that

the embryo may exceptionally be in the proximal part of the

zocecium surrounded by a membrane, really an internal ovicell.

This requires investigation.

The first polypide buds in the growing terminations are in the

proximal end of the young zoccia as usual, and are nearly always

in one of the corners, consequently the polypide grows diagonally

across the zoccium, causing a slight amount of asymmetry in the

zoecium. The buds of the Bryozoa usually grow from near to a

rosette-plate, the position of bud and polypide being thus affected

by the position and number of these plates. In S. magnilabris,

and probably generally in the genus, there are two large distal

rosette-plates. In decalcified preparations a curious large fleshy

* We must keep in mind that in the Cheilostomata the tentacular sheath is

fastened to the operculum and to the zowcial wall.

+ “ Morph. of the Cheilostomata,”’ Quart. Journ. Micr. Sc. n. s., vol. xlvi. p. 318

(1902).

¢ This was not readily made out at first, but some thick sections stained in

Chlorazol blue enabled me to see it all clearly. This is a stain that will be found

useful for staining muscles and some other tissue, but it is not a good nuclear stain.

Material can be stained and decalcitied at the same time, as acid does not affect the

colour and it may be used for intra vitam staining. This stain was given to me by

my friend Mr. Waddington, who has been unable to find out particulars as to its

constitution.

§ Loe. cit. p. 320.

|| It is interesting to find that the large “pores” in the cryptocyst of Cupularia

oweni Busk are only for the passage of depressor muscles.

BRYOZOA FROM ZANZIBAR. 501

structure is found in connection with them (fig. 16). On the

inner side (towards the older zocecium) there is a large saucer-

like portion with a fleshy club-like projection on the other. ‘The

wall of the zocecium and the rosette-plate passes between the two.

From the tissue round one of these rosette-plates the polypides

of the growing parts start, but many of the polypides have the

retractor muscles always attached to the lateral wall. The ten-

tacular part of the bud seems to be growing in one distal corner

while the gut parts are growing in the other, and in the mature

polypide a long tube connects the two (fig. 12). This narrow tube

connecting the two parts is a character of the mature polypide,

and is somewhat like that of the cesophageal tube of the

Ctenostomata.

There are very few secondary buds to be seen (as the zocecia

nearly all contain perfect polypides) ; they, however, arise from

tissue on the membrane crossing the operculum.

There are about 25 tentacles, four of which near the base are

larger and broader than the others.

In other species of Steganoporella there is also considerable

variation in the teeth of the opercula ; for example, in a specimen

of S. buskii H. from Algoa Bay, South Africa, in the British

Museum, there is a B operculum without any teeth whatever,

whereas the other opercula have small teeth.

Steganoporella is not yet known before late Tertiary, whereas

Thalamoporella is much older.

Loc. Brazil; Jamaica; St. Vincent; Florida; China Seas ;

Singapore ; Philippine Islands; Trincomalee ; Amirante Isl., 20-

25 fath. (Thornely). Wasin, Brit. East Africa, 10 fath. (500)

(520); Chuaka, Zanzibar Channel, 2 fath. (508), collected by

Crossland.

Fossil. Australian Tertiaries.

CRIBRILINA RADIATA Moll.

Loc. Cosmopolitan. Ras Osowamembe, Zanzibar Channel,

10 fath., on Adeonella platalea, collected by Crossland.

HippotTHoa DIVARICATA Lamouroux.

Loc. Wasin, Brit. Hast Africa, 10 fath. (520), on shell, collected

by Crossland.

Hipporuoa pistans MacGillivray.

For synonyms see Waters, ‘“‘ Bryozoa,” Résultats du Voyage du

S.Y. Belgica, p. 54 & pl. i. fig. 8 (1904).

Loc. Wasin, Brit. East Africa, 10 fath. (520), on shell, collected

by Crossland.

SCHIZOPORELLA UNICORNIS Johnston.

Waters, “ Bryozoa of the Sudanese Red Sea,” Journ. Linn.

Soc., Zool. vol. xxxi. p. 143, pl. xu. figs. 12, 13 (1909).

Some pieces are either one or two layered, and here again I

Sie

otk

502 MR, A. W. WATERS ON

have been able to see the zoccia superimposed, so that both lateral

and distal walls of the upper layer are above the similar walls of

the lower layer *. Reuss has shown the same thing in Cumul-

pora angulata v. M., and I have in previous papers referred to it

in Melicerititide.

In some other species of Schizoporella the superimposed layers

grow quite differently (see p. 504).

Loc. Add: “ From bottom of s.s. ‘ Juba,’ which always remains

in Zanzibar waters” (511), collected by Crossland.

SCHIZOPORELLA PERTUSA Hsper.

See Miss Jelly’s Catalogue, and add :—

Lepralia pertusa Calvet, “‘ Bry. Mar. de Cette,” Trav. Inst. de

Zool. de l'Université de Montpellier, ser. 2, vol. 1. p. 515; ‘* Bry.

Mar. de Corse,” op. cit. vol. ii. p. 26; Jullien & Calvet, “ Bry.

prov. des Camp. de l’Hirondelle,” pp. 69, 134.

Schizoporella pertwsa Calvet, ‘‘ Exp. Sc. du Travailleur et du

Talisman,” p. 416.

Loe. British; Labrador; Atlantic; Florida; Mediterranean ;

Mazatlan; Australia; Samoa; China Seas. Wasin, Brit. Hast

Africa, 10 fath. (500) (520), collected by Crossland.

SCHIZOPORELLA NIVEA Busk. (Pl. LXX. figs. 1-3, 7-9, &

P]. LXXIII. fig. 16; and text-fig. 80.)

Schizoporella nivea Busk, Zoo]. Chall. Exp. vol. x. pt. xxx.

p- 163, pl. xvii. fig. 1 (1884); Philipps, in Willey’s Zool. Results,

pt. iv. p. 440 (1889); Thornely, Ceylon Pearl-Oyster Fisheries,

vol. iv. p. 114(1905); Rec. Indian Mus. vol. i. pt. 3, no. 13, p. 189

(1907); Waters, Rep. Mar. Biol. of the Sudanese Red Sea,

‘“‘ Bryozoa,” Journ. Linn. Soc., Zool. vol. xxxi. p. 168, pl. xvii.

figs. 2-4 (1909); Thornely, ‘“‘ Mar. Poly. of the Indian Ocean,”

Trans. Linn. Soc., Zool. vol. xv. p. 148 (1912).

Zoarium in Hemescharan form. Zocecia quadrate, distinctly

separated, fairly large pores over the surface, an avicularium

near each upper corner with broad triangular to nearly semi-

circular mandibles, sometimes a small avicularium at one of the

lower corners, Below the aperture there is frequently, in the

older zocecia from (501), a mucro which may be long spreading

out at the ends, or there may be a thin lamina radiating in four

directions the whole length of the mucro. ‘These mucros are

more frequent on the ovicelligerous zocwcia. The oral aperture

is nearly circular, has a wide poster, and there are fine lines

across the operculum (fig. 2).

The ovicell is large, raised, globular, with a ridge from the

middle of the proximal edge, shortly bifurcating to each side, at

any rate in the older zocecia. ‘The pores over the surface of the

* Reuss, ‘“‘ Zur Fauna des deutschen Oberoligocans,” pt. i3.,Sitzb. d. k. Akad. der

Wissensch., Wien, math.-nat. Cl., vol. 1. Abth. 1, p. 644 (81), pl. ix. fig. 1 (1864) ;

“Die Foram., Anth. und Bryozoen des deutschen Septarrenthones,” Denk. s. d.

k. Akad. der Wissensch., Wien, math.-nat. Cl., vol. xxv. p. 179 (63), pl. vin. fig. 12

(1865).

BRYOZOA FROM ZANZIBAR. 503

ovicell are fairly large, though slightly smaller than those

of the zocecia.

The small avicularium at the distal corner (figs. 7-9) has short

and wide glands which at the lower part are joined together,

and no other avicularian glands yet seen quite resemble these

(text-fig. 80). Avicularian glands occur in Lepralia foliacea Hl.

& Sol., ZL. clivosa Waters, L. margaritifera, Smittia trispinosa

Johnst., Porella plana Hincks, P. acutirostris Smitt, Retepora

cellulosa L., and other Retepore. In all these cases there are also

oral glands. The oral glands of S. nivea are but small with the

end cells the larger and darker (Pl. LX_XIII. fig. 16, g/.).

Text-fig. 80.

Avicularian glands of the small avicularia of Schizoporella nivea.

A, the two glands separated, X 320. B, the same, X 100.

C, glands united, xX 320. D, gland, X 320.

There are 16 tentacles, no pore-chambers, but on the lateral

walls there are about eight rosette-plates near the basal wall, also

on the distal wall there are about six situated near the base.

The internal membrane of the zocecium projects in a sac-like

manner into the ovicell (fig. 16). This sac is about the width of

the zocecium, and is provided with a large number of muscles

(fig. 16, sc.) radiating from near the operculum. From a whole

preparation it seems that the ovum passes into this sac, which

is then ruptured (fig. 16, 7) to allow of the passage of the large

ovum into the ovicell. I think there is no doubt of my inter-

pretation, although a ruptured membrane has only been seen in

one ease. ‘The ovarium consists of a number of large ovarian

cells.

Some of the older zocecia are closed by a calcareous layer over

the operculum, with a calcareous bar or lamina, extending from

side to side (Pl. LXX. fig. 3) curving round to the sinus, a position

which is fairly constant, not accidental. In older zoccia there is

sometimes a calcareous bar straight across the aperture, and there

504 MR, A. W. WATERS ON

may be no operculum remaining underneath it. Frequently a

tubular projection occurs in the middle of the supra-opercular

calcareous layer, and there is a closure of the same kind in Schizo-

porella unicornis and other Schizoporellide and various other

genera besides some Tertiary fossils; and a similar structure has

been described in ‘“ Lepralia’’* syringopora Rss. and is found in

various Adeonide 7. It can be seen in SN. nivea that these raised

ledges across the operculum are the basal lines for a superimposed

layer even though the layer is seldom completed; and a South

African species which, though a larger and distinct form, is very

similar to S. nivea, explains the growth across the operculum

more fully. In it we have the commencement of a superimposed

new layer of zocecia, and in one specimen there are about fifty

zocecia together on which the walls for another layer are all

mapped out, and the wall passes over the operculum, nearly

always with the same curved line as in S. nivea.

I described and figured a similar growth:in Meliceritites $ and

apparently it occurs in various other cases. On the other hand, I

have specimens of Schizoporella unicornis Johnst. from Cape Verde

Islands and Zanzibar, in which there are many layers, and each

following layer is formed by the new walls, nearly always

growing exactly above the walls of the older layer. Nevertheless,

there is, in some cases, a curved calcareous ridge over the oper-

culum for which there seems no object. In S. wiicornis there is

often a tubule on the calcareous closure.

Some stained sections of Adeonella contorta Mich. in which

there are superimposed layers, show this tubule as an inverted

funnel with a long tube (over the operculum) attached to the

tissues below the operculum. These closures can only be compared,

in a limited sense, with the tubules of the closures of the Cyclo-

stomata, as the operculum is unaltered and there is no perforation.

There are also membranous closures, and the subject deserves

further study. The Schizoporellide and the Adeonide are

apparently the two families most likely to throw light on the

closures of the Cheilostomata. Of course we do not find the

closures in the younger zocecia, only in the older ones. In both

these families blind cells are very frequent §.

The Zanzibar forms are not separated as varieties or species,

although the one with the tall mucro might perhaps be called var.

wasinensis (Pl. LXX. fig. 1)on this account. It may be the Schizo-

porella linearis of Hincks, Ann. Mag. Nat. Hist. ser. 5, vol. vii.

p- 159, pl. ix, fig. 2, to which there is no description or locality.

* Waters, “ North Italian Bryozoa,’ Q. Journ. Geol. Soc. vol. xlvii. p. 20, pl. iii.

figs. 3, 4 (1891).

+ Waters, “ A Structure in Adeonella contorta Mich.,” Ann. Mag. Nat. Hist.

ser. 8, vol. ix. p. 498 (1912).

+ **On Cheilostomatous characters in fossil Bryozoa,’” Ann. Mag. Nat. Hist.

ser. 6, vol. vill. p. 52, pl. vi. fig. 4 (1891).

§ Sincethe above was written Levinsen has published an important work, “ Studies

on the Cyclostomata Operculata,” D. Kgl. Danske Vidensk. Selsk. Skr. 7 R., Nat. og

Math. Afd. vol. x. pt. 1, 1912, dealing with Melicerititidee.

What he calls the opercula of these fossils I should speak of as the closures, for

BRYOZOA FROM ZANZIBAR. 505

Frequently a hydroid is growing over the surface. ‘This seems

to be a species of Clava and may be the same species as one

growing in Holoporella columnaris B., and H. pigmentaria Waters,

in which it may pass through several layers of zocecia. Sections

showed that the stolons were spread under the H. columnaris,

indicating that the growth of the Holoporella occurred over

established colonies of Clava round which the Holoporella formed

calcareous tubular walls *.

(1) S. nivea belongs to a group in which all the species

have a wide sinus or arc, the surface is perforated as is

also the ovicell, there are quite small oral glands or but very

moderate sized ones. The opercula have the muscular attach-

ment close to the border and fairly near to the distal end. This

group I referred to} as including the types of Schizoporella,

and belonging to it are S. sanguinea Norm., S. linearis Hass.,

S. harmsworthii Waters, S. awriculata Hass., S. galeata B.% ete.,

anda new species from 8. Africa. Levinsen includes several other

species under Schizoporella, but I am by no means sure that all

his species will remain in the same genus, for in (2) S. wnicornis

Johnst. and var. errata Waters, S. longirostris Hincks, S. spon-

gites, S. biaperta Mich., the sinus is much narrower, and what 1s

of most importance, the muscular attachments are some distance

from the border of the operculum. In 8S. biaperta the ovicell

has a flat area with perforations round the border.

(3) In the group S. vulgaris Moll., S. viridis Thorn. , there is a

similar operculum, and it is in part the Escharina of Levinsen.

In the same place I showed that there was a group which

might be placed in a modified Buffonella Jullien, in which the

surface of both the zoarium and the ovicell is imperforate, also

there is a small suboral avicularium, and the opercula have the

muscular attachments some distance from the border as in the

last group. It is represented by S. ridleyi MacG., S. simplex

I consider that the chitinous operculum was under the closure, and I should like to

see the name operculum confined to the movable chitinous appendage. Levinsen

on Plate i. figs. 15, 16, shows Meliceritites vielbanci d’Orb. with tubules to the

closures, such as I have seen in recent Cheilostomata, and a number of similar

closures are figured. Without there being perhaps any wide divergence of view I

should not describe these as being regenerated zoccia, for while Levinsen has de-

scribed regenerated zocecia, I have not seen anything to suggest their being common,

and further, although we know that the polypides are regenerated i in the closured

zoccia. this by no means always takes place, as, for example, in the older parts of

stems and when there is more than one layer. It therefore seems unadvisable to

speak of them as regenerated zocecia when they are frequently permanently closed.

I have suggested to Professor Levinsen that the sunk walls with few openings as in

Meliceritites magnifica (op. cit. Pl. i. fig. 7), may be comparable with the perforated

wall or closure UNDER THE OPERCULUM in Vittaticella (see page 484). ‘These are

also to be compared with the partial closure I described in MW. royana Waters,

Ann. Mag. Nat. Hist. ser. 6, vol. viii. pl. vi. figs. 2,6 (1891). Levinsen in his most

valuable work has given full figures and descriptions of the structure of Melicerititidee,

and as I believe that many analogous structures are found in the Cheilostomata, a

thorough examination of the closures of livmg forms is much to be desired.

* Waters, Report on Sudanese Bryozoa, p. 254.

t Waters, Résultats du Voyage du S8.Y. Belgica, “ Bryozon,” p. 42 (1904).

506 MR. A. W. WATERS ON

VOrb., S. rimosa Jull., S. marsupifera Busk, S. twmida H.,

S. levata H., S. levigata Waters.

It is quite clear that there are several groups that can be

separated off from the old Schizoporella, but until more living

and spirit specimens have been examined important characters

remain unconsidered.

Loc. Since I gave the localities of S. mivea Miss Thornely has

reported it from Amirante, 29 fath. ; Seychelles, 39 fath.; Provi-

dence, 50-78 fath. It was procured from Wasin, Brit. Hast

Africa, 10 fath. (501); Ras Osowamembe, Zanzibar Channel,

10 fath. (504); Prison Island, Zanzibar Channel, 10 fath. (505),

and with the large mucro from Wasin, 10 fath,, which may have

to be called var. wasinensis : collected by Crossland.

SCHIZOPORELLA MONTFERRANDI Audouin.

Lepralia montferrandi Waters, Rep. Mar. Biol. of the Sudanese

Red Sea, “‘ Bryozoa,” Journ. Linn. Soc., Zool. vol. xxxi. p. 171,

pl. xvii. figs. 15-18.(1909). To which add :—

Schizoporella pachnoides MacG. Trans. Roy. Soc. Vict. vol, xxiii.

p. 180 (1886); Prod. Zool. Victoria, dec. xix. p. 314, pl. 186,

fig. 6 (1889).

This must go to Schizoporella as now limited (see p. 505).

It is closely related to S. galeata, but as I have said they must

probably be separated.

Add:—Loc. Victoria (MucG@.). Wasin, Brit. East Africa, 10

fath. (520), collected by Crossland.

GEMELLIPORA PROTUSA Thornely. (Pl. LXX. figs. 4-6.)

Gemellipora protusa Thornely, Ceylon Pearl-Oyster Fisheries,

vol. iv. Polyzoa, p. 119, pl. iv. fig. 7 (1905).

Zoarium incrusting. Zocecia rhomboidal, raised, with pores

round the border; surface with few pores and granular. Oral

aperture clithridiate, very long, with large lateral contractions,

and the peristome frequently much raised, especially at the

side. At one side of the zocecitum directed downwards a short

wide triangular avicularium with a stout bar, and sometimes a

second roundish avicularium near to the side of the aperture.

Ovicell small, raised, globose, with very thick walls, perforated,

much open in front and not closed by the operculum.

Loc. Gulf of Manaar (7/.). Wasin, Brit. Hast Africa, 10 fath.,

received dry (520), collected by Crossland.

TRYPOSTEGA VENUSTA Norman.

Lepralia venusta Norman, Ann. Mag. Nat. Hist. ser. 3,

vol. xiii. p. 84, pl. x. figs. 2, 3 (1864).

Schizoporella venusta Hincks, Brit. Mar. Poly. p. 276, pl. xxx.

figs. 6,7 (1880): Kirkpatrick, Ann. Mag. Nat. Hist. ser. 6, vol.i.

p. 76 (1888); ‘‘ Hyd. & Polyzoa from the China Sea,” Ann. Mag.

BRYOZOA FROM ZANZIBAR. 507

Nat. Hist. ser. 6, vol. v. p. 17 (1890); Proc. Roy. Dublin Soe. n. s.

vol. vi. p. 612 (1890) : Calvet, Expéd. Scient. du Travailleur et du

Talisman, vol. viii. p. 416 (1907).

Trypostega venusta Levinsen, “Studies on Bryozoa,” Vid. Medd.

f. d. Naturh. Foren. i Kjgbenhayn, p. 23 (1902); Morph. & Syst.

Studies on Chil. Bry. p. 281, pl. xix. figs. 1a-ld, pl. xxii.

figs. 13 a-13d (1909); Norman, “ Polyzoa from Madeira,” Journ.

Linn. Soc., Zool. vol. xxx. p. 299 (1909).

Gemellipora glabra, form striatula Smitt, ‘ Floridan Bryozoa,’

pt. 1. p. 37, pl. xi. p. 207 (1873) ; Thornely, “‘ Mar. Poly. Indian

Ocean,” Trans. Linn. Soc., Zool. vol. xv. p. 149 (1912).

Gemellipora striatula MacG. Prod. Zool. Vict. dec. xiv. p, 150,

pl. 138. fig. 10 (1887).

Lepralia striatula MacG. Tr. Roy. Soc. Vict. p. 134, pl. iu. fig. 17

1882).

ene striatula Waters, Q. Journ. Geol. Soc. vol. xi.

p. 301 (1885) ; Philipps, ‘‘ Poly. Loyalty Isles, &c.,” Willey’s Zool.

Results, pt. iv. p. 440 (1899).

Mollia tuberculata VOrb. Paléont. Franc. p. 388; see Waters,

Ann. Mag. Nat. Hist. ser. 7, vol. xv. p. 6.

Lepralia inornata Gabb & Horn belongs to this group.

I should have hesitated, at present, to put this in a new genus

merely on account of the small chamber above the zowcium,

which no doubt must be considered as a vestigial avicularium,

but as a genus may be required it will now cause less confusion

to use Levinsen’s name. Jepralia turgescens Reuss, ‘“ Foss. Bry.

Oest.-Ung. Miociins,” Denk. K. Akad. der Wissensch. Wien,

math.-naturwiss. Class., vol. xxxiii. p. 36, pl. viii. fig. 7, has an

avicularium, which, judging from the figure, corresponds to

the small chamber above the zoccium in 7’. venusta, and also

the avicularium of Chorizopora brongniartti Aud. is often

sunilarly situated.

Sections of the specimen from Wasin show that there are no

bands of muscles in the chamber above the zocecium, but there

are protoplasmic bands to the frontal disk, starting from two

rosette-plates at the proximal end of the small chamber, and

near the distal end there is a small group of cells at each side,

which must probably be considered as glands. There are a few

irregular large chambers almost as large as the ordinary zocecia

closed in the same way as these small supra-zocecial chambers.

There are but few ovarian cells in the ovarium, usually two,

sometimes three or four, and one may be seen developed to a

considerable size. The embryos develop fully in the ovicell.

No suboral glands have been found.

Loc. British; Guernsey; Calvados; Florida; Madeira (WVor-

man); Azores; Cape Verde Islands (C.); Lifu, Loyalty Isl. (Ph.);

China Seas (Tizard Bank), 27 fath.; Torres Straits (A’.); Amirante,

22-85 fath., Saya de Malha, 29-125 fath. (Zhornely); Mauritius.

Wasin, Brit. Hast Africa, 10 fath. (500), collected by Crossland.

Fossil. River Murray Clifts, South Australia.

508 MR. A. W. WATERS ON

ARTHROPOMA CECILIZ Audouin.

Flustra cecilia Aud, ‘ Zool. Egypte,’ p. 66 (239), pl. viii. fig. 3 ;

see Miss Jelly’s catalogue.

Arthropoma cecilii Levinsen, Morph & Syst. Studies on the

Cheil. Bry. p. 332 (1909).

Schizoporella cecilit Thornely, ‘* Mar. Poly of the Indian Ocean,”

Trans. Linn. Soe., Zool. vol. xv. p. 147 (1910).

In my Report of the Antarctic Bryozoa from the Voyage du

S.Y. Belgica, p. 50, I called attention to this group and now

Levinsen has given ita name. As I previously said Phonicosia

Jullien may belong here, but Jullien’s specimens are not suf-

ficiently complete to settle the question.

Loc. Distribution general in the north temperate zone, the

tropics, and Australia. Prison Island, Zanzibar Channel, on the

shore (513), collected by Crossland.

fossil. Kuropean and Australian Tertiaries.

OSTHIMOSIA ZANZIBARIENSIS, sp. n. (PI. LX XIII. figs. 9, 15.)

Zoarium incrusting seaweed, small, irregular. Zocecia sub-

globose, smooth. Oral aperture with distinct sinus, otherwise

nearly circular, a small semicircular avicularium at each side of

the aperture, and when there is an ovicell usually a second pair

of avicularia. Surface of the zocecia imperforate with longi-

tudinal ribs in the older zocecia, vicarious avicularia among the

zocecia having expanded spatulate mandibles. Ovicell large, wide,

globose, perforated all over.

In having the perforated ovicells this is like Cellepora avicularis

Hincks, to which it is closely allied, but the median avicularium

is absent. A perforated ovicell occurs in C. avicularis H.,

C’. coroncpus B., C. megasoma MacG., C. conica Busk, C. re-

douti* Aud. In the early stages this form resembles Schizo-

porella biaperta Mich., but vicarious spatulate avicularia are

unknown in SS. biaperta. The operculum is thinner than that

of C. conica, and the walls also are thinner. I find that C. conica

is the young form of a species determined by Busk as (’. simo-

nensis ~ B.,an erect cylindrical species. The oral aperture of both

C. conica and C. avicularis is slightly larger than that of this

species from Prison Island.

After removing from Cellepora the genera Holoporella and

Lagenipora there is the present group with a triangular sinus,

the ovicell perforated and the surface of the zocecium imper-

forate except near the border, and the group includes C. avicularis

H., C. conica Busk, C. coronopus S. Woods, leaving still another

group with imperforate ovicells, but a small semilunar area or

mark near the proximal part, and a marked sinus in the oral

aperture, as, for example, C. evewa Jull., C. eatonensis B.

* This is only a variety of C. avicularis H. I have a specimen from a sounding-

line in the Red Sea.

+ Waters, Zoo]. Chall. Exp. vol. xxxi. pt. Ixxix. p. 85 (1889).

BRYOZOA FROM ZANZIBAR, 509

Hineksin his ‘ Brit. Mar. Polyzoa’ says that the typeof Cellepora

was pumicosa, but this is not the case, for as Leyinsen points out in

his large work, Linnzeus refers to C. ramulosa as the first species

of Cellepora. Hincks incorrectly considered it the genus of

Fabricius. Levinsen is, however, mistaken in supposing that

C. ramulosa is schizostomous, as it belongs to the holostomatous

group, though not to Holoporella*. I have not as yet had any

opportunity of cutting sections of 0. ramulosa, but the operculum

has the proximal border but slightly curved, and the small

muscular attachments are close to the thicker part of the border.

In most Holoporelle the attachment is a little nearer to the

edge, on the other hand it is much closer to the distal edge than

is usual in the Schizostomous groups.

The ovicell of ramulosa is cap-like as in Holoporella ete., and

although Hincks says perforate or imperforate, none of the speci-

mens in my collection, nor any in the British Museum general

collections, including Busk’s and Hincks’s,’ have a perforate

ovicell ; also in the Norman collection a few from each locality

were examined without finding any perforate. There are two or

three species externally corresponding with C’.. ramulosa, so that

a mistake is easily made, and in three cases friends have sent me

specimens so marked, of which only a part were ramulosa.

By taking (even if provisionally) C. ramulosa as definitely

described by Hincks, and perhaps by some before him, as the

type of Cellepora, we get out of a difficulty, for when Schismopora

was created by MacGillivray Cellepora remained for the holo-

stomous division. C.ramulosa is the first of Linnzeus’ species,

the others being spongites, punvicosa, ciliata, hyalina. Linneeus’

description of femmes would do for several species of branching

forms, and we are doubtful what the other species weve meant

for. In Linneus’ copy of the 12th edition of Syst. Nature,

there is in his small writing, under C. ramulosa, a reference

SOLOS © lemme One which was hieroglyphic to me until

Dr. Daydon Jackson kindly explained that it referred to Det

Kongelige Norske Videnskabets Selskabs Skriften, 4th part,

1768-1774, in which there is a paper by Gunnerus, who supplied

Linneus with both C. ramulosa and C. pumicosa. In this work

(pl. i. fig. 6) is a figure of a Cellepora, which, judging from the

locality, “Oceano Norvegico,” is probably t C. incrassata Sm.,

although C. coronopus 8. Woods, an entirely different species

from the Mediterranean, corresponds equally well with this

figure, which shows nothing but zoarial shape. It is, however,

what Linneus described as C. ramulosa. On the same plate

the figure 7 shows a_ similar growth, though with smaller

branches, and to this under C. pumicosa Linneus refers by a

* There are various specics of holostomous Bryozoa which do not belong to

Holoporella Waters, as, for instance, C. sardonica Waters, which will fall into

Holoporellidee.

+ Waters, “ Bry. from Franz-Josef Land,” Journ. Linn. Soc., Zool. vol. xxviii.

p. 94 (1900).

510 MR. A. W. WATERS ON

similar note. A long time ago I showed* that the C. pumi-

cosa as we have understood it had nothing whatever to do with

pumicosa I., as it does not in any way correspond with

Linneus’ description, besides which he refers to a figure by

Marsigli which probably is a figure of C. coronopus. I then

referred to it as C. pumicosa Busk (non L.) and have continued

to do so. Fig. 7 may be a figure of C. ramulosa Hincks, ete.

though called pumicosa by Linnzeus; there are, however, several

other species that it might represent. After this record of mis-

takes about two well known species, showing how little we can

know what the earlier writers meant, we should be allowed to

return to our senses, and use these long established specific

names for thoroughly described and well recognised species,

namely C. pumicosa Busk, C. ramulosa Hincks, but if we retain

Cellepora for ramulosa the C. pumicosa Busk becomes Osthimosia

Jullien. This group was divided off in the same year by Jullien

as Osthimosia, and as Schismopora by MacGillivray, but it seems

that Juljien’s name was published a few months the earlier. The

opercula of this group are all of the same type, with the opercular

attachment some distance from the border of the operculum.

The group is left by Levinsen under Cellepora, through mis-

understanding C. ramulosa. The ovicell of Osthimosia is, at any

rate, nearly always punctured.

This leaves the Cellepora of Hincks divided into Holoporellidee

(holostomatous), and the schizostomatous forms into Osthimosia

and Lagenipora.

Loc. Prison Island, Zanzibar Channel, 8 fath. (505); Ras

Osowamembe, Zanzibar Channel, 10 fath. (504), collected by

Crossland.

LAGENIPORA ROTA ee eis

C ellepor a rota MacG. Trans. Roy. Soc. Vict. vol. xxi. p. 116

(11) pl. i. fig. 6 (1885) ; Prod. Zool. Vict. dec. xv. p. 184, pl. 148.

fig. 3 (1887).

Levinsen would call this Siniopelta, but it is what I have

previously put under Lagenipora, as the position and character of

the ovicells of L. socialis Hincks seem to me to be the same as in

the group which Levinsen calls Siniopelta. As Levinsen has not

agreed with me, I have also, besides again examining the British

Museum specimens, through the kindness of Professor Hickson,

examined the specimens from Miss Jelly’s collection in the

Victoria University Museum. Miss Jelly first found ZL. socialis

in Hastings, and presumably all known Hastings specimens were

collected by her. The Hastings specimens in the Victoria

Museum did not show the ovicell, ‘but one sonamed by Miss Jelly,

from Guernsey, has some ovicells and also shows the spinous

processes well.

The ovicell of the Guernsey specimen is situated on the wall

* Waters, “ Bryozoa of the Bay of Naples,’ Ann. Mag. Nat. Hist. ser. 5, vol. iii.

p. 198 (1879).

BRYOZOA FROM ZANZIBAR, Bt I

of the peristome, which frequently extends beyond it as a com-

plete tube, and the ovicell has an area which is flat or slightly

rounded and is surrounded by a ridge, while at each corner

inside the ridge there isa pit. It seems justifiable to call the

part surrounded by the ridge an area, but if there is any

objection to this it might be called a tabula. In this case the

‘“‘ calcareous base” is very slightly developed and might be over-

looked, and except that the Guernsey specimen is slightly

granular or rather nodulated there is but little difference

between it and Lagenipora lucida Hincks from Madeira, except

that Z. Jucida has pores round the area as figured by me*, but

I have not the opportunity of now comparing any ZL. lucida

having ovicells.

Now in Lagenipora nitens MacG. from Port Phillip Heads,

which I considered only a variety of lucida, the ovicell is just

the same shape and in the same position, while close to the ridge

of the area there is a row of pores; also L. boryii Aud. from the

Mediterranean has a row of pores in the same place, whereas in

L. rota MacG. and some other species there are radiating grooves.

An Australian species of Lagenipora has the area evenly perforated

all over. The pores or pits of ZL. socialis, though difficult to

trace, are no doubt small and close to the edge. When JI first

thought I saw pores it was puzzling not to find a complete row,

and there was a doubt whether they might only result from

accidental damage, but no doubt the explanation is that there is

only one at each corner, . There must be something wrong if

Lagenipora socialis and L. lucida ave placed in different genera.

Loc. Victoria (MacG’.). Wasin, Brit. East Africa, 10 fath.

(507), collected by Crossland.

HASWELLIA AUSTRALIENSIS Haswell.

Myriozoum australiense Haswell, Proc. Linn. Soc. N.S. Wales,

vol. v. pt. 1, p. 43, pl. iii. figs. 9-11 (1880).

Haswellia australiensis Busk, “ Polyzoa,” Zool. Chall. Exp. vol. x.

pt. xxx. p. 172, pl. xxiv. fig. 9 (1884); Kirkpatrick, Proc. Roy:

Dublin Soe. vol. vi. p. 612 (1890); Meissner, M., aus Semon

“Zool. Forsch, Reisen in Australien und Malay,” Jena. Denkschr.

vol. vill. p. 731 (1902); Levinsen, Morph. & Syst. Studies of the

Cheil. Bryozoa, p. 297, pl. xvi. figs. 2 a—2 6 (1909).

Porina coronata var. labrosa Waters, Zool. Chall. Exp. vol. xxxi.

pt. lxxix. p. 32 (1889).

Levinsen places this in Myriozoide t, but it seems somewhat

doubtful whether the genera grouped together by Levinsen will

remain in the same family (see p. 520).

The series of zocecia in H. australiensis are all on the same

horizon, and usually at the beginning of each branch there are

six zocecia in a whorl, there is then another whorl with six, then

one with eight when a bifurcation takes place, and then again

* Journ. R. Mier. Soc. 1899, p. 18, pl. ili. figs. 25, 27, 30 (1899).

+ Loe. cit., see page 296.

512 MR. A. W. WATERS ON

each of the new series commences with six. H. australiensis has

19-21 tentacles. Unfortunately none of the specimens now

examined have ovicells. The //aswellia group is well represented

in the North Italian Tertiaries, and although some appear to be

almost identical with living species, it seems better to keep them

distinct, as all the structures cannot be compared.

There is over the surface an outer membrane, and then under

this there are membranes of the two walls of the shell showing

when decalcified the position of the pore-tubes, Some specimens

from Wasin have a pink colour.

The operculum of the species and of the H. coronata H. or

H. gracilis, whichever we may have to call it, is identical, having

the opercular attachment very high up and elongate (see my

figure in Ann. Mag. Nat. Hist. ser. 5, WO, so TO, Wik)s Ine

operculum of H. auriculata B. has the muscular attachments much

lower down and not so near the edge ; further, the ovicell which is

but slightly raised, has a semicircular area, and the surface of

the zocecium has not poresall over. It seems exceedingly doubt-

ful whether H. auriculata should remain in Haswellia, but I have

never had sufficient material in my hands for a complete

examination. H. grandipora Waters is Haswellia, and I now

think must stand as a species.

Loc. Holborn Island, Queensland (asw.); near Torres Straits

(Chall.); Thursday Island, Torres Straits (Jeissner); Formosa

Channel (Levinsen). Wasin, Brit. Hast Africa, 10 fath. (500,

501, 507, 520), collected by Crossland.

T'UBUCELLARIA CEREOIDES, var. CHUAKENSIS Waters.

Tubucellaria cereoides var. chuakensis Waters, ‘ Tubucellaria,”

Journ. Linn. Soc., Zool. vol. xxx. p. 130, pl. xv. figs. 10-13, 18,

19, pl. xvi. figs. 20-25 (1907); Thornely, “Mar. Poly. Indian

Ocean,” Trans. Linn. Soc., Zool. vol. xv. p. 146 (1910).

Tubucellaria fusiformis Busk (non d’Orb.), Zool. Chall. Exp.

vol. x. p. 100 (1884).

Loc. Torres Straits (Busk); Grahamstown, 8. Africa; Provi-

dence, 50-78 faths. Chuaka, Zanzibar, 3 fath. (512, 524);

Wasin, Brit. E. Africa, 10 fath. (501); Chaki-Chaki, Pemba

Island (517), low water, collected by Crossland.

TUBUCELLARIA FUSIFORMIS d’Orbigny.

Waters, loc. cié. p. 131, pl. xv. figs. 1, 2, 3, 14.

Loc. Malacca, Ainirante Is., Chuaka, Zanzibar, 3 fath. (528) ;

Wasin, Brit. East Africa, 10 fath. (520), collected by Crossland.

TUBUCELLARIA ZANZIBARIENSIS Waters.

Waters, loc. cit. p. 131, pl. xv. figs. 4-7; Thornely, “ Mar, Poly.

Indian Ocean,” Trans. Linn. Soe., Zool. vol. xv. p. 146 (1912).

Loc. Saya de Malha, 145-150 faths.; Farquahar Reef ;

BRYOZOA FROM ZANZIBAR. ils

Cargados, 28 fath. Wasin, Brit. East Africa, 10 fath. (501) ;

Ras Osowamembe, Zanzibar Channel, 10 fath. (504, 514); Prison

Island, Zanzibar Channel (505); Chuaka Bay: collected by

Crossland.

SMITTINA TRISPINOSA, var. PROTECTA Thornely.

Smittia trispinosa, var. protecta Vhornely, Ceylon Pearl-Oyster

_ Fisheries, vol. iv. Polyzoa, p. 123 (1905); Waters, “ Mar. Biol.

of the Sudanese Red Sea,” Journ. Linn. Soec., Zool. vol. xxxi.

p. 173, pl. xvii. figs. 5, 6 (1908).

Smittia nitida Hincks, Ann. Mag. Nat. Hist. ser. 5, vol. vii.

p. 159, pl. ix. fig. 5 (1881).

There are two specimens with the large avicularia situated

diagonally distal to the oral aperture. The ovicells of this variety

and of typical nitida are similar, and there are some specimens

with a large, almost spinous, process at the proximal part of the

peristome, with a similar process on the ovicell just distal to the

area of pores. An identically similar form occurs off the Cape

Verde Islands, and Osburn * menticns an umbo in some forms of

nitida “behind the orifice,” and he shows how WS. trispinosa var.

nitida is subject to great variation with regard to the avicularia

and the peristome.

Loc. Gulf of Manaar (7h.); “ Africa” (J.); Red Sea (W.).

Wasin, Brit. E. Africa, 10 fath. (520), collected by Crossland.

SMITTINA TRISPINOSA, var. SPATHULATA MacGillivray.

Waters, “ Mar. Biol. Sudanese Red Sea,” Journ. Linn. Soc.,

Zool. vol. xxxi. p. 156.

The ovicell has a distinct area, the border of which sometimes

rises aS an umbo, or is divided into two sharp, erect processes.

S. trispinosa var. bimucronata Hincks belongs to this group, but

it is doubtful whether it should be separated asa variety, as there

is normally such considerable variation in the zoecia. From

(504) there are very large vicarious avicularia, directed either

distally or proximally.

Smittina is used instead of Smittia, though not including all

that Levinsen refers to it, for I consider it a group in which the

operculum is usually very thin, almost membranous, with the

lower edge straight. In the species so far examined, the oral

glands are quite small and usually more or less attached to the

tentacular sheath. In the aperture a lyrula is usually found with

the operculum under the cardelle but over the lyrula. Psewdo-

flustra solida Stimp., and Lepralia pallasiana, together with its

allies, show many differences and do not seem to belong here.

Loc. Bass's Straits; Torres Straits; Red Sea, Wasin, Brit. E.

Africa, 10 fath. (520); Ras Osowamembe, Zanzibar Channel,

10 fath. (504), collected by Crossland.

* “The Bryozoa of the Woods Hole Region,” Bull. Bur. of Fisheries, yol. xxx.

p. 246 (1912).

514 MR. A. W. WATERS ON

SMITTINA TROPICA Waters. t

Smittia tropica Waters, “ Mar. Biol. of the Sudanese Red

Sea,” Journ. Linn. Soc., Zool. vol. xxxi. p. 174, pl. xvii. figs. 10-14

(1909).

Loc. Red Sea. _ Wasin, 20 fath. (520), collected by Crossland.

SMITTINA Sp.

There is a small piece of Smittina encrusting ddeonella platalea, —

which has the peristome very much raised, especially at the distal

part, and there is apparently a triangular avicularium in the lip.

There are perforations round the border of the zocecium, and the

small ovicell is not much raised and is at the base of the peristome,

and at each side of the ovicell close to the edge there is a small

clear spot. There are two openings in the ovicell of S. oculata

MacG., and sometimes others; and I find in a specimen sent me

by Jullien as 8. longirostris J., there is a similar spot at the side

of an ovicell. Jullien’s species has large frontal avicularia,

whereas none are found on the small fragment. The lyrula is at

first very narrow but spreads out widely on each side. I hesitate

to give it a name until better specimens are found.

Loc. Ras Osowamembe, Zanzibar Channel, 10 fath. (504).

LEPRALIA FEEGENSIS Busk. (Pl. LXX. figs. 21, 22.)

Lepralia feegensis Busk, Zool. Chall. Exp. vol. x. pt. xxx.

p. 144, pl. xxii. fig. 9 (1884); Philipps, in Willey’s Zool. Results,

pt. iv. p. 446, pl. xl. fig. 7 (1889); MacGillivray, Proce. Roy.

SocwVaets mass) vole iis p: Sl, plo ox. shlosh a(S oi) seh onmelive

Ceylon Pearl-Oyster Fisheries, vol. iv. p. 121 (1905); Rec. of

Indian Mus. vol. i. pt. 3, no. 13, p. 190 (1907); “ Mar. Polyzoa

Tndian Ocean,” Trans. Linn. Soc., Zool. vol. xv. p. 150 (1912).

Hippopodina feegensis, Levinsen, Morph. & Syst. Studies on

the Cheil. Bry. p. 353, pl. xxiv. figs. 3 a—3 f (1909).

Miss Philipps described and figured the ovicell as with opaque

granulations, globular and depressed ; and Miss Thornely, not

aware that the ovicell had been seen, described it as with large

circular pitted areas punctured in their centres. In a specimen

in my collection, from “ Singapore or Philippines,” the ovicell has

the pores fairly similar to, and about the same size as, the pores

of the zocecia. MacGillivray had previously described the ovicell

of his LZ. feegensis, but as the avicularium, which is directed down-

wards, is in quite a different position, I doubt whether it is the

feegensis of Busk. He described the ovicell as large, rounded,

prominent, and marked similarly to the zocecia, In the specimens

from Zanzibar (501) there are no ovicells, glands, ovaria, or

testes, but in those from (511) there are many ovicells which are

perforated, though when the ovicell contains an ovum or embryo

these perforations look dark and opaque.

There are about 26 tentacles, and the operculum closes the

ovicell. There ave several ovarian cells in an ovarium.

BRYOZOA FROM ZANZIBAR. 515

In the submandibular part of the avicularium there is a

hammer-shaped thick chitinous piece for attachments of muscle.

The proximal muscles are in several bundles and are attached to

the base of the tentacles.

Lepralia is a genus which we have long wanted.to see brought

into order, but certainly further work is required. Hincks made

a group Lepralia, and, no doubt, on the whole the species of his

group are closely related, but he unquestionably placed some

there which should have gone in his Schizoporella and other

genera. Neviani, taking species with a horse-shoe aperture,

made a genus Hippoporina, but he still retained some which

must be separated, and very little was gained by his new

name. In Lepralia the operculum is usually thick and has a

strengthening band at each side with the muscles attached near

the distal end of the band. ‘The sides of the operculum are either

straight or much contracted where the lateral teeth of the

aperture occur. This will remain a fairly large group even when

extraneous species have been removed, but we hardly know where

Levinsen would place them, as hardly any of them are mentioned.

L. adpressa, L. hippopus, L. rectilineata Hincks, find no place.

Lepralia feegensis is made the sole. representative of a new genus

Hippopodina, but the characters given seem insufficient for

separation, as there are a considerable number with the same

general characters.

Some Lepralia Levinsen places under Smittina, a new name for

Smittia, but the true Smittia to which he alludes have a very

thin membranous operculum with straight proximal edge hardly

separated from the frontal membrane, and there is usually a lyrula,

which is perhaps a structural correlation with the thin mem-

branous operculum; further, there are very small oral glands

often partly attached to the tentacular sheath. I find it quite

impossible to place Lepralia pallasiana and its allies side by side

with Schizoporella auriculata Hass., S. linearis Hass., S. triangula

Hincks, ete., as the group seems very unnatural.

Levinsen’s Cheilopora* contains species some of which have the

ovicelligerous zocecia with a quite differently shaped aperture from

that of the ordinary zowcia, as, for instance, Lepralia cirewmeinetu

Neviani, a species found fossil, but also living, from Naples,

Capri, and Oran (Algiers) ; whereas LL. longipora MacG.= L. pre-

longa Wincks, and L. prelucida H. have large raised perforated

ovicells. This dimorphism occurs in several Lepralide, but in

Schizoporellide I only remember it in S. swbimmersa MacG. We

hardly know yet how far these characters are reliable for generic

divisions.

Such forms as LZ. adpressa Busk, LZ. hippopus Sm., L. recti-

lineata H., and the bulk of what we have known as /#pralia

should remain there until ample living and spirit material of

numerous specimens has been examined.

* Haime made a genus Chilopora and Michelotti called one Cheilopora-

Proc. Zoou. Soc.—1913, No. XXXV. 35

516 MR. A. W. WATERS ON

Loc. Philippines, 18 fath. (Chall.) ; Singapore or Philippines

(A. W. W. coll.); Hong Kong; Lifu (Ph.); Manaar (7h.);

Andamans (7h.); Cargados (Ind. Ocean) (Zh.); (¢) Nichol Bay,

N. W. Australia (ZacG.). Wasin, Brit. East Africa, on Adeonella,

10 fath., from bottom of s.s. ‘Juba,’ which always remains in

Zanzibar waters (511), collected by Crossland.

LEPRALIA TURRITA Smitt. (Pl. LX XIII. fig. 10.)

Lepralia turrita Smitt, Floridan Bryozoa, pt. 1. p. 65, pl. xi.

figs. 226-228 (1873); Kirkpatrick, Ann. Mag. Nat. Hist. ser. 6,

vol. v. p. 16 (1890); Thornely, “Mar Poly. of the Indian Ocean,”

Trans. Linn. Soc., Zool, vol. xv. p. 150 (1912).

The small specimen from Wasin has four long calcareous

spinous processes, irregularly placed round the aperture. One

or more of these processes may bear an avicularium at one side

near the base, as in the spinous process of H. albirostris Sm.

Besides these avicularia there are on the surface of the zoarium

numerous small semicircular avicularia, irregularly scattered.

The ovicell is not much raised and has an elongate elliptical

opening in the front. A specimen of Lepralia turrita in my

collection has short blunt processes with similar semicircular

avicularia, and the ovicell, which is more raised, has a similar wide

opening in the front. Perhaps, on account of the different

character of the ovicell, it should at least be made a variety, but

until more material is available it is allowed to stand.

The operculum has the sides nearly straight with the proximal

edge curved ; there are two muscular dots a little distance from

the edge and two articular thickenings ; the width is about 0°2 mm.

This operculum does not correspond with any with which I am

acquainted, for though in many respects it is like those of what I

should call, ina restricted sense, Lepralia, there is no lateral ridge

for the muscular attachment, which is not close to the border.

In the British Museum there is a specimen marked turrita,

which is Holoporella, and it seems as if there were more than one

with a series of calcareous processes round the oral aperture.

What Ridley and what I called Z. twrrita may each have to be

placed elsewhere.

Loc. Ascension Island ; Florida (Sm.) ; China Seas; Amirante,

29 fath.; Cargados, 30 fath. Wasin, Brit. Hast Africa, 20 fath.

(522), collected by Crossland.

LEPRALIA WASINENSIS, sp.n. (Text-fig. 81.)

A small piece with only seven zocecia was found on re-examin-

ing some material, when the paper was almost completed. Only

the caleareous part remains, which was probably bilaminate, but

this cannot be stated with certainty. The zoccia have the sides

straight and the distal end somewhat rounded, with thick borders

to the zoecia. The oral aperture is subrotund with a wide

curve on the distal end surrounded bya thick band. The frontal

BRYOZOA FROM ZANZIBAR. 517

surface is closely pitted, and at each side of the oral aperture,

somewhat below it, there is a large subtriangular chamber which

is probably avicularian. In the band surrounding the oral

aperture there is an opening at the proximal end which appears

to be avicularian.

Text-fig. 81.

Lepralia wasinensis, sp.n. X 25

The species seems most nearly related to Lepralia (Mucronella)

prelucida Hincks, which has a large avicularium by the side of

the oral aperture as seen in my specimens, and as Professor R.

Osburn * mentions in his specimens from Labrador.

Loc. Wasin, Brit. East Africa, 20 fath. (520), collected by

Crossland.

LEPRALIA CLEIDOSTOMA, var. INERMIS Ortmann.

Lepralia cleidostoma, var. inermis Ortmann, ‘“ Die Japanische

Bry.,” Archiv fiir Naturgesch. vol. i. p. 49 (1890).

Lepralia cleidostoma, var., Hincks, Ann. Mag. Nat. Hist. ser. 5,

vol. xii. p. (41) (1884).

This form has the surface smooth and silvery with frequently

an umbo below the aperture and often one on the ovicell. The

ovicell is scarcely at all raised and is also smooth and imperforate,

and has a small semilunate area. There is sometimes a small

knob on each side of the aperture. Both the aperture and the

operculum are about the same size and form as that of LZ. cleido-

stoma Smitt from Madeira 7, which also occurs at Florida (Sm.),

Japan (Ort.), and Bermuda (my coll.) ; but L. imermis var. differs

in having no avicularia, and it is a question whether it is advisable

to consider it a species or a variety.

Norman says ¥¢ that L. cleidostoma is the L. porcellana Busk, but

* “Bryozoa from Labrador, etc.” Proc. U.S. Nat. Mus. vol. xliii. p. 283,

pl. xxxiv. figs. 3, 3a, 3c (1912).

+ Waters, “ Bryozoa from Madeira,’ Journ. R. Micr. Soc. 1899, p. 10, pl. iii.

fig. 16.

2; “Polyzoa of Madeira,” Journ. Linn. Soc., Zool. vol. xxx. p. 305, pl. xl. figs. 1, 2

(1909).

35*

518 MR. A. W. WATERS ON

as it has been impossible to recognise it from Busk’s figures the

name cleidostoma must stand.

Loc. Queen Charlotte Island (Hincks); Japan, 100 fath.

(Ortmann). Wasin, Brit. Kast Africa, 10 fath. (520), collected by

Crossland.

PETRALIA JAPONICA Busk,

Lepralia japonica, Waters, Mar. Biol. of the Sudanese Red Sea,

Journ. Linn. Soc., Zool. vol. xxxi. p. 149, pl. xiii. figs. 10-12.

Loc. Add (Natal, as Hemeschara gigantea Busk MSS. in the

Brit. Mus.) Wasin, Brit. Hast Africa, 10 fath. (501), collected by

Crossland.

PETRALIA CHUAKENSIS, sp. n. (Pl. LXX. figs. 10-14.)

Specimens from Chuaka correspond in most particulars with

Busk’s description of Lepralia dorsiporosa *, but the notch at the

distal end of the zoccium is wanting, and in chuakensis the

proximal edge of the aperture is serrate. The avicularium

at the side is curved and is larger than those figured by Busk.

The Zanzibar specimens have ovicells which are unknown in

dorsiporosa, and the wall, which is not much raised, is finely

perforated.

The perforated area in the dorsal surface is for a broad radicle,

which, however, is not always developed ; a similar area occurs in

Petralia vultur var. armata, and also in Petralia japonica there

is often a similar large radicle. Levinsen refers to the radicle

tube in P. dorsiporosa. There is a complete bar to the avicularia

often disappearing in Eau de Javelle preparations. There are

numerous uni- or few-pored rosette-plates scattered over the

distal wall. On the lateral wall, about halfway between the frontal

and basal walls, there is a row of about 5 few-pored rosette-plates.

There are about 28 tentacles. In the ovarium there are but

few ovarian cells.

It has been thought best to give a specific name, although there

is much in common with P. dorsiporosa from the Torres Straits.

Loc. Chuaka, Zanzibar, 3 fath. (502), collected by Crossland.

PETRALIA vuLTUR Hincks, var. ARMATA, nov. (Pl. LXX.

figs. 15-20).

Type, Mucronelta vultur Hincks, Ann. Mag. Nat. Hist. ser. 5,

vol. x. p. 98, pl. viii. fig. 2 (1882).

From Chuaka, Zanzibar, there are unilaminate specimens

differing from the Mucronella vultur as described by Miss

Thornely 7, in having a triangular mandible to the avicularium

attached to the mucro. ‘There are also a large number of raised

round avicularia, with semicircular mandibles, and a distinct

cross-bar. These avicularia are sometimes on the top of a raised

* Zool. Chall. Exp. vol. x. p. 148, pl. xviii. fig. 4.

+ Ceylon Pearl-Oyster Fisheries, vol. iv. p. 124 (1905).

BRYOZOA FROM ZANZIBAR. 519

tube. There are a few large spatulate avicularia at one of the

distal corners, and these also have a distinct cross-bar. Speci-

mens of JZ. vultur in Miss Thornely’s collection show the mandible

serrate.

There is a fairly broad lyrula and two cardelle, and the lyrula

is formed before the front wall is complete. Only one ovicell has

been seen, and that is partly broken, but it is broad and minutely

perforated.

On the dorsal surface there are perforated spaces, no doubt for

radicles, but there are usually several small ones instead of one

large one as in Lepralia dorsiporosa B.

Similar ovicells with minute perforations occur in Petralia wi-

data MacG., P. japonica Busk, P. vultur Hincks, P. ellerii MacG.,

P. (Mucronella) magnifica Busk, P. (Lscharella) bisinuata Smitt

(non Busk), P. thenardii Aud., P. (JL) porosa Hincks, P. (JZ)

castanea Busk, L. crassa Thornely; and these Levinsen would

piace in his family Petraliide, and thus it would seem as though

a genus can be separated based largely on the ovicell. In the

species of this group which it has been possible to examine, there

are a large number of tentacles, viz., Petralia undata MacG.,

about 26; P. japonica B., about 25; P. castanea B., about 23 ;

P. vultur var. armata, nov., about 25; P. chuakensis, nov.,

about 25.

There seem to be two tentacles larger than the rest (figs. 15-17),

and these, instead of being triangular, have the inner surface

nearly straight, with a number of long nuclei. On this inner

surface there are cilia, but unfortunately the condition does not

admit of exact study of this point. These two larger tentacles

occur in all the species examined of the group, and larger tentacles

occur in other groups to which I have previously alluded. These

large tentacles are most marked near to the base of the tentacles

when they are commencing to divide, whereas nearer to the ends

there is but little difference.

Loc. Chuaka, Zanzibar, 3 fath., 29.38.1901 (506), collected by

Crossland.

? EsCHAROIDES OCCLUSA Busk.

See ¢Lepralia occlusa Waters, ‘‘ Mar. Biol. of the Sudanese Red

Sea,” Journ, Linn. Soc., Zool. vol. xxxi. p. 152, pl. xiii. fig. 15,

pl. xiv. figs. 1-9, 13 (1909).

The distal and lateral walls have numerous scattered uniporous

rosette-plates, especially near the inner borders. At the side of

the zoarium there are often two large spatulate opposite avicu-

laria, whereas in Adeonella the avicularia are usually on a median

line.

Levinsen would put this under Myriozoidea, but I hardly think

he can have had sufficient material before him when writing about

the family, and that is often our difficulty in considering classi-

fication.

520 MR, A. W. WATERS ON

The type of Myriozowm must, of course, be I. truncatwm, and

Tam not sure that any other living species can be put in the

same genus. It has a large number of tentacles, 26-27, which

should lead us to consider carefully whether the species with about

16 tentacles belong to the same genus, though we cannot conclude

from that alone that they do not.

Levinsen makes one of the characters of Myriozoidea, ‘“ de-

pendent avicularia seem always to be present,” but in the type

M. truncatum no avicularium has been found; another of his

characters of Myriozoum and Myriozoella is no transverse bar in

the avicularium, but Schizoporelia bituwrrita H. and what I have

called S. fuberosa Rss. and Hscharoides occlusa have a distinct bar.

In WM. truncatum the ovicell is a very large swelling with the

covering wall resembling the zowcial wall. S. crwstacea has a

prominent round ovicell with radiating ornamentation. S. biter-

rita H. and S. tuberosa have very large wide raised ovicells. In

M. simplex Busk no ovicell is externally apparent, though cal-

careous sections reveal a concealed ovicell. In S. polymon phe the

ovicell is also under the calcareous wall, giving, as a rule, no

external sign (see my suppl. ‘Challenger’ Rep. vol. xxxi. pl. i.

figs. 22, 23). In S. biturrita and S. polymorpha B. there is an

avicularium at each side of the oral aperture, and in J/. mario-

nensis B., at each side well within the peristome, there is a

small avicularium with a mandible rounded at the end.

Now, in ? Escharoides occlusa there is an avicularium on the

lip of the peristome at one side*, the ovicell is but moderately

raised, with a few large openings, the avicularia have a bar and

there are very large glands, at first paired, but often ultimately

uniting to form a large multilobular gland. So far as my sections

go, namely, S. crustacea, M. coarctum, M. subgracile, and M. trunca-

twm, there are no glands in the species considered as Myriozoidea

except in H. occlusa, and these are quite different from any others

known. Also the general absence of glands indicates that Myrio-

zoidea of Levinsen are not, as considered by him, closely allied to

the Reteporid, in which glands are so well developed, and which

usually have a sublabial pore and a lamina to the ovicells, also

vibices indicating two distinct layers. ‘The operculum of Z. occlusa

is quite different from that of any other species placed by Levinsen

under Myriozoidea. It has a thick border all round, and near

the distal edge are two knobs for the attachment of the muscles.

Myriozoum truncatum has a very large operculum of a shape

quite different from that of the others mentioned, the proximal

portion of the operculum being somewhat triangular with the

attachments near to the proximalend. M/. subgracile, M. coarctum,

and MM. crustacea have the sides of the moderate-sized operculum

nearly straight, with the muscle near the side. Haswellia

australiensis has the muscular attachment near the border of the

* The position of the avicularium is similar in Rhamphostomella, etc., but we do

not yet know the systematic value of this character.

BRYOZOA FROM ZANZIBAR. SPAM

operculum. J/yriozowm marionensis would seem to be Cellarinella

or related to it. Gephyrophora polymorpha has a large operculum

nearly round, with a wide vanna fitting into the poster, and the

muscular dot is some distance from the edge, in fact it is of a

Schizoporellidan character.

It seems probable that H. occlusa must be made the type of a

new genus, and I expect that the examination of material suit-

able for cutting will result in a re-arrangement of classification

of the Myriozoidea of Levinsen, as the ovicells present such great

differences. I am loth to make new genera until we can see from

sufficient material what is to be grouped together ; at any rate, I

fail to see good reason for placing Hscharoides occlusa * under

Myriozoum.

Loc. See Waters, ‘“‘Red Sea Bryozoa” p. 156. Wasin, Brit.

East Africa, 10 fath. (501) (520); Ras Osowamembe, Zanzibar

Channel, 10 fath. (504), collected by Crossland.

HoLoPoRELLA COLUMNARIS Busk.

Cellepora columnaris Busk, Zool. Chall. Exp. vol. x. pt. xxx.

p. 194, pl. xxix. fig. 11, pl. xxxv. fig. 16 (1884); Waters, Ann.

Mag. Nat. Hist. ser. 6, vol. iv. pl. il. figs. 1-6 (1889); Ortmann,

“Die Japanische Bryozoenfauna,” Archiv f. Naturgesch. vol. i.

p- 55 (1890).

Cellepora cidaris MacG. Prod. Zool. Vict. dec. xvii. p. 243,

pl. 165. fig. 4 (1888) ; Thornely, ‘‘Ceylon Pearl-Oyster Fisheries,”

vol. iv. Poly. p. 126 (1905); Records of Indian Museum, p. 195

(1907).

The oral aperture has a denticle at each side. As shown in my

paper mentioned above, the columns are often very thick, ex-

tending through several layers of the zoarium. A specimen

of Holoporella in my possession, named in manuscript celosia by

Busk, has similar columns passing through several rows of zocecia,

and the avicularian mandible has a double columella, while the

shape of the operculum also shows that it is a distinct species.

This is not, however, the species so named in the Busk collection

in the British Museum, where there are two other species named

celosia, MSS.

I have had the opportunity of comparing Busk’s types and

MacGillivray’s co-types (7. e., duplicates) in the British Museum,

and there can be no doubt as to the identity of columnaris and

cidaris.

In a specimen of colwmnaris from Wasin there are numerous

small hydroids extending far into the zoarium within, protecting

internal walls, formed by the WHoloporella, prolonged slightly

* Hincks considered H’scharoides, a name given by Milne-Edwards for a subgenus

of Cellepora, as being now the genus of Smitt, who used it for species with an

avicularium in the lip on one or both sides, but Smitt never fully diagnosed it. As

Escharoides, a name originally meaningless, has been used in a quite different sense

from Levinsen’s, it would be much better if he recalled his genus Peristomella and

used it instead of E’scharoides.

Oo, MR. A. W. WATERS ON

beyond the zoarium. Sometimes they start from the base, and

the poiypides stand out from the upper surface of the zoarium.

A similar case from the Red Sea is mentioned in the addenda to

my Report*. Probably the hydroids in both are Clava.

There are about 18 tentacles. There are a great number of

ovaria with two small ovarian cells, and in only one or two cases

has a moderate-sized ovarian cell been seen, and the ovaria are of

the Bugula type. The ovaria may be surrounded by testes.

Loc. Bass’s Straits, 38 fath. (Chall.); Port Phillip Heads

(MacG’.); Port Jackson, N.S.W. (Waters); Sagamibai, Japan

(Ort.); Manaar, 34 fath.; Ceylon Coast, 32-34 fath.; off Port

Blair, 100 fath.; St. 59, 32 fath., St. 77, 35 fath. (Zhornely).

Wasin, Brit. Hast Africa, 10 fath. (501), collected by Crossland.

HoLoporeEuuA APERTA Hincks.

Waters, ‘Mar. Biol. of the Sudanese Red Sea,” ‘“ Bryozoa,”

Journ. Linn. Soc., Zool. vol. xxxi. p. 161, pl. xvii. figs. 20-23

(1909).

Loc. Additional. Wasin, Brit. Kast Africa, 10 fath. (500); Ras

Osowamembe, Zanzibar Channel, 10 fath. (504), collected by

Crossland.

HoLoporELLA ALBIROSTRIS Smitt. (Pl. LX XIII. fig. 11.)

Cellepora albirostris, forma typica Smitt, ‘ Floridan Bryozoa,”

pt. ii. p. 70, pl. xii. figs. 234-239 (1873).

Cellepora albirostris Busk, Journ. Linn. Soc., Zool. vol. xv. p. 347,

pl. xxvi. fig. 2 (non fig. 1) (1881); Zool. Chall. Exp. vol. x. pt. xxx.

p. 193, pl. xxxiv. fig. 7, pl. xxxv. fig. 3 (1884); Waters, Quart.

Journ. Geol. Soc. vol. xli. p. 304 (1885); op. cit. vol. xlii. p. 68

(1887); Ann. Mag. Nat. Hist. ser. 5, vol. xx. p: 197 (1887);

MacGillivray, Prod. Zool. Vict. dec. xvii. p. 249, pl. 167. fig. 1

(1888) ; Mon. Tert. Poly. p. 109, pl. xiv. fig. 11 (1895); Thornely,

“Mar. Poly. Indian Ocean,” Trans. Linn. Soc., Zool. ser. 2,-vol. xv.

Oise hs fos Ltaysy (ILS) :

In HZ. albirostris there is great variation in the colour, some

being very dark with the ends of the rostra white, others may be

nearly white.

There are vicarious duck-bill avicularia and the border of the

rostral avicularia is denticulate ; the oral glands (fig. 11) are very

long, and there are 16-17 tentacles.

A specimen is partly adnate, from which bilaminate branches

arise. The unilaminate portions, when seen from below, are quite

regular with the sides of the zocecia parallel, as in so many

Schizoporellide.

Both Smitt and Busk expressed doubt as to whether //. albi-

rostris and H, bispinata were synonyms, but this is not the case,

for albirostris has a long spine below the oral aperture, while

bispinata has a short avicularium ; further, im 7. albirostris there

* Journ. Linn. Soc., Zool. vol. xxxi. p. 254, p. 168, pl. xv. fig. 16 (1910).

BRYOZOA FROM ZANZIBAR. De

isa pair of the minute avicularia at the side of the zocecium

to which I referred in my Report of the Sudanese Bryozoa, but

none were found in H. bispinata. Further, the operculum of

H, albirosiris has a muscular ridge turning sharply inwards and

ending off short, whereas the operculum of H. bispinata has a

ridge, very thick at the base, extending as a thin band nearly to

the apex. H. albirostris has a very long oral gland nearly the

length of the polypide, while the gland of H. lispinata, though

also cylindrical, is much smaller.

Loc. Florida, 25-35 fath. (Sim.); Heard Island, 35 fath. (Chall.);

New South Wales (Waters); Victoria (ZacG'.) ; Amirante, 12-18

fath. (Zhornely). Wasin, B. East Africa, 10 fath. (501); Chuaka,

Zanzibar, 3 fath. (525), collected by Crossland.

Fossil. Australia, N. Zealand (WV); Victoria (JlacG.).

MiICROPORELLA CILIATA Pallas.

Loe. Northern Seas; British; Atlantic; Mediterranean ; Red

Sea; Ceylon; Australasia and varieties in Japan, Australasia ;

Zanzibar (H.), the N. Indian Ocean. Wasin, Brit. KE. Africa, 10

fath. (500), collected by Crossland.

Fossil. Pliocene and Miocene of Europe, and Tertiary of

Australia and New Zealand.

RHYNCHOZOON PROFUNDUM, var. LAMINATUM, nov. (PI. LX XII.

Hees)

The specimens from Wasin may be the 2. profundum as first

described by MacGillivray, but it is not the profundwm of the

Prodromus, which MacGillivray considers is the same as 2. longi-

rostre Hincks.

The species of Rhynchozoon are difficult to distinguish, but the

ovicell has a lamina in the front like the lamina of Retepora, to

which it is related, and the shape of the lamina furnishes specific

characters of value. In this Zanzibar variety the lamina is long

and rounded at the end, whereas in 2. profundum it is shorter,

very broad, and spreads out at each side. In &. bispinosum Johns.

it is quite short.

In this variety as in &. profundum the denticle is broad and

uncinate, and the ovicell is but little raised above the general

surface. ‘There are narrow pointed avicularia over the surface,

as usual in Lynchozoon.

Loc. Wasin, Brit. Hast Africa, 10 fath. (520), collected by

Crossland. Noumea (in A. IW. W. coll.).

RETEPORA HIRSUTA Busk.

Retepora hirsuta Busk, Zool. Chall. Exp. vol. x. p. 119, pl. xxvi.

fig. 4 (1884); Waters, ‘‘ Mar. Biol. of the Sudanese Red Sea,”

Journ. Linn. Soc., Zool. vol. xxxi. p. 175, pl. xviii. figs. 24-26

(1909).

Specimens from (500) and (520), which are undoubtedly the

524 MR. A. W. WATERS ON

same as Busk’s ‘ Challenger’ 2. hirsuta, have besides minute cir-

cular avicularia round the fenestra, also on the inner side of each

of the zocecia bordering a fenestra, an elongate avicularium with

the beak bifurcate but with an acute mandible. This was de-

scribed by Busk for F. harsuta and is one of the most important

characters. Large pieces were examined without finding any of

the long antenniform spines described by Busk, and then in

some places they were found in abundance. This is much like

Rk. monilifera var. umbonata MacG., but in that there are only

occasionally the long bifid avicularia on the zocecia bordering on

the fenestra, and they must be separated because, among other

reasons, the branches have more zoccia in A. wmbonata; the

more delicate hirsuta is a tropical form, whereas wmbonata is

Australian.

The monilifera group of Retepora seems to be well defined, and

perhaps should have generic value given to it. All have the

well known “ trifoliate stigma” on the ovicell, and in most there

is a minute avicularium on the lip to one side; the opercula

generally are fairly similar with a nearly straight proximal edge,

and in shape rather wider than long, with the muscular attach-

ments rather high up and near the border. In &. formosa

MacG. and R. contortuplicata B., the operculum is more curved

on the proximal edge, and no labial avicularia have been found

in these; apparently all have the labial pore, which is often the

end of a long tube opening into the zocecium * proximally to the

operculum, and I agree with Jullien and Calvet in speaking of

this as the Reteporidan pore, as it is characteristic of a large

number of Retepore.

There is another group with widely open ovicells, and in these

none have a labial avicularium, unless 2. incrustata Calv. belongs

here; the proximal edge of the operculum is not straight, and in

some it is contracted in the middle like the opercula of some

Lepralia (for example L. hippopus Sm.). In this group are

R. imperati B., R. elongata Smitt, LR. tessellata Hincks, R. solan-

deria Risso, R. inordinata Calv., having a broad thickened border ;

while &. sinwosa Kirkp., R. novezelandie Waters, R. hippocrepis

Waters, R. gelida Waters, 2. lepralicides Waters, have long bands

at the sides of the operculum, for the muscular attachments, and

lateral contraction of the oral aperture. The Reteporidan pore

has only been observed in this group in &. solanderia Risso.

Another group has a fissure in the ovicell as in #. cellulosa L.

and the proximal edge of the operculum is nearly straight, and

very similar throughout this group, labial avicularia occur in

some but not in all. There are a few Retepore with imperforate

ovicells. Throughout the genus the oral glands are very well

developed.

The minute round or oral avicularia with thin membranous

* Waters, “ Medit. & New Zeal. Reteporide,’’ Journ. Linn. Soc., Zool. vol. xxv.

pp. 255-6; Expéd. Antarct. Belge, pl. vi. fig. 6 a (1904).

BRYOZOA FROM ZANZIBAR, 525 8

mandibles occur very frequently in the genus and may be called

Reteporidan avicularia. There are other small avicularia with

solid chitinous semicircular mandibles like those near the oral

aperture. These have two muscular tendons, whereas the large

triangular ones have a single tendon, and this muscular distine-

tion between the round and triangular mandibles is very frequent

in the Cheilostomata.

In the monilifera group the growth of the ovicell can be well

followed. Looking down the peristome the opening of a large

pore is seen in the distal wall (Pl. LX_XJI. fig. 1) until the ovicell

commences to grow (figs. 1 & 2), when the pore is more or less

hidden by a disk growing on a small stalk (fig. 2). This disk

becomes concave as it grows, showing this to be the commence-

ment of the inner wall of the ovicell (fig. 3), then in a later

stage part of the front is open (fig. 5), next the slit of the upper

part of the trifoliate stigma is seen (fig. 6), and soon the ovicell

is complete (fig. 7).

More or less similar commencements of the ovicell occur in

other genera, and sometimes specimens show these disks when

no ovicells are developed.

There is one large rosette-plate at the distal end of the

zocecium, and the muscles for withdrawing the polypide are

arranged on each side of this with a chitinous thickening to

which they are attached.

The avicularia are very variable in etepora, and we must not

trust to them too much in determining species.

Loc. Off Cape York, 8 fath. (Busk): Red Sea (Waters). Wasin,

Brit. East Africa, 10 fath. (500), 20 fath. (520), collected by

Crossland.

Rerepora PropuctaA Busk. (PI. LX XII. figs. 9, 10.)

tetepora producta Busk, Zool. Chall. Exp. vol. x. pt. xxx.

p. 108, pl. xxv. fig. 7 (1884); MacGillivray, ‘‘ Tert. Polyzoa of

Victoria,” Trans. Roy. Soc. of Victoria, vol. iv. p. 115 (1895) ;

Thornely, Records of Indian Museum, vol. i. pt. 3, no. 13, p..193

1907).

Retepora porcellana, var. laxa MacGillivray, Prod. Nat. Hist.

Victoria, dec. x. p. 15, pl. 95. fig. 6 (1885).

The small semicircular avicularia are very much raised, which

is not usualin Retepora, and are crenulate, as is frequently the

case in the round avicularia of the genus. These avicularia do

not occur on the younger zocecia but are common on the others.

The zocecia are much smaller than those of 2. porcellana MacG.

and the spines are ridged, but Iam not sure that it is right to

speak of them as articulated. In the zowcia near the border

there are often triangular avicularia with a single beak. The

ovicells in the Wasin specimens are not very much raised and

have a distinct median slit. On the dorsal surface there are

long narrow avicularia; the meshes are large, but “ papillary

eminences ” are not found as a frequent character.

326 MR. A. W. WATERS ON

Loc. Tongatabu (Polynesia), 18 fath. (Chall.); Samboangan,

10 fath. (Chall.); Port Phillip Heads, Victoria (J/acG.); San Pedro

Shoal, 25 fath.; Daros Island, Amirante Group (in my collection,

sent to me under another name). Wasin, Brit. East Africa, 10

fath. (507), 20 fath. (520), collected by Crossland.

Fossil. Victoria, Tertiaries (MacG.).

RETEPORA DENTICULATA Busk. (Pl. LX XII. fig. 8.)

Retepora denticulata Busk, Zool. Chall. Exp. vol. x. p. 109,

pl. xxvi. fig. 1 (1884); Philipps, “Rep. on Polyzoa,” Willey’s

Zool. Results, p. iv. p. 449, pl. xliii. fig. 13 (1899); Calvet, “ Bry.

@Amboine,” Rev. Suisse de Zool. vol. xiv. p. 620, pl. xxi.

figs. 2-9 (1906); Thornely, ‘“‘ Mar. Poly. of the Indian Ocean,”

Trans. Linn. Soc., Zool. vol. xv. p. 147 (1912).

The lamina of the ovicell is not quite so long as that figured

by Calvet, nor has it the orifice mentioned by him. Only a

few ovicells have been found. The trabecule are not barren.

There are large bifid avicularia on many of the zocecia near to

the distal end, which are probably only the small labial avicularia

modified ; also there are small semicircular or oval avicularia in

the older zocecia, near the middle of the front surface, but in the

younger zoccia they are wanting. Similar avicularia also occur

in the ‘ Challenger ’ specimens.

There are about 11 tentacles.

Internally, at the proximal end on each side of the zocecium,

there is a chitinous thickening (fig. 8) for the attachment of the

muscles which withdraw the polypide, and between these two

sets of retractors there is a thick funicular bundle passing to

the next older zoecium. The interior wall of the zocecium is

lined with very large square cells, and very similar cells line the

zoecial and vibracular chambers and the tube-pores of some

Cupularie, recent and fossil.

Reteporella dendroides Ortmann is similar in many respects.

Loc. Sandwich Is., 20-40 fath. (Calvet); Lifu, Loyalty Is.

(Ph.); Amboina (Calvet); Amirante, 15-35 faths., Seychelles, 34

fath. (7h.). Wasin, Brit. East Africa, 10 fath. (501), collected by

Crossland.

RETEPORA JERMANENSIS Waters.

Waters, “ Mar. Biol. of the Sudanese Red Sea,” Journ. Linn.

Soc., Zool. vol. xxxi. p. 176, pl. xviii. figs. 11-16.

Loc. Red Sea (Waters). Prison Island, Zanzibar Channel (513),

the shore, collected by Crossland.

RErEPoRA TUBULATA Busk, var.

Ketepora tubulata, type, Busk, Zool. Chall. Rep. vol. x. pt. xxx.

p. 121, pl. xxviii. fig. 2 (1884); Thornely, “‘ Ceylon Pearl-Oyster

Fisheries,” vol. iv. p. 125 (1905); Records of Indian Museum,

vol. 1. p. 193 (1907).

BRYOZOA FROM ZANZIBAR. 527

The fureate avicularium at the proximal end of the fenestra

is not so large or as much divided as in the ‘Challenger’ specimen,

but this divided avicularium is a character of much use in dis-

tinguishing the species from F. hirsuta B. and some others about

the same size with various zoarial characters similar. There are

no other large avicularia on the fenestral border, and there is a

delicate spine at each side of the oralaperture. As Miss Thornely

found in the specimen from the Indian Ocean, the growth some-

times brings the celluliferous surface to the outside as in

R. philippensis B.

Ortmann has described a species as FR. axillaris which has

an avicularium at the proximal end of the fenestra, but the

shape of the avicularium and mandibles differs from the species

from Zanzibar in which the avicularia are similar in shape to

those of 2. hirsuta.

There are 11 tentacles, and the oral glands are moderately

large but not much elongated.

Loc. Cape York, 8 fath. (Busk); Gulf of Manaar (7.); Gaspar

Straits, Malacca Straits (7h.), var. Wasin, Brit. East Africa, 10

fath. (501), Ras Osowamembe, Zanzibar Channel, 10 fath. (501),

10-20 fath. (504, 514), collected by Crossland.

? BIFAXARIA VAGANS Thornely. (PI. LXV. figs. 13, 14.)

Thornely, ‘‘ Mar. Polyzoa of the Indian Ocean,” Trans. Linn.

Soe., Zool. vol. xv. p. 145, pl. viii. fig. 7 (1912).

Zoarium erect, cylindrical, about 0-3-0-4 mm. in diameter,

branches dividing dichotomously at nearly right-angles, continuous

in the younger portions, but in the older the junctions are broken

across and there is a chitinous tube inside the articulation.

Zocecia face opposite all round the stem in four directions; the

surface of the zocecia has longitudinal ridges and large pits. The

secondary orifice is round with a small mucro bearing a small

avicularium, with an acute triangular mandible extending into

it, while the primary orifice, which is some distance down the

peristome, has the proximal edge straight. There are no ovicells

on the specimens examined, which were growing on Retepora,

nor were any ovaria or testes found in the sections made. There

does not appear to be any bar to the avicularium. There are

about 16 tentacles.

This is much like the ‘ Challenger’ Lifaxaria reticulata * Busk,

but in the British Museum specimens there are no avicularia,

and the surface is reticulate and apparently not perforate.

Bifaxaria of the ‘Challenger’ is not a satisfactory genus, and

will be broken up. It might be retained for B. submaucronata B.

and £. corrugata B., which are closely allied, and have the

avicularia at the proximal end of the zocecia, not at the distal

end as supposed by Busk. JB. denticulata B. has been removed

* Busk, Zool. Chall. Exp. vol. x. pt. xxx. p. 82 (1884).

528 MR. A. W. WATERS ON

to Sclerodomus by Levinsen; B. papillata B. should perhaps be

put to Urceolipora; B. abyssicola B. probably belongs to the

Cellarinella group; B. levis B. must be placed elsewhere; B.

minuta B. perhaps belongs to the B. reticulata B., B. vagans

Thornely, group.

Loc. Seychelles, 34-39 fath., Amirante, 34 fath. (Thornely).

Wasin, Brit. East Africa (500); Ras Osowamembe, Zanzibar

Channel (504), collected by Crossland.

ADEONID4.

In a short paper in the Annals and Mag. of Nat. Hist. for May

1912*, I have given grounds for believing that the Adeonide

will be found to be a much more important group than has

generally been supposed, and this in spite of none of the characters

upon which it was founded being of the importance then supposed.

A large number of the species are pigmented, most have long

pore-tubes, there are triangular avicularia on the front, and

often vicarious avicularia on the zoarial border, and both kinds

are without any cross-bar. There are no external ovicells,

the embryos being developed in an ovicellular sac which about

half, or, in some cases, nearly wholly fills up the zocecium ; and in

a considerable number the zocecium in which the embryo is

developed is larger than the others, with a differently shaped

aperture, together with a difference in the frontal pores, and these

larger zoccia have been called goneecia. The number of species,

however, in which there is no difference in the ovicelligerous

zocecia 1s very large, perhaps as many are indistinguishable as are

externally recognizable. There are no oral glands in any species

examined. I had previously shown that in the Adeonella of

Busk’s ‘ Challenger’ Report there were species in which the pore

entered into the zocecial chamber, while in others the pore is

above the operculum, also that the opercula in some are nearly

straight on the proximal border, others have a broad curve. At

the time there was not sufficient material for complete generalisa-

tion, but it is now clear that I was right in separating from

Adeonella species with a pore entering into the zocecial chamber,

and it is now seen that they also have a more or less straight edge

to the operculum. This group is now known as Adeonellopsis of

MacGillivray ; and Adeonella, much the same as I restricted it, is

accepted.

The wall of the sac (Pl. LX XIII. figs. 3, 5, s.) containing the

embryo (figs. 3, 5, emb.) is much thicker than that of the sac in

such species as Lepralia cucculata B., which also has no external

ovicells. The very earliest stages of this sac have not been seen,

and while it does not ever seem to hang from the opercular

region as a small pendant, yet a comparison with the sacs of

* Ser. 8, vol. ix. p. 493 (1912).

BRYOZOA FROM ZANZIBAR. 529

Diplodidymia complicata Rss. (text-fig. 79, p. 491) may throw

some light upon the growth.

Blind cells, that is cells without an oral aperture, are very

frequent throughout the family, and sometimes the central or

stalk portion is composed of such cells. A few have a closure

somewhat like those of the Cyclostomata, but not entirely so, as

a membrane from below the operculum spreads to the front to

make the funnel-like closure. Membranous closures are also

occasionally found, and these may be perforated. These closures

with tubuli occur in Adeonella contorta Mich., and I have found

them in Lepralia syringopora Rss., one of the Adeonide.

The gonecia in Adeonide, when distinguishable, frequently

are situated near the border, but in one species they are all

situated on the median line, while in another they are in groups

in various places, distinguishable by the naked eye.

In my recent paper dealing with Adeonide I alluded (p. 496)

to the occlusor muscles being, as a rule, single in triangular

mandibles, whereas the semicircular mandibles usually have two

tendons. A slightly different statement would have been better,

as I was aware of some exceptions, and since writing I have come

upon more—in fact in Cellepora and Holoporella many of the

triangular mandibles have two tendons, but in Flustra, Membrani-

pora, Retepora, and Cribrilina, etc., and in most genera the rule

seems to be, only one tendon to the triangular mandibles. Some

of the exceptions may be in semicircular avicularia modified

into triangular ones. I have not got a record on this point of all

the mandibles prepared, and must make further examination.

ADEONELLA PLATALEA Busk. (Pl. LUXXIII. figs. 3-5, 8 &

text-fig. 82.)

Eschara platalea Busk, Brit. Mus. Cat. Mar. Pol. p. 90, pl. ev.

figs. 1, 2 3, pl. eviii. fig. 4 (fide Busk) (1854); MacGillivray,

Prod. Zool. Vict. dec. v. p. 41, pl. xlviil. fig. 4 (1880).

Eschara hexagonalis Haswell, Proc. Linn. Soc. N.S. Wales, vol. v.

p. 41, pl. ili. figs. 1, 2 (1881).

Adeonella platalea Busk, Zool. Chall. Rep. vol. x. pt. xxx.

p. 184, pl. xxi. figs. 4, 4a & fig. 50 (1884); Meissner, M., “ Liste

der von Herrn Prof. Semon bei Amboina und Thursday Island

gesammelten Bryozoen,” Jenaische Denk. vol. viii. p. 731 (1902) ;

Waters, “ A Structure in Adeonella (Laminopora) contorta Mich.,”

Ann. Mag. Nat. Hist. ser. 8, vol. ix. p. 489 (1912).

Escharinella cecilleana VOrb. Pal. Frang. vol. v. p. 207 (1850) ;

and see Waters, Ann. Mag. Nat. Hist. ser. 7, vol. xv. p. 5

(1905).

I have not had the opportunity of re-examining the Z. cecilleana

d’Orb. since this collection was taken in hand, but the name

cecilleana cannot be retained, as the description was insufficient.

The ovicellular sac (fig. 3) is formed near the distal end

and ultimately fills up nearly the whole of the zoccium, but

530. MR. A. W. WATERS ON

the very earliest stage has not been seen. The wall of the

sac is formed of short cells and at one part touches the embryo.

The embryo in early stages (fig. 5) is surrounded by large and

long cells, and in some eases is filled up by formative tissue. In

later sta ges, when the embryo is nearly ready for a free existence

the large cells of the calotte are readily followed (fig. 6).

Text-fig. 82.

Fi iota

ise

Ve ee 2s

Lateral surface of Adeonella platalea. X 25

When the ovicellular sac has grown to a moderate size there

may be, near to the opercular wall, an ordinary polypide, but

often there is only a very small one looking like a simple bud,

being sometimes not more than a straight tube. There are strong

muscles at the distal end on one side for contracting the walls of

the ovicellular sac, to which they are attached, as well as to the

distal wall. The ovieells of A, polystomella Rss., are smaller than

those of the other species, but there is the same thick-walled

ovicellular sac. Also, in a specimen. of Beania magellanica B.,

from Chatham Islands, there is a large embryo in a thick-walled

cellular sac nearly filling the zoecium. This I do not find in any

of my specimens from | Naples, but there is a pair of sac-like

bodies near the distal end, and in the Antarctic specimens these

bodies are large and were thought to be for the formation of

testes, although there are ordinary testes near the distal end in

which the spermatozoa can easily be seen.

It has been supposed that all the large zocecia of the Adeonida,

namely the goneecia, are merely for embryos, but this is by no

means certain, as some may be for the testes, with which some are

nearly filled, but the material available does not permit of my

saying much on this point. However, it is only in a limited

number of species that there are large zocecia, and the same changes

may take place in these as in ordinary zocecia ; for as the gonecia

BRYOZOA FROM ZANZIBAR. 53h

contain polypides and buds, the embryos are only a stage in the

life of the goneecia.

In the present species the two outer rows of zocecia are, in parts

of the colony, larger than the rest, forming goneecia with a larger

and wider oral aperture, also the pore below it is wider, and a

short tongue from the distal end sometimes divides the pore into

two. In: many species of the Adeonidee where there are goncecia

they occur at the border of the zoarium or near a Rigarestions but

apparently in no species are they continuous all along the border.

When the zoccia are blind or closed there is often a dise with a

central tubule in about the position of the oral aperture.

In A. platalea there are at the border of the zoarium, besides

the large avicularia, very minute ones between the two zocecia,

and sometimes the small chamber has a round opening and no

mandible.

Loc. Bass’s Straits; Cape Capricorn; Victoria, Queensland

(MacG.) ; Port Darwin (Brit. Mus. Coll.); China Seas, 82-102

fath. (Chall.); Thursday Island (Meissner); Ye, Burmah Coast

(Thornely); Basilan, Indian Ocean (dOrb.). Wasin, Brit. E.

Africa, 10 fath. (501,520); Ras Osowamembe, Zanzibar Channel,

10 fath. (504); Prison Island, Zanzibar Channel, 8 fath. (505):

collected by Crossland.

ADEONELLOPSIS CROSSLANDI, sp. n. (Pl. LX XIII. figs. 1, 2, 6, 7,

12-14.)

This is, in many respects, like 4. swbsuleata Smitt, but there

seems sufficient reason for separation, for the ridge surrounding

the area in swbsulcata is here wanting and the suboral avicularium

is directed laterally.

The zoarium consists of flat branches (a little over 1 mm. wide

and less than | mm. thick) frequently bifurcating and sometimes

anastomosing.

All the zocecia are about the same size, no difference being

discoverable externally in the ovicelligerous zocecia, although in

sections zocecia steele filled by the embryo are found occurring

indifferently in any position except in the outer zocecia. The

proximal border of the aperture is crenulate and this, as mentioned

by Miss Thornely, is the case in subsulcata ; also the broad median

pore is denticulate ; however, in some zocecia, especially those next

to the outer row, there is no apparent pore.

There is a small triangular avicularium immediately below or

within the aperture, and often a few of the zoccia near a bifurea-

tion, and some of the bordering zocecia have also an avicularium

at the proximal edge.

The zocecia at the sides of the zoarium are opposite as, in fact,

are all the zocecia, so that the oral apertures on both sides rise to

the same height. On the side of the zoarium there are no

vicarious avicularia, whereas they occur in A. distoma, A. sub-

sulcata, and A. imperforata.

Proc. Zoou. Soc.—1913, No. XXXVI. 36

532 MR. A. W. WATERS ON

The tentacular sheath is attached to the operculum close to its

border, and on the inner side of the opercular region there is a

small globular body (fig. 2), immediately below which the new

bud grows. We see the same thing in various species, among

others, in ? Bifaxaria vagans Thorn., A. polystomella Rss., ete.,

and it is very large in Adeona foliacea var. fascialis Kirchenpauer,

being about 0-4 mm. in diameter, which is about double the

diameter of the globe in the present species. The wall is thick

and it may give support to the new bud.

There are about 13-14 tentacles. 4. distoma has 14-16.

Tt would seem that the ovicellular zocecia very frequently occur

near to a bifurcation, and there may be a group of three or four

ovicellular zocecia close together. The few ovaria seen consist of

two very small ovarian cells, at first situated at the distal end of

the zocecium, but others, no doubt older, occur by the proximal

end of the ovicellular sac. The ovaria are similar to those of

Bugula, and are of about the same size in Adeonella platalea,

A. polystomella*, and A. crosslandi. The early stages of develop-

ment have not been made out.

The ordinary zoccia may have the polypide entirely surrounded

by a testis, in fact, may be practically filled with it.

Loc. Wasin, Brit. East Africa, 10 fath. (500, 501), collected by

Crossland.

EXPLANATION OF THE PLATES.

Prats LXIV.

Fig. 1. Aetea anguina L. Showing the ovicell (ov.) at the end of the tubular pro-

jection; the retractor muscles (7m.) attached just

below the tentacles, and also attached at the other

end near the commencement of the basal expansion,

where the ovaria (oa.) also occur. X85. Fron

Wasin (500).

2. Do. do. Front view, showing the operculum and the tentacular

sheath (¢s.) to which the muscles are attached. 250.

3. Aetea truncata Landsborough. X25. From Prison Island (505).

4. Brettia tropica, sp.. X85. Dorsal surface. From Wasin (501).

5. Do. do. < 25. Anterior surface.

6. Catenaria diaphana Busk. X25. From Ras Osowamembe (504).

al Do. do. Lateral view, X 25.

8 Do. do. Operculum, X 85.

9 Do. do. X 25. Section showing embryos (/.) in the ovicell;

also an ovum (ov.) in the zocecium, and ovaria below

the ovicell (oa.).

10. Do. do. Ovarium, X 250.

11. Do. do. < 85. Section of zocecium showing ovicell (ov.),

diaphragm (d.), compensation-sac (es.).

12. Synnotum pembaensis, sp. n. X25. From Chaki-Chaki Bay, Pemba

Island (517).

118} Do. do. Mandible, X 250.

14, Do. do. Avicularium, X 85.

15. Do. do. x 3.

16. Synnotum contorta, sp.n. X 25. From Chaki-Chaki (617).

Wife Do. do. Avicularium, < 150.

18. Do. do. Mandible, X 250.

* A. polystomella occurs in the Red Sea, as I find from the examination of a

specimen in the British Museum, since my Red Sea paper was written. There is

also a specimen from Brishane (65.5.29.4).

BRYOZOA FROM ZANZIBAR. 533

PLATE LXV.

Wig. 1. Vittaticella elegans Busk. XX 25. Showing large avicularium. From

Prison Island, Zanzibar Channel (503).

2. Do. do. Mandible of large avicularium, X 85.

3), Do. do. Small avicularium, X 150.

4. Do. do. Distal end, showing the avicularium, X 86.

5: Do. do. Showing internodes composed of many zocecia with

ovicells, X 15. (503).

6. Do. do. Section of embryo im ovicell, X 400.

7 Do. do. Section showing zowcia with polypides and ovicells

(ov.) in position, X 85.

8. Vittaticella elegans, var. zanzibariensis, nov. Distal end, showing the

avicularium, X 85. From Wasin, Brit. KE. Africa

(500). a, mandible, X 250.

9. Do. do. do. x 50. Showing small avicularium on the

dorsal surface near the distal end.

10. Do. do. do. Upper half of the contents of the vitte, « 250.

From Prison Island, Zanzibar Channel (505).

ik Do. do. do. < 85. Showing internal zocecial muscles.

(505.)

12. Do. do. Semidiagrammatic transverse section through the

vitte (v.), X about 200.

13. 2 Bifaxaria vagans Thornely. X 25. From Wasin (500.)

14, Do. do. Operculum, X 85.

15. Chlidonia cordieri Aud. Section showing muscles (m.) from the parietal

wall to the base of the chamber, X 250. From

Wasin, Brit. E. Africa (500).

16. Do. do. Operculum, X 250. a, mandible, X 85.

Prats LXVI.

Hig. 1. Stirparia exilis MacG. 25. Only the lower part of the tuft is shown.

From Wasin, Brit. E. Africa (522), 20 fath.

2. Do. do. x 2.

3. Do. do. X 85. Stem showing disc from which a fresh branch

grows.

4, Stirparia dendrograpta, sp.n. X 25. Only the lower part of the tuft

is shown, and for clearness ouly a few of the spines

are drawn full length, but are shown as broken off,

which is frequently the case. A pair of radicles are

shown growing from the base of a node. From

Chuaka, Zanzibar (508), 2 fath.

5. Do. do. < 250. Avicularium.

6. Do. do. x 25. First zowcium of a tuft, showing long spines.

ie Do. do. < 85. Dorsal surface, showing the polypide in the

zocecium with ovaria (ovar.).

8. Do. do. < 250. Ovarium from below the polypide, with one

ovarian cell.

9. ote dos xX 2. On the left a few zocecia are shown in a natural

position; the others are spread out as in a mounted

slide. This piece is entirely unattached.

Prats LXVII.

Fig. 1. Cellaria wasinensis, sp. n. X25. Specimen prepared with Eau de

Javelle, showing the openings of the ovicell in an

enlarged part of the zoarium. From Wasin.

2 Do. do. Mandible, & 85.

3 Do. do. Showing the frontal membrane, with the trabecule and

the tessellated area below the operculum, X 86.

4. Do. do. Tessellated area, X 250.

5 Do. do. Avicularium prepared with Eau de Javelle, showing

the plate im the submandibular part. XX 8d.

6 Do. do. x 3.

7. Cellaria gracilis, var. tessellata, nov. ‘Tessellated area, X 250. From

Ras Osowamembe.

Figs. 8,9. Farcimia oculata Busk. Stalk growing from stolon, X 265.

36%

Fig.

MR. A. W. WATERS ON

Membranipora armata Haswell. Opercular region, seen from the interior.

The opercular muscle is attached to a linear sclerite (sc.) on the mem-

branous operculum, and from the sclerite there is also a muscular band

to the tentacular sheath (¢.s.). Two protoplasmic bands (0.) pass to the

distal rosette-plate. XX 50.

11. Diplodidymia complicata Rss. X 6. Colony from Chuaka (506).

12. Do. do. X 25. Lower part of colony.

13. Do. do. x 85.

14. Do. do. < 250. Operculum.

15. Do. do. X 250. Mandible.

Pratt LXVIII.

1. Stirparia zanzibariensis, sp. n. X 85. From Chuaka, Zanzibar, 8 fath.

(508).

Do. do. <2:

Serupocellaria pilosa Aud. X85. Dorsal surface. From Wasin, Brit.

KE. Africa, 10 fath. (500).

Do. do. X25. Anterior surface.

. Scrupocellaria macandrei Busk. X85. Dorsal surface, showing the

groove extending beyond the vibracular chamber.

From Prison Island, Zanzibar Channel (505).

Do. do. < 85. Anterior surface.

. Bicellaria chuakensis, sp.n. Avicularium, « 250.

Do. do. < 25. From Chuaka Bay, Zanzibar Channel, (515).

. Scrupocellaria wasinensis, sp.n. X25. Dorsal surface. From Wasin,.

Brit. E. Africa, 10 fath. (522).

Do. do. X 26. Anterior surface.

. Scrupocellaria ferov Busk. X25. Anterior surface. From Prison

Island, Zanzibar Channel, 8 fath. (805).

12. Do. do. Mandibles—(a) anterior, (4) lateral, X 85.

13. Do. do. Base of vibracular seta, * 250.

14. Do. do. Decalcified piece, showing the articulation with the

chitinous tube already formed to the inside zoecia

of the new branch, X 25.

16 Do. do. Dorsal surface, & 25.

PratE LXIX.

1. Canda retiformis Pourt. X25. From Ras Osowamembe, (504).

2. Do. Do. Dorsal surface, X 28.

3. Serupocellaria cervicornis Busk. X 25. From Wasin (522).

4, Do. do. Dorsal surface, showing the articulation, < 25.

5. Caberea ellisii Flem. Vibracular chamber seen from the inside: a, },.

vibracular chamber; 4, ¢, continuation of the groove beyond the

chamber.

6. Canda retiformis Pourt. Dorsal surface, to show the articulation. X about 6.

7. Scrupocellaria ferox B. Do. do. _X about 6.

8. Scrupocellaria jolloisii Aud. Do. do. X about 6.

9. Flabellaris (Menipea) cuspidata B. Do. do. X about 6.

10. Bugulopsis peachii Busk. Do. do. X about 12.

11. Menipea patagonica B. Do. do. X about 12.

12. Menipea buskii W. Th. Do. do. X about 25.

13. ? Menipea cyathus W. Th. Do do X about 25.

. Stirparia zanzibariensis, sp.n. Showing basal stalk from which there are

delicate rootlets, and also capsules nearly filled with a bright yellow

substance. XX 12.

. Bugula robusta MacG. Showing the earlier zocecia, X 25; a, natural

size. From Wasin (501).

16. do. do. Avicularium, X 8d.

17. Serupocellaria wasinensis, sp. n. Zocecia showing the position of the

ovaria (ov.), testes (¢.), rosette-plates (7.). > 85.

18. Do. do. Ovarian cells with protoplasmic threads from the

rosette-plates, K 250.

19. Do. do. Ovaria showing two nucleated ovarian cells, X 250.

20. Serupocellaria ferox Busk. Band which starts near the distal end and

passes down the side of the zocecium; also small bundle of protoplasmic:

threads running parallel with the granular band. 250.

BRYOZOA FROM ZANZIBAR. 535

PLATE LXX.

Fig. 1. Schizoporella nivea Busk. X25. From Wasin (501). This may perhaps

be called var. wasinensis.

2. Do. do. x 85. Operculum.

3. Do. do. X 85. Oral aperture closed by calcareous wall.

4. Gemellipora protusa Thornely. X 25. From Wasin (520).

5. Do. do. X< 85. Operculum.

6. Do. do. < 85. Interior of the zoccium showing the oral

aperture.

7. Schizoporella nivea Busk. X 25. From Wasin (501).

8. Do. do. X 150. Basal avicularium.

9: Do. do. X< 150. Lateral avicularium.

10.8 Petralia chuakensis Busk. > 25. From Chuaka, Zanzibar.

Make Do. do. x 85. Operculum.

12. Do. do. x 85. Mandible.

13. Do. do. < 250. Mandible showing the crenulated border.

14, Do. do. X< 25. Dorsal surface showing spaces for the attach-

ment of radicle tubes.

15. Petralia vultur, var. armata, nov. X 85. Section through the zocecium,

showing the tentacles, with two larger than the

others.

16. Do. do. do. < 85. Operculum.

AWE Do. do. do. < 550. Section of tentacles near the base

showing the two large fan-shaped tentacles with

long nuclei near the edge.

18. Do. do. do. x 25. Eau de Javelle preparation. From

Chuaka.

19. Do. do. do. > 85. Mandible of oral avicularium.

20. Do. do. do. > 85. Mandible of round avicularium.

21. Lepralia feegensis Busk. X 85. Operculum.

22. Do. do. X85. Mandible.

Prate LXXI.

Fig. 1. Membranipora savartii Aud. Zoccia showing the network of protoplasmic

threads. In the upper zocecium histolysis of the

polypide has commenced, and in the right hand

zocecium (7) histolysis has progressed further, and

now considerable granular masses are formed. In

the lower zocecium, of which only the distai end is

shown, the polypide is in full vigour with digestion

active and the tentacles unaltered. Drawn trom a

decalcified mount, < 85. From Zanzibar Channel

(528).

2 Do. do. Section showing plasma threads by the side of finely

granular cord and parts of narrower cords, X 1000.

3 Do. do. The granular masses shown in the right zocectum in

fig. 1, X 150.

4. Do. do. A similar mass shown in transverse sections sur-

rounded by protoplasmic threads, x 1000.

5. ? Membranipora armata Haswell. Mandible, X 85.

6 Do. do. Decalcified avicularium showing retractor muscle (a),

divaricator (6), with tendon attached to the mandible

in the middle of the base, and the “peculiar body”

(ec), X 85.

Uke Do. do. Avicularium, X 85.

8. Do do. Lower portion of lateral gland, X 320.

9. Do. do. Decalcified zocecium showing the avicularian chamber

on the left with muscles as in fig. 6, and the glan-

dular chamber (ge.) on the right, X 86.

10. Do. do. X 25. From Wasin, Brit. East Africa (500).

536

ON BRYOZOA FROM ZANZIBAR.

Puate LXXII.

Fig. 1. Retepora monilifera, var. umbonata MacG. X 25. Showing in the lower

10.

Do.

do.

zoccia the large pore within the oral aperture, and

in the upper zoccia the stalked dise which grows in

front of this pore. From Victoria.

x 85. Aperture showing the stalked disc.

x 50. The disc is shown larger and can now be

recognised as the commencement of the wall of the

ovicell.

X 50. The same rather larger.

x 50. The sides of the ovicell are commencing to.

grow, but the ovicell is entirely open in front:

x 50. The ovicell showing a large cruciform opening..

x 50. The ovicell shows the complete trifoliate-

stigma.

. Retepora denticulata Busk. X 250. Section of the proximal end of the-

zocecium showing the muscles (m.) attached to the base of the polypide

and to the wall of the zocecium, which has a chitinous thickening where

the muscle is attached. Between the two sets of muscles a thick funi-

cular cord (f.) passes through a large rosette-plate.

Do.

. Retepora producta Busk. X 25. From Wasin (520).

do.

xX 50. Ovicell.

. Rhynchozoon profundum, var. laminatum nov. Diagrammatic figure of

the aperture and ovicell.

. Steganoporella magnilabris Busk. X about 50. Showing the tentacular

8}, Do.

14. Do.

15. Do.

16. Do.

Y/, Do.

18. Do.

19. Do.

20. Do.

do.

parts attached to the side wall of the proximal

corner, while the stomach, etc., is in the other corner

connected by a long narrow cesophageal tube.

x 50. Operculum of B zoccium.

x 50. Decalcified distal end of the zocecium as seen.

from the front with the polypide seen through the.

wall and (se.) sclerites to which the frontal muscles:

are attached.

X 250. Section of the edge of the operculum.

X 85. Section of the distal wall through two rosette-

plates. Probably the fleshy parts belong in part to:

the distal and in part to the proximal wall.

X 50. Section cut through an operculum showing a

membrane (m.) between the main sclerites.

x 50. Operculum of A zoccium.

< 85. Decalcified distal end seen from inside with

the basal wall removed. The tentacular sheath (é.s.);

is held by four bands (6.) attached to the distal wall,

also there are muscles (¢.s.m.) from the distal end of

the tentacular sheath to the wall (a.) which divides

the zowcium. The large retractor muscles (7.) are

seen attached to a tendon, while below there is a

smaller pair of muscles (d.) not attached direct to,

the operculum, but to the frontal wall quite close

to the operculum.

X 25. Growing end with the membrane covering the

surface. The end zocecia are as yet quite empty

and the operculum of a B zocecium (B.) is growing,

but no basal sclerite is yet formed. In the second

row the frontal calcareous wall only extends a slight

way into the zoccium and here the zoccium is still

quite empty. Next the short tube (¢.) connecting

the two parts of the zocecium grows and the lip

commences, but is still some distance from the

frontal membrane, though later it rises right up to:

the proximal edge of the operculum. The frontal

sclerite (sc.) to which the opesial muscle is attached

does not occur in the younger zocecia, but is seen im

the mature zocecia.

Fig. 1

)

<6 OO NI

ON TWO AFRICAN ANTELOPES. DOT

Prare LXXIII.

. Adeonellopsis crosslandi, sp. n. X 250. Showing an ovarium in the

distal end of the zocecium. From Wasin.

X 250. Showing round body under the operculum

with the protoplasmic threads to the buds below.

. Adeonella platalea Busk. X 150. Showing the early stage of the ovicel-

lular sac (s.) near the distal end containing the

embryo (emb.). From Wasin (501).

X 85. Section showing the ovicellular zowecium nearly

filled up by the embryo, with an ovarium (ovar.)

now proximal to the embryo. From Wasin (501).

< 150. Section of embryo showing the zocecial wall

(z.w.), the ovicellular sac (s.), and the embryo

(emb.). The muscles are seen at the distal end on

the right in the zocwecial chamber. From Wasin (501).

. Adeonellopsis crosslandi, sp. n. X 250. Thick section of embryo

showing the aboral end. Fig.6a, 250. Diagram-

matic outline of vertical section.

< 85. Opereulum.

. Adeonella platalea Busk. X85. Operculum.

. Osthimosia zanzibariensis, sp.n. X 85. Operculum.

10. Lepralia turrita Sm. X 85. Operculum.

14,

. Holoporella albirostris Sm. X 85. Polypide showing the long oral

glands.

. Adeonellopsis crosslandi, sp.n. X 25.

13.

x 50. Transverse decalcified section showing the

opposite zocecia at about the same stage of develop-

ment.

Natural size.

15. Osthimosia zanzibariensis, sp.n. X 50.

16. Schizoporella nivea Busk. X50. The two zoccia figured were near

together but not in the same row, and are placed

together to save space. A sac-like structure (sc.) is

shown at the base of each ovicell (oc.), but of the

numerous muscles in this sac only about half of

those in focus are drawn. It seems that the ovum

(ov.) passes into this sac which is then ruptured (7.),

and the ovum is pressed forward into the ovicell

where it segments. The small oral glands are

shown (g/.).

35. Notes on Albinism in the Common Reedbuck (Cerw-

capra arundinum), and on the Habits and Geographical

Distribution of Sharpe’s Steenbuck (Raphiceros sharpez).

By Major J. StevENson-Haminton, C.M.Z.S., Game-

Warden of the Transvaal.

[Received April 8, 1913: Read April 22, 1913.]

INDEX. Page

(CAP UORLY DTH GHROKDCIODTID, "53 oon soutien ves 2o0ane oob oan esdnsoson SEH

IMO YOOGUROS SUCH VED ogi aroaae neddossanoadaxscousuee soxdu0%90 bon AY)

Albinism in the Common Reedbuck (Cervicapra arundinum).

During the early weeks of 1911 a rumour reached me of three

white Reedbucks having been seen near the south-western border

of the Sabi Game Reserve. I gave orders that they should be

538 MAJOR J. STEVENSON-HAMILTON ON

kept under observation so far as possible, though the very broken

and hilly nature of that area rendered any close following of

their movements impracticable.

A month or two later a troop of hunting dogs (Lycaon pictus)

passed through that part of the country and, it is believed, killed

one of the animals, as only two were afterwards seen. I then

made preparations for an expedition to capture the survivors, if

possible, for the Pretoria Zoological Gardens, but almost imme-

diately afterwards heard that they had wandered out of the

Game Reserve and that the male had been shot by some unknown

person. Towards the end of 1911, the last survivor, a female,

came back to the Reserve. The Museum authorities having

expressed a desire to obtain the skin, failing success in capturing

the animal alive, I proceeded to the ground. Under the circum-

stances it seemed that the capture would be very difficult, and

that failure would almost certainly drive the animal out of the

Reserve, where it would be killed at once by natives or white

men, and so be irretrievably lost. I therefore shot it, and

sent the skin, bones, and measurements to the Pretoria Museum,

where it is now set up.

The animal proved to be a female about two years old, and of

a perfectly pure white colour throughout. The muzzle and inside

of the ears, membrane round the eyes, ete., were pinkish or flesh-

colour.

The skin on the back of the ears was scabby and unhealthy-

looking. Eyelashes white. Pupils reddish black. Inner irides

grey blue. Outer irides pearly with darker rays. White of eyes

normal. Hoofs and lateral hoofs pale horn colour, the former

nearly white at bases.

Measurements for a two year old animal about normal.

Head and body 52 inches; tail 73 ; fore girth 32.

When found she was running with an ordinary ram, and

contained a perfectly formed male feetus, which was normal in

all respects. I should be inclined to think, in view of the time

which had elapsed since the disappearance of the male albino,

that she had been impregnated by the normal ram with which she

was found ; but this isa point I cannot speak on with any certainty.

It is worth recording that within a mile of the spot where I

saw this animal, and in the midst of the comparatively small

area where these Reedbucks had been born and bred, was a

village in which lived an albino male native, his hair nearly

white, and his eyes of almost the same curious light grey-blue

shade as those of the Reedbuck. Native information states that

albino children have been born before in the same village, and

that white Reedbucks have also been seen in former years in the

district. ;

Some two months later (January 1912) a report came to hand

of another albino Reedbuck at a point 60 or 70 miles north of

where the female was secured. ‘This animal was always noticed

alone, and was repeatedly seen by passengers by the Selati train.

TWO AFRICAN AN'LELOPES. 539

T sent a ranger up to endeavour to capture it alive, but attempts

were unsuccessful, owing to the mesh of our net being too large.

It was seen close enough, however, to be recognized as a female,

nearly pure white and about one year old. After the unsuc-

cessful attempts to capture her alive, she was noticed at intervals

for another month ; but before another attempt could be made,

she disappeared, probably killed by wild dogs.

These animals, which may have been born in 1909 in the first

ease and 1910 in thesecond, were found at a considerable distance

north and south of one another, and the incidents almost certainly

had no connection whatever, but both occurred within a few miles

of or among the foothills of the Drakensberg Mountains, on the

western border of the Reserve. In nearly eleven years’ caretul

observation of Reedbucks in the Reserve, these are the only

eases of albinism that have been brought to my notice.

Sharpe's Steenbuck (Raphiceros sharpei).

Spiti-pite or Pitsi-pitsi of the Thonga tribes of the North-

east Transvaal and neighbouring Portuguese East Africa.

Isigulane of the Swazis.

(reographical Distribution. —The most northerly habitat of the

species appears to be British Nyasaland, where it was discovered

and recorded by Sir A. Sharpe. Thence it extends down through

Mashonaland, possibly hugging the vicinity of the eastern hilly

country, to the North-eastern Transvaal. It is there found all

along the course of the Lebombo Hills (but never in the Drakens-

berg Mountains, sixty miles further west) as far as the Crocodile

River at Komati Poort. It becomes very numerous between the

Limpopo and Letaba Rivers, and is there spread through broken

ground to as much as thirty miles from the Lebombo. South of

the Olifants it becomes progressively scarcer, and is very rare

indeed between the Sabi and Crocodile Rivers. It reappears,

however, in Swaziland (Mr. R.'T. Coryndon), and is found all

along the border of that country and Portuguese East Africa on

both sides of the Lebombo. Mr. Coryndon is of the opinion that

it occurs also in Northern Zululand—that is to say, as far as the

end of the Lebombo Hills.

Whatever be the case in Nyasaland and Mashonaland, in the

Transvaal and Swaziland it occurs only in one narrow strip com-

prising the Lebombo Hills and their immediate surroundings.

No antelope answering to the description seems to be reported

from Natal or Northern ‘Pondoland, and in the south of the latter

we arrive at the extreme known northern limit of the Grysbuck

(2. melanotis), at Port St. John’s.

The geographical distribution of Sharpe’s Steenbuck, therefore,

seems to be between 14° and 28° south latitude, and between

about 312° and 332° east longitude, the line of distribution

following more or less the south- -westerly trend of the coast,

though at some distance from it.

540 MAJOR J. SLEVENSON-HAMILION ON

There seems to be a gap of 4° of latitude in which no small

grizzled antelopes are found between the most southerly habitat

of Sharpe’s Steenbuck and the most northerly of the Grysbuck

(32° south).

Description.—The animal found in the North-eastern Trans-

vaal bas been described from specimens obtained by Mr. Grant

(collector for Mr. Rudd). Females and immature males are,

generally speaking, of a brighter red colour than old males.

The distribution and profuseness of the white in the coats tends

to vary in different specimens, and some show more than others.

Individuals display small patches of white hairs here and there

on back or sides.

In a half-grown female the dark horseshoe mark on the crown

was found to continue backwards in the form of a narrowing dark

brown line as far as the centre of the back, where it suddenly

broadened and then disappeared. In the same animal the face-

markings were much more highly contrasted than in older

specimens.

There are four mamme present in the females.

I fancied that I detected a very slight thickening of the tissues

at the points where the lateral hoofs are found in other species.

The horns of the males are very short and rather conical in

shape. They seldom exceed an inch and a half in length, and

I should imagine two inches to be about the maximum length.

Mr. Selous has recorded shooting certain small grizzled animals

in the Mashonaland Hills which possessed horns over three inches

long, and this seems to point, perhaps, to an interesting variation

of the type in that country.

Habits, etc—Sharpe’s Steenbuck is found either in patches of

thick bush, which may be at a considerable distance from any

hilly country, or among the stones and boulders of the lesser

ridges of the Lebombo Hills and the thick covert sometimes

growing at their bases (North-eastern Transvaal). In no case has

it been noticed at any very great distance from water. In the hill-

country it especially favours the rough boulders interspersed with

rank grass and bushes, crowning the crests of long stony ridges

which gradually merge into grassy slopes on either side. Its

colour almost exactly matches the red tint of the Lebombo rocks,

and consequently, when it chooses to lie close, it is a most difficult

creature to see. One or more individuals have a custom of

returning to the same spot to deposit their droppings, but the

habit is not a constant one. It feeds at night or in the very late

evenings and early mornings, and, except on dull days, always

lies up for the daylight hours among boulders or long grass.

In cloudy weather, especially in spring, when the grass is young

and fresh, it may be observed quite late in the day feeding close

to one of its refuges, moving slowly about, and lying down at

frequent intervals. In the stomachs examined, I have found

about half the contents to consist of young grass and the rest of

the small leaves of ground-shrubs.

TWO AFRICAN ANTELOPES. 541

When disturbed while lying up for the day, Sharpe’s Steenbuck

usually springs up 30 or 40 yards away, and makes off at once.

Occasionally, however, when well concealed, it lies very close,

and only gets up whenapproached within a few yards. In no case

that I have observed did it ever pause before going, resembling

many other small buck and hares in this respect. It always goes

at best pace for a considerable distance, squatting again suddenly

when it has found suitable covert. Its gait isa scuttling run, and

it never bounds like a Steenbuck or a Duiker. Nevertheless, its

speed is considerable, and, in the rough country affected, it takes

a very good dog to run down an adult of either sex.

It is very solitary in habit, and even when a pair are put out.

of the same patch of bush, they seem generally to have been lying

in different parts of it. Bush and rocks seem to be regarded

equally as natural refuges, and in following up individuals I

could not discover any preference for one over the other. Even

in the heat of the day the shade of a large stone on a hillside,

destitute of a blade of grass or a scrap of bush, is sometimes

sufficient for the animal’s requirements.

Most of the females appear to be in lamb(North-eastern Trans-

vaal) in October and November, and I imagine the bulk of the

young to be born in the early or mid summer months, though I

should hesitate to affirm that they do not, like many other small

buck, breed more or less all the year round.

From observation of appearance and habits in the field it would

be impossible to recognize its kinship with the true Steenbuck.

In habit it approximates much more closely to the Grysbuck, for

which it used often to be mistaken by hunters. The native

(Thonga) name is Spiti-pite or Pitsi-pitsi for Sharpe’s Steenbuck,

while they call the Steenbuck Jnginana and the Livingstone

Antelope Jnhlengana.

Mr. R. T. Coryndon, the Resident Commissioner of Swaziland,

confirms these observations of the animal’s habits in the North-

eastern Transvaal as being applicable to what he has himself

observed in Swaziland, and gives the Swazi name for it as

Isigulane, for the Steenbuck /ngcina, and for the Livingstone

Antelope Znhlengana. The Zulus call the common Steenbuck

Iqina. The natives therefore recognize, judging as they do

merely from habits and outward appearance, no affinity between

the two species of Steenbuck.

Sharpe’s Steenbuck is generally spoken of by Colonists in South

Africa as “Grysbuck” or “ Grys Steenbuck,” and in this Province

it received its proper title for the first time in the Game Laws.

of 1912.

542 MR. H. J. ELWES ON THE

EXHIBITIONS AND NOTICES.

April 8, 1913.

Sir Jouy Ross Braprorp, K.C.M.G., M.D., D.Sc., F.B.S.,

Vice-President, in the Chair.

THe Secrerary read the following report on the additions that

had been made to the Society’s Menagerie during the month of

Wlevela JUSS} 3

The number of registered additions to the Society’s Menagerie

during the month of March last was 133. Of these 45 were

acquired by presentation, 58 by purchase, 11 were received on

deposit, 6 in exchange, and 13 were born in the Gardens.

The number of departures during the same period, by deaths

and removals, was 167.

Amongst the additions special attention may be directed to :—

1 Kandt’s Cereopitheque (Cercepithecus kandti), from Lake

Kivu, new to the Collection, deposited on March 7th.

2 Canadian Poreupines (Hrithizon dersatus), from North

America. presented by W. O. Danckwerts, Esq., K.C., F.Z.S.,

on March 19th.

1 Kordofan Giraffe (Giraffa camelopardalis antiquorum) 3,

born in the Menagerie on March 13th.

1 Greenland Falcon (Hierofalco candicans), captured in Mid-

Atlantic, and presented by Lt.-Col. F. B. Drage, R.H.G., and

the Hon. John Hubert Ward, C.V.O., on March 30th.

2 Purple Kaleege Pheasants (Genneus horsfieldi), from the

Himalayas, presented by the Marquess of Tavistock, F.Z.S., on

March 3rd.

2 Keysser’s Cassowaries (Casuarius keysseri), from German

New Guinea, received on deposit in immature plumage last

September, but identified as new to the Collection on March

12th.

4 Banded Trichogaster (7'richogaster fasciatus), from India, new

to the Collection, purchased on March 14th.

1 Central-African Mud-fish (Protopterus cthiopicus), from

Uganda, new to the Collection, presented by C. W. Woodhouse,

Esq., on March Ist.

2 Gibbous Scorpions (Buthus gibbosus), from Sardis, Asia Minor,

new to the Collection, presented by Miss Flora Russell on

March 11th.

Mr. H. J. Exwes, F.R.S., F.Z.S., exhibited the head of an

Asiatic Wapiti (text-figs. 83 & 84), remarkable for the outward

extension of the bay or bez tines, and made the following

remarks :—

“The antlers I exhibit were bought in Moscow on my return

from Formosa in May 1912 of Mr. Lorenz, who informed me that

ASIATIC WAPITT. 943

they had come direct from the Sayansk Mountains, east of the

Upper Yenesei River. The only original account I know of the

Deer of this region, though no doubt more recent information

exists in Russian, is by Radde in ‘ Reisen im Siiden yon Ost-

Sibirien,’ St. Petersburg 1862, vol. i. p. 284. Radde considered

the species to be C. elaphus, and says that it was found up to

Text-fig. 83.

Front view of antlers of Asiatic Wapiti from the Sayansk Mountains.

and above tree-level on the Sayansk, Baikal, Apfel, and Chingan

Mountains, frequenting the thickest forests and islands on the

Amur, but wandering in summer as high up as the glacier

of Munku-Sardik and over the bare peaks of the Sayansk

Mountains.

544 ON THE ASTATIC WAPITT.

Semi-profile view of antlers of Asiatic Wapiti.

MR. R. H. BURNE ON MALFORMED ANTLERS. 545

“He gives the rutting cry, as imitated by the native hunters

with a hollow stem of Calisace daurica, as follows, which seems to

me to have some resemblance to that of the American Wapiti :—

“In a paper read by me at the Linnean Society on Dee. 15,

1898*, I spoke of some horns from the Yenesei Valley in the

St. Petersburg Museum, which [ thought had more resemblance

to those of Cervus elaphus, having a distinct cup or crown

of 6 or 7 tines branching from the same point on the beam, as

sometimes seen in old Kuropean specimens of C. elaphus, but

never, so far as I know, in any other Asiatic form of Wapiti.

But I have not been able to examine them recently; and the

horns shown to-night belong, without doubt, to a race of the

Asiatic Wapiti known as C’.. eustephanus Blanford, which name I

think preferable to C. canadensis, var. siberica Severtzoft, though

the latter has priority. But until the races of Asiatic Wapiti

are better known, I think it is premature to name them definitely,

and it cannot be done without careful examination of specimens

in the St. Petersburg Museum.

“The bez or bay antlers in my specimen (text-fig. 83) are

clearly abnormal in their position. The development of the back

tines, though clearly of Wapiti type, is also poor.”

April 22, 1913.

EH. T. Newron, Esq., F.R.S., in the Chair.

Mr. R. H. Burne, M.A., F.Z.8., exhibited two pairs of

malformed antlers of an Axis Deer (Cervus axis), lately presented

to the Museum of the Royal College of Surgeons by Capt. Stanley

S. Flower, F.Z.8. The deer was born in the Giza Zoological Gardens

in 1899, the malformed antlers being shed in 1905 and 1906. The

second pair showed a common malformation, i. é. duplicity of the

brow tine, but the first pair suggested rather an injury during

growth than a congenital malformation, Ata similar point in

each antler the beam was abruptly bent inwards upon itself at a

very acute angle. The apical tines also were stunted, probably

owing to an interference with the blood-supply brought about by

the sharp bend in the beam.

* Journ. Linn. Soc., Zool. xxvii. p. 23.

546 DR. R. BROOM ON

Mr. E. G. Boutencer, F.Z.S., Curator of Reptiles, exhibited a

number of living specimens of the Leaf-Insect (Phyllium cruri-

folium), presented to the Society by Dr. Alfred Russell, which

had been reared from eggs laid in captivity, and which showed

various stages of development.

May 6, 1913.

Dr. Henry Woopwarp, F.R.S., Vice-President

s ? i ? ?

in the Chair.

Mr. E. G. Boutencer, F.Z.5S., Curator of Reptiles, exhibited a

living melanistic specimen of the Green Lizard (Lacerta viridis)

recently received from Dalmatia.

A new Species of Golden Mole.*

Dr. R. Broom, C.M.Z.S., exhibited an example of a new species

of Golden Mole from the Transvaal, of which he gave the following

account :—

‘“‘ For a couple of years I have known that a species of Golden

Mole occurred near Johannesburg in the Transvaal, but until a

few weeks ago the only specimen I had seen was a badly stuffed

skin without the skull. Recently I found that two specimens

had been ploughed out on the farm Vischkuil, near Springs, and

had been skinned by a native. Fortunately the complete carcase

of one was still to be had, and most of the skin. On examining

the skull I found to my great surprise that the Mole is nota

typical Chrysochloris, but belongs to the subgenus Bematiscus,

hitherto only known from Natal and EKastern Cape Colony. The

species is a near ally of Lematiscus villosus Smith, one of the

least known South African forms.

‘““So few specimens of BL. villosus are known that we do not at

present know the degree of variability of the species, and thus

cannot be quite sure whether the few specimens at present placed

under that specific name really all belong to one species. The

type specimen described by Smith is in the British Museum. It

is nearly full grown, and is stated to have come from Natal. The

underfur is of ver y fine texture, and of a slaty-grey colour.

The long hairs, which are compar atively few in number, have the

outer, flattened portion 10 to 15 mm. in length and of a pale

greyish-brown tint, rather darker towards the tips. The fewness

of the bristly hairs and their length give the fur a rough harsh

feeling.

* [The complete account of this new species appears here; but since the name

and a preliminary diagnosis were published in the ‘ Abstract,’ ‘it is distinguished by

being underlined.—Ep1ror. |

A NEW GOLDEN MOLE, DAT

“The skull of Smith’s type is well figured by Dobson, and

though barely mature is sufficiently near full size for purposes of

comparison,

** Dobson described and figured a second specimen which he

believed to be Chrysochloris villosa, but he recognised differences

which he thought might be specific. When working at the

Golden Moles some years ago, I thought it well to keep the

varieties of 6. villosus under the one species, though I called

attention to the fact of one of the specimens in the Maritzburg

Museum having a dark slaty, almost black fur. The specimens

are so few in number—there being, so far as I am aware, only

five skins in the museums of the world—that one hesitates to split

up the Natal supposed B. villosus specimens into two species, but

it seems likely that when more specimens are obtained at least

two subspecies and possibly even species will be recognised.

““ Whatever be the case with the Natal specimens, there is no

doubt that the present Transvaal specimen is not Smith’s Chryso-

chloris villosa, and almost as certainly it is not the same as

Dobson’s type.

“The Transvaal specimen, for which I propose the name

BEMATISCUS TRANSVAALENSIS,

Abstract P. Z.S. 1913, p. 25 (May 13),

has a thick soft fur very unlike that of Smith’s B. villosus. The

underfur is much shorter and less woolly, and the terminal

flattened portion of the hair is less bristly and very much shorter,

being only about 7 or 8 mm. in length, instead of 10 to 15 mm. as

in B. villosus. The terminal portion of the hair isa rich reddish-

brown, becoming very dark towards the tip. The whole dorsal

side of the animal has thus a reddish-brown tint. On the

abdominal surface the reddish tinge has almost disappeared and

the fur has a slaty tint. The fur from the sides of the head

and nose has been destroyed.

“The claws of the manus of &. transvaalensis measure:

Ist, 3°7 mm.; 2nd, 16°5; 3rd, 10; 4th, 1:5. The hind foot

measures 16 mm., and the length of the body is about 155 mm.

The type is a female.

“The skull differs from that of b. villosus in a number of

characters. The huge crest which rises up from the zygomatic

arch is even better developed than in B. trevelyani, and a plane laid

across the tops of the crests is 2 mm. above the cranial wall. In

B. villosus the crests pass forwards and outwards, and ata distance

of 7 mm. in front of the top of the occiput are 10°5 mm. apart

in Smith’s type. In Dobson’s type they are 10 mm. apart. In

B. transvaalensis the crest runs forward for some distance nearly

parallel, and at a point 7 mm. in front of the occiput are only

6-5 mm. apart. Another important point in the’skull is the very

much larger temporal bulla. In J. villosus the bulla is about

7 mm. in diameter, in B. transvaalensis it is 10 mm. in diameter.

Proc. Zoou. Soc.—1913, No, XX XVII. 37

548 ON A NEW GOLDEN MOLE AND YOUNG LIZARDS.

‘¢he followine skull measurements (in millimetres) indicate

the slight differences in size :—

Length. Breadth. Height. Dental Series.

Ga SSOUMIEUS UNOS go soc0oeosa0 c55000 33 20°5 15:5 13°5

Dobson’s specimen............ 34 22 17 1B

B. transvaalensis ............ Bylo, D355 es 14:5

“The teeth in the specimen of B. transvaalensis are in front

partly the milk and partly the permanent set. The Ist upper

premolar has no internal cusp. The molars are appreciably larger

than in B. villosus. In B. villosus the 2nd molar measures 2 mm.

in width; in B. transvadlensis it measures 2°5 mm.; and there

is an even greater difference in the size of the last molar.

“The type has been deposited in the British Museum.”

Dr. Broom also exhibited an adult female of the large 8. African

Lizard, Zonurus giganteus, with two newly-born young (PI.

LXXTIV.), and three adult male specimens of the allied species

Pseudocordylus microlepidotus.

He gave the following particulars of Zonuwrus giganteus :—

“The mother was given to me by Mr. F. W. Fitzsimons, F.Z.8.,

of the Port Elizabeth Museum. On opening the box containing

the specimen on arrival in London it was found that two young

had been born on the voyage. The young differ in appearance

very considerably from the mother, being much more brightly

coloured. Young specimens in the British Museum collection

are described by Mr. G. A. Boulenger as “light yellowish,

marbled and cross-barred with blackish brown ; the spines not at

all developed, and all the scales more strongly imbricate.”

‘These young also show the marked imbrication of the scales

which is largely due to their having no supporting dermal ossi-

fication. This gives the tail especially avery different appearance

from that of the adult. The colour may be described as yellowish,

with, on the back, irregular cross-bands of black. The top of the

head is blackish, but in most of the antorbital region the yellowish

colour predominates. The scales round the eye are mostly bright

yellow, but there is a black spot on the upper eyelid and a less

distinct one on the lower. The 4th lower labial has a large black

mark, and a black mark extends above this on the upper jaw to

the eye. The scales above the tympanum are yellow. On the

back the light bands, especially towards the lower half, become

very distinctly reddish, and on the tail about half the scales are a

light brick-red. The legs are irregularly banded yellow and

black, and the front of the body is pale straw-coloured except the

neck, which has a number of large black spots. The young

measure about 5 inches in length. Mr. HE. G. Boulenger assures

me that the young are feeding satisfactorily and look lively.”

“SNALNVOID SNYNNOZ

‘jap “uaadyy “£7

Cinquvg “pry ‘u0g Gg au0rg &

INDXOXal aid Suey sSeZid

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= y

ON MAMMALIAN CESTODES. 549

PAPERS.

36. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea. By Frank HE. Bepparp, M.A.,

D.Sc., F.R.S., F.Z.8., Prosector to the Society.

[Received April 8, 1912: Read May 6, 1913.]

(Text-figures 85-94.)

X. On Two Spectres oF TAPEWORMS FROM GENETT.A DONGOLANA.

INDEX.

Page

Dipylidiim dongolense; Sp. Ne ...-----2--never se cee 2-2 549

Diplopyliduuni Sen NOVe seers eee eseeee ee eeske ead OOO

DDG CWECEE ASPs, Men det rir siasisteateceee meee TIT eeiee 559

An examination of the small intestine of a Dongolan Genet

(Genetta dongolana) yielded about a dozen small tapeworms and

the hinder part of the body of a larger worm. ‘The latter I have

not studied ; the former belong to two species which are described

in the present paper. Inasmuch as the Genet, which died in

November of last year, had been about four years in the Gardens,

it is uncertain whether the parasites are to be regarded as indi-

genous to Africa or had been acquired in captivity. From this

particular variety of the Genet I believe that no tapeworms have |

been recorded; but from allied forms species of Cestodes are

known. v. Linstow, in his ‘Compendium der Helminthologie,’ *

mentions three from “ Viverra genetta,” viz. Mesocestoides ambiguus

Vaillant 7, Tenia platydera and 7’. geneite, both of Gervaist. The

latter, as I point out later, may be identical with my Dipylidiwm.

Tenia platydera seems$ to have no rostellar hooks, and may

quite possibly be an Anoplocephalid. I refer later to other species

from Genets in my description of Dipylidiwm dongolense.

(1) Dipylidium dongolense, sp. n.

There were altogether five specimens of this species, all of

which I have carefully examined either entire or cut into series

of transverse and longitudinal sections. The largest specimen,

after preservation in alcohol, measures just 5 mm. in length

and 1 mm. in breadth at the broadest part. During life

individuals reached a length of 6 mm. The accompanying

illustration (text-fig. 85) represents the individual referred to as

the largest. The scolex is not sharply marked off from the

ensuing neck, which is very short and soon passes into the

* Hannover, 1878, p. 37.

+ Comptes Rend. Soc. Biol. Paris, 1863, p. 48.

{ Mém. Ac. Sci. Montpellier, 1847.

§ Cf. Diesing, Syst. Helminthum, i. p. 519.

Sit

550 DR. F, E. BEDDARD ON

strobila, The neck is at once wider than the scolex. The pro-

truded rostellum has several rows of the usual minute thorn-like

spines characteristic of the genus Dipylidium. The rostellum has

a three-pointed appearance, which will be obvious from the

drawing, the narrowest part being the apex.

Text-fig. 85

Dipylidium dongolense.

View of entire worm, magnified.

p. Extruded cirrus. 7. Rostellum.

T believe that there are four rows of spines upon the rostellum.

The four suckers are not very large. The shape of the terminal

segment of the body seems to me to prove that the specimen

which I here figure is a complete individual with no posterior

segments shed. It will be noticed that the proglottids are

nowhere longer than broad, and that they overlap laterally. The

segmentation of this species is thus very different from that of

MAMMALIAN CESTODES. 551

the type species of the genus, Dipylidiwm caninum. An in-

spection of the figure will render a detailed description of the

proglottids of the present species unnecessary ; it will be noticed

that their number is small, not more than 28. In another

specimen I only found 22. In this example (or in one very like

it) the last nine segments became suddenly very much more

elongate, and were three to four times as long as broad. In

transverse sections the body shows an elliptical form pointed at

the two ends laterally and nearly as deep as wide.

The general structure of the body shows certain differences in

the less mature and more mature segments, which do not appear

to me to be altogether due to the greater tension in the latter.

The specimen which I selected for examination by means of

transverse sections was much like that of which I give an entire

view in text-fig. 85. But in the posterior segments, though

short, the sexual organs were well developed, and there were

ripe ova lying in cavities (of which a full description will

be given later). In these transverse sections the cuticle is

very thick and the layer of subcuticular cells very conspicuous

and deeply stained; they have the usual flask-shaped form

and lie in a dense layer, being closely adpressed. Their thinner

outer ends are in contact with the cuticle above them. Upon

this layer follows a layer of lax tissue, and then a strong

longitudinal layer of muscular fibres, which are themselves

separated from the medulla by a thinner layer of transverse

muscular fibres. These fibres are associated together in bundles

of three or four fibres, which are very stout ; there appear to be

also a few fibres to the outside, which are not associated in

bundles but implanted singly. In longitudinal sections it is

rather easier to count the number of fibres which lie in a single

radial row of this longitudinal layer, and I find that there are not

more than five or six. Here and there between the bundles are

parenchymal cells whose deep staining as contrasted with that

of the muscular fibres emphasizes their existence. In sagittal

sections the subcuticular layer is also obvious in the more anterior

proglottids. I have not been able to get any transverse sections

of the more elongated posterior proglottids (owing to the limita-

tion of my material); but in longitudinal (sagittal) sections I

could find no trace whatever of elongated flask-shaped subcuticular

cells. The general cortical parenchyma reaches absolutely up to

the (here) very thin cuticle. There are, I think, only two possible

explanations of this appearance of vanished subcuticular cells.

First, that they have altered their form owing to the pulling

out of the segment, and have become broader and round like the

cells which secrete the calcareous bodies; or secondly, that they

are really absent from this region of the body. It is clear from

the observations of Lonnberg that the subcuticula varies among

tapeworms. It is, for example, in Tetrarhynchus tetrabothrius* ~

* “ Anatomische Studien tiber Skandinavische Cestoden,” K. Svensk. Ak. Handl.

xxiv. 1891, pl. i. figs. 6, 11.

552, DR. F. E. BEDDARD ON

a layer two or three cells deep, of which the outer ones are close

to the cuticle. On the other hand, in Ptychobothrium belones the

more scanty subcuticula lies in the parenchyma some way below

the cuticle. I find on a re-examination of sections of Hyracotenia

hyracis* that the subeuticula occurs in patches on the mature

segments, and seems, therefore, to be in course of disappearance.

It is, moreover, mentioned by Gough? that in Séilesia the sub-

cuticular layer is not to be found in ‘older portions of the strobila

in segments where the paruterine organ is fully developed.”

Gough holds that this absence of the subeuticula in mature

segments is a secondary character. The state of affairs seems to-

bear out the suggestion that in my Dipylidiwm the subcuticula

has actually disappeared ; in the mature segments.

The testes occupy all the available space in the proglottids, and

are thus mainly between and posterior to the ovaries. In the

more elongated and riper proglottids the testes are only one row

deep and not more than 6—9 (according to the length of the

segment) in a single row; they are thus evidently not so

numerous as In some species.

The two generative orifices lie one on each side of the body not

far from the anterior boundary of the proglottids. The ewrrus-sac

is in front of the vagina, and, as is shown in sagittal sections, in

the same straight line with it, being neither dorsal nor ventral.

The cirrus-sac has very definite, rather thick muscular walls,

which are even quite obvious when the sac is dilated in mature

segments, though not so thick in appearance. The cirrus is long

and much coiled within the cirrus-sac. The sperm-duct is also

much coiled, and the two sperm-duct coils wey nearly come into.

contact in the middle of the segment.

The ovaries of this Dipylidiwm are, as a rule, clearly two in each

proglottid lying rather laterally and anteriorly. They are, in fact,

like those of other species, such as D. zschokkei of Hungerbihler,

as to their position in the proglottid. But I do not find in cases

which I have particularly examined that there is an absolutely

complete separation between the two ovaries of a given proglottid.

In this matter I refer to rather anterior proglottids which are

not elongated and stuffed with ripe eggs containing embryos.

However, in other proglottids I found a distinct separation between

the two ovaries, the fact being that they approach very near to

each other in the middle line of the segment. There is, however,

no doubt about the entire separation of the vitelline clands, each

of which lies behind the ovary of which it is an adjunct. The

ovaries occupy a good deal of the space in the short proglottids

where they occur when full sized. They are posterior to the coils

of the vas deferens, but reach very nearly to the posterior end,

where the testes occur ; the testes also occupy the available space

between them.

The receptaculum seminis penetrates the middle of the ovary

* For species, see P. Z.S. 1912, p. 593.

+ “A Monograph of the.... Avitelline,”’ Q. J. M.S. lvi. 1911, p. 345.

MAMMALIAN CESTODES. 553

and is greatly swollen so as to be almost spherical in this region.

It is several times the diameter of the vagina.

The wterws in the genus Dipylidium is thus described by Ran-

som in his definition of the genus * :—‘“ Uterus at first reticular,

later breaking up into egg capsules, each containing one or more

Text-fig. 86.

Dipylidiun dongolense.

A completely mature proglottid seen in sagittal section.

m. Longitudinal muscles. o. Ripe embryos, each in a separate cavity.

¢. Remains of testes.

eggs.” Nearly the same definition, but not quite so explicit, is

given by Fuhrmann t, who writes :—‘‘ Der Uterus lést sich in

* “Cestodes of N. American Birds,” Bull. U.S. Nat. Mus. no, 69, 1909.

+ “Cestoden der Végel,” Zool. Jahrb. Suppl. vol. x. 1908.

554 DR. F. E. BEDDARD ON

einzelne ein oder mehrere Hier einschliessende Siickchen auf.”

These definitions are, as I presume, chiefly based upon the many

observations made upon the best-known species of the genus,

viz. Dipylidium caninum (=Tenia cucumerina, mult. auct.),

of which they are a correct restatement. But Fuhrmann has

himself described in an Avian Dipylidium* a series of sinuous

uterine tubes preceding a series of ege-capsules each containing a

single egg. Furthermore, although he gives no details, Diamare f

asserts that the development of the uterus in other species is as

in D. caninum.

Text-fig. 87.

Dipylidium dongolense.

A portion of the section represented in text-fig. 86, more highly magnified

to illustrate the absence of a special lining membrane to the egg-holding spaces.

Lettering as in text-fig. 86.

In the species of which the present paper is an account I can

find no confirmation of Diamare’s statement concerning the de-

velopment of the uterus, though his figures of the completely :

mature uterus of D. trinchesti and D. pasqualii agree very closely

with what I have seen in mature proglottids of D. dongolense (see

text-fig. 86). In the largest specimen which I have been able to

examine, in which the last nine proglottids were elongated, it was

only the last two which agreed really closely with the figures given

by Diamare and just referred to. In these alone the embryos were

contained each in its own separate compartment, which it did not

by any means fill. I lay particular stress upon the fact that they

* “Taenien der Raubvoégel,” CB. Bakt. u. Paras. xli. 1906.

+ “Tl genere Dipylidium,” Atti R. Acc. Napoli (2) vi. 1894.

MAMMALIAN CESTODES. 55D

were embryos, since in Diamare’s figures they are clearly eges

which have not yet segmented to form emb1 ‘yos. In Dip alidlinm

caninum (as 1 suppose), an example of which I have studied from

the Asiatic Jackal, the separate chambers of the mature uterus

were very plainly lined with a cellular layer. J may observe of

this Dipylidium incidentally that the number of embryos in a

given chamber was very much greater than what is figured in

that species by Diamare, who only represents 3-7. I found quite

30 or more.

Text-fig. 88.

Dipylidium caninwn.

A portion of uterus in a mature proglottid.

ep. Epithelial lining of uterine cavities. O. Embryos.

In Dipylidium dongolense there was not, so far as I could see,

any special lining membrane to the small chambers which lodged,

each of them, a developing embryo. It will be observed in the

text-figure (text-fig. 87) which illustrates the mature “ uterus”

of the present species, that the appearances presented are con-

sistent with the view that the chambers which contain the eggs

are merely interstices in the general medullary network. Their

walls are perfectly continuous with the medullary tissue lying to

the inside of the longitudinal muscles and forming the peripheral

layer of the medulla. Though the walls of the cavities often

contain imbedded in them crowded nuclei, it must be borne in

mind that the peripheral layer of the medulla is also crowded

556 DR. F. E. BEDDARD ON

with quite similar nuclei. Furthermore, the nuclei (see text-fig.

87, p. 554) do not give the impression that they line the cavities ;

they belong rather to the tissue of the walls: there are, moreover,

often considerable tracts without any nuclei at all, as is often

the case with the medullary tissue of tapeworms. This is, of

course, opposed to the idea of a lining membrane.

Text-fig. 89.

Dipylidium dongolense.

A portion of a section through a proglottid less mature than that represented

in text-figs. 86 & 87.

m. Longitudinal muscles. o. Egg-holding spaces.

All this contrasts greatly with the uterine spaces of D. caninwm

(see text-fig. 88, p. 555), where there is invariably a close row

of nuclei with accompanying protoplasm actually lining the

cavity and in which the nuclei of the medullary tissue are

not numerous and, as a rule, different from those of the lining

membrane. ‘The thedulla, in fact, forms a reticulum in éhe

meshes of which lie the developing embryos; there is not,

at any rate in the fully mature stage, any uterus at all. A

1) |

MAMMALIAN CESTODES. DDT

consideration of the disposition of the eggs in proglottids anterior

to the two that have just been described bears out this view.

In the two proglottids anterior to the two terminal ones, the

arrangement of the eggs, or rather developing embryos, in the

medullary parenchyma was as follows :—The medullary region is

not so entirely given up to these eggs as it isin the case of the

last two segments that have been described above. In the two

latter but slight traces are left of the testes; in the less mature

proglottids more testes are found. They have, however, com-

menced to disappear, for in the proglottids antecedent to these the

testes are more abundant and therefore take up more room. The

space apparently thus left by the degeneration of the testes is

occupied by the developing eggs. It is not, however, a continuous

cavity running from end to end of the proglottid (see text-fig. 89).

It is divided here and there by delicate trabecule into compart-

ments, each of which contains a considerable and varying number

of developing embryos*. Further forward still, the embryo-con-

taining spaces are less conspicuous owing to the full development

of the testes, which thus occupy more room. JI can draw no line

of demarcation between these cavities in which lie the developing

embryos on the one hand, and spaces which lodge ova and form

actually a part of the ovary on the other hand, in the more

mature proglottids which have begun to become elongated.

As already mentioned in describing the ovary of this species

of Dipylidium, the ovaries are mature or, at any rate,

possess plenty of full-sized ova in proglottids which are not

at all elongated and are, indeed, still broader than long. In

these proglottids, however, there is no trace that I could find

of a definite uterus, and nothing at all like the obvious retiform

uterus of not fully mature proglottids of D. caninum. However

we may interpret the embryo-holding spaces of D. dongolense, it

is clear that they cannot arise in the same way that the egg-

sacs of D. caninum arise, i.e., by a breaking up of a continuous

retiform uterus into a number of discontinuous sacs.

With what, then, can we compare the spaces which surround

the developing ova in this and other species of Dipylidiwm—in

fact, possibly in all with the exception of D. caninum? I do

not think that any genus has been described as possessing an

exactly similar mode of lodgment of the developing eggs. But

there are nevertheless some comparisons to be made with other

forms.

The final stage shown in Dipylidiwm dongolense, where the

eggs are uniformly scattered through the medullary region, each in

its own separate compartment, is suggestive of Oochoristica, with

which genus Dipylidiwm is usually considered to be properly

associated. The comparison, however, can hardly be exact, since,

as we have seen, this final stage is preceded by larger spaces con-

taining each of them many developing embryos. Jt seems to

* Tn one specimen I have also seen ova enclosed by threes and fours in cavities in

the cortical layer. But this seems exceptional.

558 DR. F. E. BEDDARD ON

me that we must come to the conclusion that a growth of the

medullary parenchyma between the individual eggs produces the

final result seen in the terminal and penultimate proglottid of

the specimen which I have studied. It might be argued that

this process is really comparable to the formation of paruterine

organs which are growths of medullary tissue encapsuling the

eges though with histological change. In Dipylidiwm we have

no histological change, but all the same encapsuling growths of

parenchyma.

Having given an account of the structure of this Dipylidium,

it remains to be decided whether it be a new species or identical

with one of those already described. It is, as I think, hardly

necessary to say much of D. caninwm, with which there can be no

possibility of confusing the present species. The much greater

size of that species and the form of its uterus at once distinguish

it from that which is the subject of our present communication.

It seems indeed to me that the genus Dipylidiwm requires really

division into two genera—J/. caninum belonging to one, and all

the other species that are adequately known to the other. It is

from the latter that it is not so easy to distinguish D. dongolense

of the present paper. If D. genette of Gervais * and D. gervaisi

of Setti were not much larger species than mine, I should have

been inclined to identify it with one or possibly both of them.

D. echinorhynchoides (from Megalotis cerda) is much larger than

my species, has a much longer rostellum and a longer neck.

D. trinchesti is a smaller species, but still considerably larger

than D. dongolense, measuring 2°5 cm.; it has, however, a

short rostellum, like that of my species; but it is to be

distinguished by the greater length of the mature segments,

which do not overlap, and the invariable inclusion of each ripe

ovum in a separate compartment. Though a receptaculum

seminis 1s said to be present, it is hardly represented in the

figure. D. pasquali, on the other hand, has an obvious recep-

taculum {, but it is a much larger species, 7.e. 200 mm. The

uterus is as in the last, and therefore differs from that of D. don-

golense; but it is to be noted that the ripe ova extend into the

cortical layer, as do the egg-chambers of D. dongolense. Occurring

as 1t does in the intestine of a closely allied species, it might be

expected that D. gervaisic of Setti $ would be more closely allied

to D. dongolense than the other species of Dinylidiwm referred to.

It is not, however, as it would appear, any nearer in structure.

For in this species, as in most others, the ripe ova are figured as

embedded separately in the parenchyma, without any previous

inclusion in larger cavities. It is also rather larger, being from

1-4 em. in length.

Nor can the present species be confused with D. zschokkei ||.

* Mém. Ac. Sci. Montpellier, 1847.

+ Diamare, loc. cit. pl. 1. fig. 9. { Loe. cit. pl. i. fig. 8.

§ “ Dipylidium gervaisii,’ Atti Soc. Lig. Sci. Nat. vi. 1895.

|| Hungerbiihler, “Studien au Gyrocotyle und Cestoden,”’ Denkschr. Ges. Jena,

xvi. 1910, p. 516.

MAMMALIAN GESTODES. 559

For the latter reaches a length of 120 mm., and the ripe pro-

glottids may be four times as long as broad. Moreover, the neck

of D. zschokket “muss als sehr lang bezeichnet werden,” It

appears also that my species has a much more pronounced

receptaculum, which is mentioned by Hungerbihler as existing

in D. zschokkei, but is hardly indicated in his figure*. This

species occurs in Cynictis penicillata. Dipylidium triseriale of

Lihe also appears to odin a species which occurs in the African

Civet cat. For Liihe says of that species that the ripe eggs are

imbedded singly in the parenchyma, and makes no mention of

previously existing spaces, a state of affairs which I have described

in my species. Moreover the genital pores of D. triseriale are

further back than in my species, and it is larger in size, In

D. triseriale, moreover, the proglottids are described by Liihe

as being “ungefihr quadratisch,” which I presume means that

they do not overlap as In my species.

A second species, Dipylidium monoophorum, 1s also described by

Liithe from the same host, which only measures 10 em. in length

and therefore more nearly approaches the dimensions of ). dongo-

lense of the present paper. But in this species, as in D. triseriale,

the length of the ripe proglottids is much greater; and further-

more, the two separate small round ovaries (with the vagina to

the outside of each) are not what I have seen in D. dongolense.

Diamare = has described, besides the well known JD. caninum,

D. echinorhynchoides Sonsino, two species named by himself,

viz. D. trinchesti and D. pasqualii (both from the common cat),

and two doubtful species, which are D. (?) genette Gervais and

D, monticellir,

Diplopylidium genettz, gen. et sp. n.

I refer what I believe to be a new species of Tapeworm to the

new genus Diplopylidiwm for reasons which I shall discuss after

setting forth the structure of this worm. The intestine of the

Dongolan Genet contained fewer specimens of this worm than of

the Dipylidiwm which has been described above. They were,

however, of about the same size, measuring up to 6 mm. or so

in length, and consisting of about 28 segments at most.

The accompanying drawing (text-fig. 90) represents an average

sized example to which the above statements apply. The scolex

is as broad as the body which follows, and there is only a slight

increase up to the end of the body. The worm has thus a sturdy

form, and is so far much like the Dipylidiwm which has just been

described. It agrees with it, moreover, in thickness, the trans-

verse sections being oval in outline and not much flattened from

above downwards. The scolex is furnished with an armed ros-

tellum, which is large in contrast to the comparatively small

* Toc. cit. pl. xix. fig. 18.

+ “Beitrage zur Helminthenfauna des Berberei,’ SB. Ak. wiss. Berlin, 1598

p- 626.

ft “Il Genere Dipylidium,” Atti R. Acc. Napoli (2) vi. 1894.

560 DR. F. E. BEDDARD ON

suckers. These latter are unarmed and show no peculiarities of

moment; they face outwards and slightly forwards. The hooks

of the rostellum are large and arranged in two rows, 17 to each

row. Their shape and relative size can be understood from an in-

spection of text-figure 91, which represents a longitudinal section

Text-fig. 90.

Diplopylidium genette.

View of entire worm, magnified.

g. Extruded cirri, 7. Rostellum.

through the retracted rostellum of this species. The rostellum is

a muscular one, but with fewer external layers of muscles than is

sometimes met with, which act as protractors of the rostellar bulb.

The stout retractors (see text-fig. 91) are of course continuous

MAMMALIAN CESLODES. 561

with the longitudinal muscular layer of the body, which is poste-

viorly not thick but quite evident. In comparing the present

species with Dipylidium dongolense in the living state, I noted a

difference in the dispersion of the calcareous corpuscles. In both

species they were abundant throughout the body generally; but

in the Diplopylidium alone did I succeed in detecting any in

the scolex.

Text-fig. 91.

Diplopylidium genette.

Longitudinal section through anterior end of body.

FH. Hooks on rostellum. S. Sucker.

The éestes of this species are numerous in proportion to the

size of the proglottid. They are ripe in the earlier seg-

ments, where the ova are not mature, but still attached to the

ovaries. In such segments the testes are closely crowded

together and fill all the availabie space of the proglottid from

one end to the other. They lie, however, within the ovaries,

thus occupying the middle of the proglottid. I counted up to

twelve or so in a single longitudinal row. The cirrus-sac of

562 DR. F, E. BEDDARD ON

the worm is large and has an obvious and thick muscular coat.

A peculiarity about this organ is that it does not he in a straight

line extending inwards from the point of opening on to the

exterior, and thus at right angles to the long diameter of the

proglottid, as is so usually the case. The sac, on the other hand,

often bends at right angles near to its external pore and passes

backwards ; at other times it is more irregular in its folding.

Its large size is responsible for this folding, as the two cirrus-sacs

of a given segment would meet in the middle line did they run

straight inwards from the external pore. The cirrus itself is a

very fine tube upon the end of which I could detect no spines ;

it is coiled within the cirrus. The sperm-duct after emerging

from the cirrus-sac forms a coil which is anterior to the sac and

near to the front of the segment.

The vagina of this Cestode together with adjacent parts of the

genital system is shown in text-fig. 92, The most remarkable

fact is that that tube opens anteriorly to the cirrus, a position

which is made plain in the text-figure referred to. It would

also appear from a study of a series of sagittal sections—which

are the best in which to study these particular relationships—

that the vagina has not a separate orifice on to the exterior of

the body. But the truth of this conclusion will be doubted after

an inspection of text-fig. 92. For in that figure the vagina

seems to open separately, and of course anteriorly to the cirrus-

sac. It may be that this is at times the case; but it is also

clear that in tracing the vagina in a series of sagittal sections the

tube is lost sight of in the walls of the male duct before the latter

reaches the exterior. I believe that among the Tetracotylea the

genus Tetrabothriwum is the only genus in which the female

openings are in front of the male. It is true that in the Ichthyo-

teniide generally, if not universally, there is an irregular alter-

nation in these conditions, the female duct lying in one segment

in front of and in another behind the male pore. But it is

doubtful whether this family is to be safely referred to the

Teeniade. The figure referred to (text-fig. 92 C) shows the narrow

vagina lying in front of and parallel with the stouter cirrus-sac,

the two forming in this region a perfectly straight line, so that

their relative positions is a matter of ease to ascertain. I found

the same relations in other series of sections besides that from

which the text-figure referred to has been taken. Thus I am

able to state that there is not an alternation in the positions of

the male and female openings, as in Jchthyotenia. The vagina

is darkly stained and of small calibre in this tapeworm, the small

bore of the tube bearing a relation to the fineness of the cirrus.

Another remarkable circumstance is the nature of the recep-

taculum seminis and less mature and more mature proglottids.

In text-figure 92C the slender vagina is seen to open into a minute

spherical chamber or dilatation and to issue from the opposite side

as an equally slender tube. This chamber is, as I suppose, to be

compared to a receptaculum seminis; but it has in this section

MAMMALIAN GESTODES. 563

the appearance of the mere beginning of such a chamber. There

is no trace of sperm to be found in it, and it is so small that I

have not been able to detect it in other sections of neighbouring

segments. The particular section to which attention has been

Text-fig. 92.

(E Wipics

Three sections through the generative region of Diplopylidiwm genette.

-A. Section to show relations of vagina (@ ) and cirrus-sac (¢), the latter lying

behind the former.

B. A transverse section through the receptaculum seminis and adjacent parts

of the generative system.

ce. Cirrus-sac. o. Vitelline gland. 7.s. Receptaculum seminis.

v. Vagina.

vit. Ovary.

C. Transverse section showing immature receptaculum seminis (7.s,) and

anterior position of vagina (v.).

c. Cirrus-sac.

N.B.—There is an appearance of a diverticulum of the cirrus probably due to

irregular staining.

Proc. Zoot. Soc.—1913, No. XXX VIII. 38

564 DR. F. E. BEDDARD ON

called is of a proglottid filled with completely mature eggs,

enclosed in their shell, and which have developed into embryos.

It is rather remarkable to find that the receptaculum is so small

and immature ina section which is otherwise quite ripe. For

the receptaculum does not suggest a reservoir from which the

contents have been recently expelled; it distinctly suggests an

incompletely developed receptaculum. On the other hand, text-

figure 92.B is from a section of which the maturity was less

advanced. The testes were fully ripe, and the ovaries quite

developed ; but there were no ova scattered through the paren-

chyma at all. And if I have missed any in the examination of

the section, they must at most have been few. Yet in this

section, as will be seen from the drawing cited, the receptaculum

seminis is very large and quite distended with abundant sperm.

Nor can this difficulty be explained away on the assumption that

one of the two vagine in the fully-ripe proglottid was more

mature than the other, and may have been the storehouse of the

sperm after copulation. For I ascertained that the vagine on

both sides of the body were in an identical state of maturity.

The ovaries in this species are distinctly double, and lie one on

each side of the proglottids a little way behind the point of

opening of the generative ducts. The ovary is in each case

inmediately Follow ed by the vitelline gland. There seem to be no

features of special interest about either the ovaries or the vitelline

glands. On the other hand, the cavities lodging the ripe eqys are

remarkable, and like those of but few other tapeworms that have

neon described. I do not give to these cavities the name of

“uterus,” for it does not seem to me to be proved that they

actually ‘correspond to the uterus of such genera as Choanotenia,

Hymenolepis, etc. ‘The spaces indeed to which I here refer are more

comparable, as it appears to me, to those of the Dipylidiwm, which

have already been deseribed in the present paper, and to be rather

lacune in the medullary network than deliberate pre-existing

cavities at first united to form a single cavity. I believe, imoleeal

that there is no such fraementation| here of a pre-existing uterus

as, for example, I have lately described * in my genus Otiditenia.

For it is hard to believe that a uterus like that of so many

tapeworms can exist in the present species. As already men-

tioned, in the more anterior proglottids, where the testes are

fully ripe and the ovaries fully formed, there is no trace of any

sac which might be considered to be a uterus. Nor, indeed,

among the closely adpressed testes which occupy so much of the

interior of these proglottids does it seem that room exists for

the development of a uterus. In any case nothing obvious of

the kind is visible. It is in riper proglottids which are more

elongated in their form that the “ uterine” spaces referred to are

first visible. They le (text-fig. 93) scattered among the testes,

* “Ona New Genus of Tapeworms (Otiditenia) from the Bustard (Hupodotis

kori), P.Z.S. 1912, p. 194.

MAMMALIAN CESTODES. 565

and are developed in the posterior region of the proglottid

only, but all the same occupy the greater part of such proglottids.

There is no regularity in the dispersal of these cavities, so far as

I have been able to make out. They lie here and there, and are

never crowded like the testes ; among them lie the remains of the

testes in the most posterior segments, where the testes are most

fully reduced. The cavities increase in size pari passu with the

development of the embryos contained within them. In the more

anterior part of the body the spaces are small and contain but a

single egg or embryo just commencing to divide, which does not

nearly fill the cavity.

Text-fig. 93.

Diplopylidium genette.

A portion of a nearly ripe proglottid showing ova (e.) lying im spaces in

the medulla.

In the centre of the field is one ovum which has no cavity round it.

Further back the cavities are larger to accommodate the large

embryo with its wide egg-shell. Nowhere is there anything of

the nature of a lining membrane to these cavities. No nuclei

are distinguishable as forming a layer immediately surrounding

them, though the nuclei of the medullary parenchyma are, of

course, to be found frequently in the neighbourhood of the

limiting layer of the cavities; but in these cases there is no

38*

566 DR. F. E. BEDDARD ON

relation to be observed between these cells and the egg-holding

spaces. I have never found more than a single embryo in one

cavity ; if there are occasionally more, such instances must be

rare. The accompanying drawing (text-fig. 94) shows a number

of the embryo-holding spaces in a fully ripe proglottid. It will

be noticed that between the embryo itself and the wall of the sac

in which it lies is a tumbled mass of a membranous appearance.

Text-fig. 94.

Po Oi

/ a ids

do i

™

Diplopylidium genette.

A portion of a more fully mature proglottid. The embryos are now surrounded by

a delicate often crumpled membrane, and the spaces in which they lie are

larger.

e. Embryo. m. Egg-membrane. w. Cavity surrounding embryo.

This is not all stamed by the hematoxylin, which has amply

stained the embryo itself and the surrounding tissues of the

medulla. The inference, therefore, appears to be that this

membranous material is in reality the egg-shell and is of

a chitinous nature. In some cases it has a more regular

MAMMALIAN CESTODES. 567

appearance, and seems to lie in concentric layers surrounding the

embryo; in others this is not apparent, and it even simulates a

retiform tissue in the laxity of its arrangement.

The ultimate arrangement of the uterine spaces in this worm

is, therefore, the same as we find in the species of Dipylidiwm

that has just been described. But the two genera differ in the

fact that this is preceded in Dipylidium by another developmental

stage not represented in Diplopylidiwm. In the latter we find

the scattered cavities lying among the remains of the discharged

testes, and as both a final and an initial stage. There is no inter-

mediate condition that I have found where the ova are contained

in larger cavities, which subsequently become reduced to chambers

each containing a single egg such as is found in Dipylidium.

The cavities in the two cases are, therefore, not exactly equiva-

lent; otherwise, as has been said, there is a close agreement

between the two worms. In neither can any lining membrane

be detected forming a wall to the ege-cavities. The actual size

of the individual spaces is much the same, but in Dipylidiam

they are more closely packed together. In formulating this

difference between the two genera I rely upon what I have seen

and not seen in my sections. It would be unwise—because of

the negativity of my conclusions—to assert this fact in a more

positive way by using it as part of the generic definition of

Diplopylidium. It is not only in Dipylidiwm that we find a

series of egg-holding spaces like those of Diplopylidium. The

same kind of arrangement occurs in Oochoristica, so named

on account of the scattering of the mature ova through the

parenchyma. But in this genus there is an obvious uterus which

exists before the final scattering of the eggs. Moreover, when

imbedded in the parenchyma, the appearance is as if the eggs

were firmly imbedded in the homogeneous ground material of the

medullary tissue and not received into cavities. The eggs, in

fact, are immediately surrounded by solid tissue and not by the

rest of a cavity. More like the disposition met with in Dipy

lidiwm and Diplopylidium is that which is to be seen in J/ono-

pylidium, a genus to which the present genus, as I point out

later, is probably related. But it should be remarked, first

of all, that the absence or presence of a cavity round the singly

imbedded ova is not a matter of great importance. For, as will

be seen by an inspection of text-fig. 93, it happens occasionally

in Diplopylidiwm that imbedded ova have no cavity round them.

This may well be a preliminary state of affairs, the subsequently

appearing cavities being due to the growth of the egg into the

embryo. In this case there will be no doubt whatever that the

egg-holding cavities of Diplopylidiwm are totally different from

those of Dipylidiwm dongolense, since the latter are formed as

subdivisions of an earlier larger cavity.

In Monopylidium the ova are in the same way scattered

through the parenchyma in cayities which contain one, or in

some cases more than one, ovum. It is evident, however, from

568 DR. F. E. BEDDARD ON

the definition of this genus by Ransom *, that he regards these

spaces as disjected portions of a pre-existing uterus. It seems

clear, from the illustrations given by Mola‘, that the egg-holding

cavities are not lined by cells in Zenia marchali (a species which

Fuhrmann refers to the genus J/onopylidiuwm), and in this fact

we find an agreement with the two tapeworms described in the

present paper. A very remarkable condition is described and

figured by Fuhrmann? in Monopylidium rostellatum. In this

species the eggs are scattered singly through the medullary region,

and each egg fits closely into a hexagonal mass of delicate tissue,

which itself fits equally closely into the medullary parenchyma,

leaving no gaps or spaces anywhere. It seems to me to be just

possible that this “‘ parenchyma vacuolaire”” which immediately

surrounds the egg may be simply the outer egg shell, which

I have myself observed in Diplopylidiwm, when crumpled, to

present quite the appearance of a retiform tissue. It is finally

to be noted that the disposition of the mature ova in Diplo-

pylidium is perhaps also to be compared to the scattering of the

eges in Inermicapsifer capensis § before the inclusion of these in

the parenchymal organs. But it must always be borne in mind

that in the case of Diplopylidium, Oochoristica, and Monopylidium,

completely mature proglottids may not yet have been seen. A

final stage may be the formation of paruterine organs, though

this does not seem to be very likely.

We may characterise this new form as follows :—Size small, up

to 6 mm.; number of segments up to 28. Scolex with retractile

muscular rostellum armed with two rows of hooks, 17 in each

row. Suckers unarmed. Neck short; posterior proglottids two

or three times as long as broad. They do not project posteriorly

or overlap. Generative orifices lateral and paired in each pro-

glottid, opening at or rather in front of the middle of the

proglottid. The ducts pass between the longitudinal water-

vascular tubes. The female orifice is situated in front of the

male, The water-vascular tubes are two on each side, the dorsal

being of narrower calibre; they are placed one above the other.

A longitudinal muscular layer, only a few fibres thick, separates

the cortical and medullary regions. The testes are numerous and

large, filling up all the proglottid not occupied by other parts of

the generative system ; testes disappear in posterior proglottids

when egg-chambers appear. The cirrus-sac is large, with muscular

walls, not lying in one straight line, but bent backwards in rela-

tion to the fact that there are two in each proglottid. The cirrus

is very slender and coiled within its cirrus-sac. The sperm-duet

is coiled and lies on each side in front of cirrus-sac. Ovaries

distinctly paired in each proglottid, lying laterally behind the

* “The Teenioid Cestodes of N. American Birds,” Bull. U.S. Nat. Mus. no. 69,

1909, p. 76: “ Uterus breaks down into egg-capsules.”

+ Bull. R. Ac. Belg. 1907, p. 806.

+ “ Nouveaux tenias d’Oiseaux,” Rev. Suisse Zool. xvi. 1908.

P. Z.S. 1912, p. 586, text-fig. 76.

MAMMALIAN CESTODES. 569

generative pores and at the middle of the segment ; each ovary

lies i in front of the corresponding vitelline gland. The vagina is

narrow and passes forward, running in front of and parallel to

terminal section of cirrus- pouch ; ‘poster iorly it dilates into a

receptaculum seminis lying close to and in front of ovary and

behind dilated region of cirrus-sac. Uterus as a simple cavity is

absent (2). The developing ova are lodged singly in cavities of

the medullary parenchyma, which they nearly fill, and are without

any lining epithelium. Eggs with very wide but delicate shell.

From this general résumé we may now endeavour to form a

generic definition of Diplop ylidium by eliminating from the above

characters those which are in all probability to be regarded as

only specific in value.

Diplopylidium, gen. nov.

Retractile muscular rostellum armed with two circles of hooks.

A double set of reproductive organs in each proglottid. Genital

canals pass between dorsal and ventral excretory tubes. Vagina

lying in front of cirrus-sac. Cirrus-sacs large and muscular, bent

upon themselves ; cirrus coiled, very slender, and unarmed ; sperm-

duct forms a coil. Testes numerous, filling wp available space in

progiottid. Ovaries two, in front of vitelline glands. Vagina

narrow, with receptaculum seminis. Uterus represented by nume-

rous cavities, each containing one egg only.

We have now to consider the affinities of this tapeworm and

the family within which it should be placed.

There is not a very large number of genera in which the

reproductive organs are double in each segment. We find, how-

ever, such forms in nearly all of the families into which the

Tetracotylean Cestodes are divided. We may at once place on

one side those genera, such as Cittotenia and Monieza, which

belong to the Anoplocephalide ; for in the genus which forms the

subject of the present communication there are not, as in these

forms, persistent uteri; and, moreover, the worms of the family

Anoplocephalide are not provided with a hooked rostellum. Nor

can we place the present genus, which I propose to term Dzplo-

pylidium, with either Diplophallus* or Diploposthe, two genera

which are nearly allied to each other, though placed by Ransom +

in separate families, viz. the Acoleide and Teeniade. It must be

noted, however, that Fuhrmann {, in his well-known revision of

the tapeworms of birds (those belonging to the Cyclophyllidea),

regards Diploposthe as not strictly referable to the Tzniade, but

as forming an intermediate type between this family and that of

the Acoleide.

* For anatomy of Diplophallus, see Cohn, Zeitschr. wiss. Zool. ]xvii. 1897, p. 277,

‘and Wolfhiigel, “‘ Beitr. z. Kenntniss d. Vogelhelminthen,” Inaug.-Diss. Freiburg-

im-Br. 1898.

+ Bull. U.S. Nat. Mus. 1909, no. 69.

t Zool. Jahrb., Suppl. vol. x. 1908.

570 DR. F. E. BEDDARD ON

Both of these genera have a rostellum which is armed with

ten hooks. The ‘preater number of hooks in Diplopylidiwm

might be regarded as a difference of merely specific value. But

while in Diploposthe * and Diplophallus the uterus is persistent,

and in mature segments occupies an enormous amount of space

in each ripe proglottid, Diplop, ylidium has probably not a per-

sistent uterus at all; or, if the spaces containing eggs which have

been described above are to be looked upoh as remnants of a

uterus or of uteri, the conditions are obviously very different.

This important diacronce: as it appears to me, renders any con-

fusion between these forms quite impossible. I have referred

above to certain other comparisons between my genus and the

two that have just been compared with it.

My genus presents certain points of likeness to the genus

Cotugnia, founded by Diamare in 18937. The chief point.

of likeness is that in Diplopylidium the ripe eggs are imbedded

singly in the medullary parenchyma, as is stated to be the case

with Cotugnia. But in the latter genus, Fuhrmann {, though he

gives no figures showing detail, speaks of a ‘“ parenchymkapsel ”

as surrounding the eggs. This is probably to be compared to

that of the allied Davainea, and, therefore, presumably is not

like that which I describe in the present paper. The vagina, too,

of Cotugnia has a dilated receptaculum seminis. Otherwise I do

not think that the two genera can be confused. The worms are

rather large species, and the rostellar hooks are numerous and

minute, and have the typical Davaineid form which is unlike

that which I find in Diplopylidium. But Cotugnia appears to:

differ from other Davaineids in having no hooks upon the suckers

[‘“‘ Ventose grandi e inerme” (Diamare)]|, and thus to approach

Diplopylidium, to which its double generative pores and possibly

the nature of the egg-cavities affine it.

It may be furthermore pointed out that the genus Co otugnics

seems to be characterised by its short and broad proglottids, in

which it clearly differs from the worm upon which I report im

the present communication. DiamareyT, in his paper upon

Cotugnia (and other tapeworms), assigns to his new genus

a species described by Monticelli§$ from material named by

von Siebold. This species (‘‘ Tenia bifaria”) has, as it appears

to me, rather more claims to be allied to my genus Diplopylidium

than to Cotugnia, if Diamare is right in supposing Monticelli to-

have overlooked the rostellar hooks. For in his figure || of the

generative organs (which is repeated in Bronn’s ‘ Thierreichs’ @ ),

* For the anatomy of Diploposthe, see Jacobi in Zool. Jahrb. x. Anat. Abth-

1897, p. 287; Kowalevsky, Bull. Ac. Cracow, 1903, p. 518, for a brief account of

Diploposthe sui-generis n. sp.(?); and Fuhrmann, Centralbl. Bakt. xl. p. 218, for

a general résumé of this genus.

+ Boll. Soc. Nat. Napoli (1) vii. 1893, p. 11.

t “Neue Davaineen,” Centralbl. f. Bakt. u. Paras. xlix. p. 115; and Zool. Anz.

xxiv. 1901, p. 273.

§ Boll. Soc. Nat. Napoli (1) v. 1891, p. 151. || Loe. cit. pl. vill. fig. 12.

¥ Pl. lvi. fig. 9.

MAMMALIAN CESTODES. 57}

Monticelli places the vagina in front of the cirrus-sac as I have

found it in Diplopylidium. But this tapeworm has a cirrus

which is armed with spines which I have not found, and which

also does not seem to occur in Cotugnia. Nor does Monticelli

say anything of the eggs: which makes any comparison more

difficult. The segments, however, are more elongated than in

Cotugnia and thus like those of Diplopylidiun.

Passing by the family Amabilide, of which the type-genus

Amabilia has double reproductive organs, and to which Diplo-

pylidium does not, as it appears to me, show any affinities, we

come to the Hymenolepide. To this family, and especially to the

subfamily Dipylidiine (in the sense of Ransom), Diplopylidiwm

shows points of likeness. In this subfamily there are two genera,

viz. Dipylidium and Pancerina, which have reduplicated gene-

rative organs. After the description of a Dipylidium, which I

have given in the present paper, I do not think it necessary to

emphasize further the distinctness of Dipylidiwm and Diplo-

pylidium. As to Pancerina, that genus has no hooks upon the

vostellum ; it may be merely an Oochoristica with double gene-

rative organs, or, as I have ouegesied elsewhere * an Ichthyo-

teniid. Our ‘knowledge of its anatomy is at present insufficient

to place it with accuracy.

But although Diplopylidiwm is certainly not identical with

Dipylid ium, it agrees with that genus and some other Dipylidiinze

in the characters of the ese holding spaces (which I do not term

uterus for reasons given ¢ above )r. It is obviously quite near in

this respect to the species of Dipylidiwm which I describe in the

present paper. Oochoristica also has the ripe ova scattered at

random through the medullary parenchyma ~. Monopylidium has

also the same “type of egg-holding cavities van I have referred

to more at length above in describing this part of the generative

system of Diplopylidium. Furthermore, in Monopylidiwm there

are species (e. g. Jf. macracanthum $) where the hooks are not

very numerous (22) and arranged in two circles. On the whole,

Tam inclined to place my genus in the near neighbourhood of

Dipylidium and Monopylidium.

P. Z.S. 1913, p. 6.

Supra, p. 567 ete.

Cf. e. g. Beddard, P. Z.S. 1911, p. 633, text-fig, 150 e.

Fuhrmann, Centralbl. f. Bakt. u. Paras. xlv.

tt %

572 MR. J. A. MILNE ON THE

37. Pacific Salmon: An Attempt to evolve something of

their History from an Hxamination of their Scales,

By Joun ApvAm Mitne, of Ardmiddle, Turriff,

Aberdeenshire *.

[Received April 9, 1918 ; Read May 6, 1913].

(Text-figures 95-118.)

SALMONIDA: SeRuctuRE, DevetorpmeEnt, EvHonoey.

The above title indicates what is perhaps a somewhat bold

venture on the part of one who lives on the east side of the

Atlantic, and who has only once seen a freshly killed Pacific

salmon during a short visit to Vancouver in 1893. The

reasons for my undertaking it are that, so far as I have been

able to ascertain, the study of salmon scales has not yet made

much progress in America, that undoubtedly much may be dis-

covered from them, and that even such inadequate observations

as I have been able to make may afford most valuable hints to

others better able to procure materials for the study of the habits

of the so-called salmon of the Pacific. I say so-called, because with

one exception, Salmo gairdneri, the Steelhead Trout, the Pacific

salmon do not belong to the same division of the genus Salmo as.

the salmon of the Atlantic Ocean. Excepting the Steelhead,

they belong to the subgenus Oncorhynchus, while our salmon and

trout belong to the subgenus Salmo.

I shall have to notice five species of Oncorhynchus which

breed in the rivers and streams of Western North America.

They are O. quinnat, or O. tschawytscha, the Quinnat, King, Tyee,

or Spring Salmon ; O. nerka, generally known as the Sockeye,

from the sunken appearance of the eyes, and also called the

Blueback and the Red Salmon ; O. kisutch, the Cohoe, Silver,

White, or Fall Salmon; O. gorbuscha, the Humpback, so called

from a peculiar hump which appears on the backs of the males

at spawning time; and O. keta, the Dog or Chum Salmon, A

sixth species, O. masu, is found on the Siberian coast and in

Japan, but I shall not deal with it here, as I intend to confine

my remarks to the salmon of the Pacific coast of North America.

It is there that full knowledge of the habits of the Pacific salmon

is of the greatest importance on account of the magnitude of the

canning industry, to which every one of the native species

now contributes its quota.

When the canning industry was first started in the West, the

Quinnat only was cared for, but soon the Sockeye was recognized

asa fish of much greater importance. The value of the remaining

species has only recently been appreciated.

It is generally believed that none of the fish of the genus Onco-

rhynchus that go to the rivers ever return and that all die after

* Communicated by the SECRETARY.

HISTORY OF THE PACIFIC SALMON. Diliey

spawning. But although the upper tributaries of the Fraser River

are almost unapproachable for some time after the spawning

season, on account of the numbers of dead and putrefying fish

which they contain, I do not think it is absolutely proved that

all the fish do die after spawning. Later on I shall produce

something approaching a proof that they do not. Most of them

undoubtedly do die then. Many of our Atlantic salmon are so

exhausted after spawning that they promptly die. Much more

so must that be the case with these Pacific salmon, which ascend

the rivers not for a few tens of miles as our fish do, but for some

hundreds, or even for many hundreds of miles. The probability

certainly is that when they go very far from the sea, none of them

return. But they do not all ascend to extreme distances; and 1

can see no reason why some of those which have not had far to

go, or great difficulties to surmount on the journey, should not

have sufficient strength to recover and to spawn again. I well

remember the manager of one of the largest canneries on the

Fraser saying to me that the idea that all the fish died was based

on the statements of Indians only; that no one else knew any-

thing about it (I speak of twenty years ago), and that millions of

kelts might come down the middle of the Fraser with the stream,

and not a soul be any the wiser. To my mind the fact that no

kelts are ever seen does not prove their absence. No netting is

going on when the kelts would come down, so, as no Pacific

salmon can jump, and as no Pacific salmon has ever been known

to take a bait in fresh water, it is most improbable that they

would be seen.

It was in the hope that some definite evidence might be forth-

coming upon this point that I first took up the study of the

scales of Pacific salmon. One certain spawning mark would

disprove the idea that every fish dies. But even on our Atlantic

Coast the percentage of fish that are recaptured after having

spawned is very small; and on the Pacific Coast the percentage

must, for the reasons mentioned, be much smaller. Supposing

it to be as high as one in a hundred, which it probably is not, it

might be necessary to examine the scales of some thousands of

fish before that one happened to be among them. I have not

been able to examine the scales of more than a few dozens, but I

think that I have been so lucky as to find a spawning mark

on the scales of a large Quinnat. One of its scales is shown

in text-fig. 95 (p. 574), and I believe that the mark about a

quarter of an inch from the edge of the photograph is a spawning

mark, I shall have more to say about it later on when I come to

deal with the Quinnat salmon in detail.

As against the general belief that Pacific salmon never survive

spawning, I have heard the argument that the largest Quinnats,

which weigh from 50 to 100 lbs., must be very old fish and that

they must almost certainly have spawned. In text-fig. 96 (p. 575)

is shown the scale of one which weighed 623 Ibs. It has certainly

not spawned, and it is also certainly not old. It appears to be

574 MR. J. A. MILNE ON THE

only in its fifth year but to have grown very regularly and fas

both in summer and winter throughout its life.

As I have to reason largely from analogy, I must now digress

for a time from my immediate subject to say a little about

what has been discovered from the scales of our salmon, Salmo

salar, which is also the salmon of the east side of the American

continent.

Scale of a Quinnat (Oncorhynchus tschawytscha) 33+ lbs. Length 415 ins.; girth

252 ins. Captured at New Westminster, B.C., in 1912. Supposed spawning

mark shown at S.

It appears to have occurred to Leuwenhoeck so long ago as

the year 1696, and to Réaumur in 1716, that the concentric lines

which are to be found upon the outer surface of the scales of most

fishes are formed with some relation to the age of the scale, and

therefore must give an indication of the age of the fish to which

HISTORY OF THE PACIFIC SALMON. 575

the scale belongs. But no very close or reliable investigations as

to how the ages of salmon might be read from their scales

were made until Mr. H. W. Johnston took up the study

a few years ago. Although I am not actually quoting from

anything he has written, I think his authority for every-

thing I am going to say, unless I may quote someone else, will

be found in one or other of his contributions to the Reports of

the Scottish Fishery Board. (See Parts II. of the Reports for

1904, 1906, and 1907.)

Text-fig. 96.

Scale of a Quinnat (O. tschawytscha) 623 lbs., 6. Length 502 ins.; girth 31 ins.

Captured at New Westminster, B.C., in 1912.

A salmon scale is divided into two areas, one, the anterior and

larger part, being enclosed in a pocket in the skin, to which it is

loosely attached; the other, the posterior area, being the only

part of the scale which we see while it is still attached to the

576 MR. J. A. MILNE ON THE

fish. The anterior part is covered by a mass of concentric lines °

or ridges, from the relative positions of which much may be

learned. ‘The posterior part is almost without lines. This

formation is common to the scales of all members of the

genera Salmo and Oncorhynchus. The whole scale is covered

by a membrane and grows with the fish. When the fish—and

with the fish, the scales—grows slowly the concentric lines, which

seem to be produced at a fairly uniform rate as to number, are

situated closely together ; when they grow quickly the lines are

placed further apart. When the fish does not grow at all, the

scales also cease to grow, and no lines are added to their surfaces.

The scales first appear upon young salmon as minute bony plates

under the skin. They may first be noticed when the fish is

about three months old, and about #? inch in length (Vogt,

‘Embryologie des Salmones,’ 1842, Klaatsch 1890, and Dahl,

‘Age and Growth of Salmon and Trout in Norway,’ 1912).

When about four months old and 1; imches long the little fish

may have from 2 to 5 rings round their scales ; and when about

eight months old, that is just before the winter comes on, there

may be, roughly speaking, from 8 to 20 lines already visible on

each of their scales. Within small limits the number of lines

varies in different scales even from the same fish. We know that

the yearlings when kept in ponds feed well and grow rapidly

during the summer months, and that as winter approaches they

take less nourishment, finally at times fasting completely for

three or four days on end. We also know that their growth

progresses in proportion. This mode of life is clearly depicted

on their scales. Surrounding the nucleus, the lines formed in

summer are at some small distance from each other, and each

line can usually be traced right round the scale. The lines

formed in winter are much closer together and are usually more

numerous on the anterior portion of the scale.

When the growth of the next year begins in the spring, the

lines first formed are again wider apart and generally continuous

all round the scale, and, later on, the winter formation of the

first year isrepeated. Thus there is formed an area of well-spaced

lines followed by a band composed of lines very close together.

This band is usually most noticeable in front, and is followed by

another area of open lines and asecond band which is probably not

uite so well defined as the first. The accompanying photograph

(text-fig. 97) of the central part of the scale of a Steelhead Trout

(S. gairdnert) well illustrates the formation. The line A points

to the completion of the first winter band, B of the second winter

band, © of the third, and the end of the line D shows the point

in the next year when the smolt left the river and commenced to

grow rapidly in the sea.

Tn the sea growth proceeds as before, but at a much increased

rate, and the difference between fast summer and slower winter

growth continues to be apparent on the scales.

Fish that are born, and spend all their lives in the sea, show

HISTORY OF THE PACIFIC SALMON. Bite

annual winter bands like the others ; but, except for these bands,

the spacing between the lines shows no sudden increase due to

any abrupt change in the mode of life. Compare text-fig. 98,

the photograph of a scale of a large haddock, with the other

photographs illustrating this paper, and the difference in the

Text-fig. 97.

tis)

: Aye a ge PS ;

ae. The amie a ae 4 ted atte ae is ee

Centre of scale of a Steelhead Trout (S. gairdneri) from the Fraser River, much

magnified, showing three winter bands before migration to the sea. (For references

see the text, p. 576.)

i. ed » nee a et

eae ry

: . Hi

centres will be at once apparent. I wish to emphasize

this difference because the Pacific salmon scales, if we except

those of O. keta, the Dog Salmon, show a sudden change

of growth just as do those of the true salmon; and this, to my

578 MR. J. A. MILNE ON THE

mind, at once disposes of the idea that these fish go down to the

sea almost as soon as they are hatched and commence feeding

only when they get there.

Text-fig. 98.

iis

neal

i :

i ei

ANE

aes

Scale of a large Haddock (Gadus eglefinus). XX about 40.

Mr. Johnston kindly gave me some of the scales taken from

salmon which had been marked as smolts in the estuary of the

Tay in the early summer of 1905. The smolts were then on

their way to the sea, and the marks were still attached to the

fish when subsequently recaptured as salmon. ‘Thus the exact

times spent by them in the sea were known, and it was found,

as had been expected, that the summer and winter bands on the

HISTORY OF THE PACIFIC SALMON. 579

scales agreed with the known times. Text-fig. 99 is a scale from

one of these fish which was captured in the Tay nets on May 7th,

1907. It had been marked as a smolt when the scale had grown

to the point C, and when the fish was just over two years old.

The winter band formed in the winter 1905-6 is apparent

between the points marked D, and the band formed in the winter

Text-fig. 99.

Scale of 124 Ib. salmon (Salmo salar) caught in the Tay nets on May 7th, 1907.

Previously marked as a smolt in May 1905. (For references see the text, supra.)

1906-7 between the points E. Outside the last band is seen

the quick growth made in the summer of 1907 previous to

May 7th, when the fish was netted. Summer growth is the term

used in distinction to winter growth. It does not mean growth

taking place only in June, July, and August, our summer months.

Tt evidently starts and finishes at varying times, the probable

Proc. Zoou. Soc.—1913, No. XX XIX. 39

580 MR. J. A. MILNE ON THE

limits being about the middle of March and the middle or end

of September.

The foregoing remarks may suffice to explain the regular

markings on salmon scales, and how they show the age and time

of migration of the salmon. Scales, however, may show much

more.

Mr. Johnston noticed that the scales of the older fish some-

times seem to have stopped growing, and become jagged and

broken at the edges, and then to have gone on growing again.

When this has happened the lnes of the new growth do not

exactly follow the contours of the lines of the old growth, and

thus a clear mark is formed round the anterior portion of the

scale, often also apparent as a thickening round the posterior,

the unlined, portion. Further, it was noticed that when such a

mark occurred the number of lines between the winter bands on

each side of it was frequently far from normal. ~The normal

number differs somewhat in different fish, and also within certain

limits on scales from different parts of the same fish; but never-

theless it is fairly constant, and any wide departure from the

usual state of things required explanation. A reason for the

mark suggested itself to My. Johnston from the known fact that

salmon, whether or not they take food in fresh water, do not

take it in sufficient quantity, after the smolt stage is passed, to

nourish them. Therefore neither salmon nor their scales can

grow in fresh water. It was further noticed that most scales

taken from kelts---that is to say from salmon which, having

spawned, have not yet returned to the sea—were broken and

torn at the edges. From these facts Mr. Johnston argued that.

the sort of marks illustrated at C and D on the accompanying

photographs (text-figs. 100 & 101) showed that the fish bearing

them on their scales had entered fresh water and stopped growing

there; that they had spawned and become shrunken after

spawning, so that, from mechanical reasons, the scales being im-

bricated, 7.¢e. overlapping each other like tiles on a roof, had become

frayed at the edges; and, lastly, that on the salmon’s return to

the sea the new growth had started again evenly round the scale

and thus left the mark. Spawning operations mean a winter

at least spent in fresh water and, therefore, if the fish enters

a rviver early in the year there will be fewer than the normal

number of lines between the winter bands formed on each side

of the spawning mark. If it comes in early in the spring there

will be none in that year, and in the year following there will

be fewer than usual on account of the fish taking some time

after its return to the sea to make up lost condition before it

starts again to imerease in size. Of course, the later the

salmon came into the river the more lines would have been

formed behind the mark, and the later it returned to the sea the

fewer would there be in front of it.

Besides the irregular arrangement of the lines on the scales

when a fish has spawned, there is a thickening round what was

HISTORY OF THE PACIFIC SALMON, 581

the edge of the scales at the time the salmon left the sea, and

this thickening is apparent both on the anterior and posterior

parts of the scales if the edges have not become too much

worn during the stay in fresh water. JI account for it by

supposing that the materials which go to form the scales

are still being secreted in fresh water, and that as the skin

Text-fig. 100,

Scale of 33 lb. Salmon (Salmo salar) caught in the Tay in August 1903.

This fish had spawned twice, and the marks are shown at C and D.

pockets have ceased to grow they can be deposited only on the

scale edges already formed. The cells from which the concentric

lines on the upper surface are evolved are already dead, except

close to the periphery of the scale (Klaatsch, ‘ Zur Morphologie

der Fischschuppen,’ 1890, and Stuart Thompson, ‘ Journal of the

Marine Biological Assoe.’ vol. vii. no. 1, 1904). This in itself

3o™

582 MR. J. A. MILNE ON THE

would account for the marginal thickening if the cells still remain

active in fresh water and the scale cannot expand,

Text-fig. 101.

eee

Wee

A, 7.

Hix

GALT

Scale of a 334 lb. Salmon (Salmo salar) caught at Port Gordon, Aberdenshire, on

August 19th, 1908. Previously marked as a kelt on the Deveron, March 23rd,

1908. This fish had also spawned in the previous year, and the two marks

are shown at Cand D. The original scale has been lost at the end of the first

summer in the sea, and its position filled by a new scale without lines.

On both text-figs. 100 & 101 two spawning marks are to be

seen, but the figures illustrate what I have just said as well as if

they had but one mark. Scales which like these show two marks

are far from common. The central part of text-fig. 101 is

without lines, so that the parr scale does not show at all. ‘This

is because the original scale was lost by the fish towards the end

of its first summer in the sea, and the place occupied by it in the.

skin pocket was filled by a new scale which, the line producing

HISTORY OF THE PACIFIC SALMON. 583

cells having died, is devoid of concentric lines. In both text-

figs. 100 & 101 the first winter band formed in the sea is shown

between the ends of the lines AA. The second winter band is

between the lines BB. The first spawning mark in each case

is at the end of the line marked C and the second spawning mark

at D. Text-fig. 100 is a scale from a 33 ]b. salmon caught in

the Tay in August 1903. It can be seen, from the few widely

spaced lines between the band BB and the spawning mark C,

that when it first spawned it was a summer fish about 43 years

of age. After spawning the scale had become much worn, so

much so that, except in ” front, not only the summer growth but

the band BB had disappeared, and the new growth formed after

the fish returned to the sea can easily be distinguished, its lines

being again continuous round the scale. In foxt- fis. 101 the

winter band of the 5th year had begun to form before the

spawning mark C, so it was evidently a late autumn salmon.

In text-fig. 100 the second spawning mark is indicated by the

line D. The fish must have returned to the sea after the first

spawning early in the year, and the lines between the two

spawning marks show it to have come in to spawn the second

time late in the summer of the same year. The following summer

it was coming back to spawn a third time when it was captur ed,

The thickening on the posterior part of this scale, due to both

spawning periods, is clearly visible in the form of dark lines.

In text-fig. 101 the second mark D, like the first mark C, denotes

an autumn fish. It is not a very clear spawning mark, but I

have chosen it for that very reason, because it is so like the mark

on the scale of the Quinnat which I believe to have spawned

(see text-fig. 95, p. 574) and because I have undoubted evidence

that mark D on text-fig. 101 is a true spawning mark. The fish

was caught as a kelt, weighing 222 lbs., and marked by my own

samekeeper, Peter Bowie, at “Netherdale on the River Deveron,

on March 23rd, 1908. Tale put in its fin one of the Seotbish

Fishery Board labels numbered 3613 B. The scales then showed

only the first spawning mark C. The fish was recaptured as a

clean salmon, weighing 333 lbs., in a bag-net at Port Gordon on

the Aberdeenshire coast on August 19th, 1908, after only five

months’ interval. The mark D thus certainly represents the

second spawning period during which my keeper caught and

marked the fish.

Many other instances have occurred in which fish that have

been marked with distinctive numbers as kelts have been caught

again, and in every case they have proved the truth of Mr. John-

ston’s theory by showing a spawning mark on their scales. On

the other hand, no fish that one knows from its age could not

have spawned has ever been found to bear quite the same sort

of mark, although various marks indicating sudden checks in

feeding may appear.

I have entered somewhat fully into the above description of

spawning marks, because I hope that now many scales of Pacific

584 MR. J. A. MILNE ON THE

salmon will be examined upon the chance of finding one. Much

can be done with an ordinary high-powered pocket- -lens, and for

close observation only a very lor powered microscope is either

necessary or desirable. I find a 14-inch objective powerful

enough for all purposes. One wants to be able to see as much as

possible of the scale at the same moment.

Mr. Knut Dahl (‘ Age and Growth of Salmon and Trout in

Norway, London, 1912) has shown that the size of a fish at

any period of its life may be deduced from its scales.

A fish does not change its seales. From the time they form

they are retained through life unless removed accidentally. If a

fish does by chance lose one it is replaced by another of the same

size and shape but without the concentric lines—see text-fig. 101.

It follows that as the fish grows, but remains covered by the same

number of scales occupying the same relative positions, the scales

must grow with the fish, and the growth of each scale be pro-

portionate to the growth of the fish. If, then, the protected

part of the scale be measured along its length from the centre to

the anterior edge and again from the centre (by which I mean

the nucleus) to, say, the first winter band, the lengths of the

scale at these two points will be proportionate to the lengths of

the fish at corresponding ages. An example will make this more

clear. Suppose we have a salmon 75 cm. long and that one of

its scales, when magnified, measures 55 mm. from the centre to

the anterior edge, and 11 mm. from the centre to the point at

which the rapid growth, consequent on migration to salt water,

is seen to begin. Then as 55 is to 11 so is 75 cm. to the length

of the fish when it entered the sea. The smolt was therefore

15 cm. (or just under 6 inches) long. Another scale from the

same fish magnified to the same degree, might be only 45 mm.

long, but then the other measure would be found to be 9, and the

sum would work out just the same.

Dahl has proved his theory by measuring the scales of

hundreds of salmon and trout from different rivers. He has

always found that the actual average lengths of the fish in

different rivers at various ages, agree almost exactly with the

lengths calculated by him from the scales of the older fish.

I have been trying to apply this method of measurement to

the scales of Pacific salmon, and I will give the results when I

come to deal with each species separately. First, however,

I must make some criticisms on what I have set out above as

my understanding of the claims made by Dahl. I tried to

check his theory on the scales of two salmon which had been

marked as kelts and subsequently captured as clean salmon.

Photographs of both of these scales form illustrations to

Mr. Johnston’s second paper on salmon scales in the 25th

Annual Report of the Fishery Board for Scotland. The lengths,

on the occasion of each capture, are given elsewhere in the

24th and 25th Reports, so it is known that the salmon marked

No. 9194 was 27 inches long when taken as a kelt, and 31 inches

HISTORY OF THE PACIFIC SALMON. 585

long when recaptured. The other fish, marked 1180, was

264 inches long as a kelt and 36 inches long when recaptured.

The measurements on the photographs of the scales from the

centre to the anterior edge, and from the centre to the spawning

mark are 130 and 116 for No. 9194, and 198 and 175 for No. 1180,

the unit of measurement being ~;th inch. This would give the

kelt measures as 272 inches and 32 inches respectively. The

former measure is only 2 inch wrong, but the latter is nearly

6 inches wrong, and shows either that the scale is abnormal or

that Dahl’s system of measurement is not applicable to a fish

that has spawned.

Another criticism is that measurements from several different

scales of the same fish seldom all agree exactly, and I have there-

fore come to the conclusion that it is very unsafe to rely on the

measurement of one scale in estimating the size of the fish at

various ages. The reason may be that it is not easy to recognize

either the exact centre of growth, or the exact limits of the

Text-fig. 102.

Quinnat (O. tschawytscha). 25 bs. 13th October, 1911. Shuswap,

South Thomson River.

The black lines show the variations in the long axis of the scale.

various bands, but Ido not think that the scales grow quite

equally. Still the idea seems so well founded in theory, and to

have worked out so exactly in practice on a large scale in Norway,

that I believe I may consider myself justified in drawing con-

clusions from the average measurements of a considerable number

of scales taken from the same fish, and whenever possible, I have

measured 20 scales. A smaller number might suftice in the case

of a true salmon, but the scales of the Pacific salmon are more

difficult to read. As a rule, the limits of the bands are less

586 MR. J. A. MILNE ON THE

clearly defined, and the change of growth, which I can only

suppose to be due, as in the case of the true salmon, to the

commencement of sea life, is more gradual and not so well marked.

In addition to this, the long axis of the scale frequently shows

more than one change of direction, the scale apparently being

liable to get turned round in the skin-pocket. The accompanying

photograph (text-fig. 102) will show my meaning. I have not

measured such scales as these.

To come now to what is known, and to what I think I may claim

to have found out about the various species of Pacific Salmon.

THE SockEYE. (Oncorhynchus nerka.)

First I take the Sockeye, because it is the mainstay of the

canning industry of the West. The number of Sockeyes canned

on the Pacific coast in 1909 must have attained to the enormous

total of about 50 millions. The exact weight was214,980,448 lbs.,

or nearly 100,000 tons.

Text-fig. 103.

Sockeyes (Oncorhynchus nerka) running up Scotch Creek, a tributary of the Fraser

River, British Columbia. (Photograph reproduced by kind permission of

My. Frank Parry.)

The Sockeye is not much in evidence south of the Columbia

River, in which it is known as the Blueback, but is plentiful from

there the whole way north to Bering Sea. In Alaska it is known

as the Red Salmon. The chief Sockeye stream is the Fraser

River, which these fish ascend to spawn in countless myriads.

The feeding-ground of the Sockeyes is somewhere far out in the

Pacific, and the fish seem to cease feeding before they approach

the coast, for even when caught in the sea near the coast their

HISTORY OF THE PACIFIC SALMON. 587

stomachs are invariably empty. This, however, may be from

lack of means rather than from want of will. When one hears

of a shoal of fish, seven miles broad and of unknown length,

heading for the land through the Straits of San Juan, one can

well imagine difficulties in the commissariat. Part only of this

big shoal enters the Fraser River, the remainder moves on up the

coast of British Columbia. The photograph (text-fig. 103), which

I give by kind permission of Mr. Parry, late of Granite Creek

Hatchery, shows a detachment on its way up Scotch Creek, a

subtributary of the Fraser River, about 300 miles from the sea.

The big run of which I have been speaking strikes the south-

west coast of Vancouver Island in J uly and August, coming from

the north-west, but a few Sockeyes run as early as April. In

the Fraser River itself the main run is in August, and some con-

tinue to come in until October. In the far north of Alaska the

main run is as early as June, which goes some way towards

showing that the feeding-grounds are in the north, unless a

natural instinct to get spawning over before the winter sets in has

determined the habits of the fish fr equenting the most northerly

rivers. In the Fraser district spawning begins in August, and

may go on until November; spawning takes place only in streams

running into or out of lakes. In this district, from which

most of my information and all my specimens have come, several

hatcheries have been established, which in 1910 liberated

134,639,200 Sockeyesin British Columbian waters. Besides these

4,544,825 were liberated from U.S. hatcheries in Puget Sound,

and 257,021,790 in Alaska.

Tam indebted to Mr. W. J. Sim, who was employed at Granite

Creek for some years, for the information that the fry are liberated

from the Canadian hatcheries as soon as the yolk-sac has been

absorbed, at a period of the year which varies from January to

April, according to the date of spawning. The main liberation is

about the first week in February. In that district the fry may

remain in the creek for two months or less; then they move into

Shuswap Lake. The exodus from the lake takes place from June

to September. The fry are supposed to be under one year old at

the time of their exodus from Lake Shuswap, but no one really

knows how long they may have remained in this, or in any other

lake, and it is admitted by some observers that they may be in

their second year. Dr. Greene, writing in the Bulletin of the U.S.

Bureau of Fisheries, vol. xxix. 1909, on the Migration of Salmon

in the Columbia River, and quoting Evermann, says the fry

begin their seaward journey not sooner than September of their

first, and not later than July of their second year. That means,

I presume, that they leave the lakes when from nine to eighteen

months old. But others, e. g. Rutter, Bulletin U.S. Bureau of

Fisheries, vol. xxii. 1902, p. 102, say that they begin to descend the

rivers as soon as they can swim, and reach the sea in about three

months. My. Sim says he is ori that there are no fry in the

rivers after September, and that they could not possibly avoid

588 MR. J, A. MILNE ON THE

being swept down with the current as soon as they leave the

lake.

The question as to the age at which the fry enter the sea is

one which scale-readings should easily settle. So far, I have been

Text-fig. 104.

ALSk.

Sari

Scale of Sockeye (O. nerka), $, 35 lbs. Length 202 inches; girth 103 inches.

A=end of first year’s growth. B=nmigration to sea.

C=end of second winter, D=end of third winter.

unsuccessful in my attempts to procure any specimens of fry of

known ages for the purpose of seeing how their scales do actually

grow, but I give some illustrations of Sockeye scales from full-

grown fish (text-figs. 104-106). I have no doubt but that each

HISTORY OF THE PACIFIC SALMON. 589

one of these Sockeyes spent a year at least in fresh water, and

the first of them pr obably not less than 15 months. I can

conceive of no other way in which scales with centres similar to

these could have been formed. They are the only Sockeye scales

which I have from measured fish, but I have others from six fish

of which I know only the weights, and a great many more from

Sockeyes that unfortunately were neither weighed nor measured.

All have similar centres.

ary

ni a Py A

Scale of Sockeye (O. nerka) 2. 5; lbs. Length 24+ inches; girth 121 inches.

A=end of first winter. B=migration to sea.

C=end of second winter. D=end of third winter.

The Sockeye from which the scale, shown in text-fig. 104, was

removed, was caught last August at New Westminster, near the

mouth of the Fraser River. It weighed 3} lbs. and measured

202 inches in length and 104 renee in birth when captured.

I have measured 20) scales from this fish, the aver age length of

the enlarged images to which [applied the measure being 33 mm.

from the centre to the anterior edge. The other average

measurements to the points denoted by the lines drawn on the

590 MR. J. A. MILNE ON THE

figure were 5 mm., 6°5 mm., 10 mm., and 20 mm., and the lengths

of the fish at the various ages indicated work out as follows :—

At the end of the first year 2? inches; when it entered the sea

34 inches, at the end of the second year 6 inches, at the end of

the third year 124 inches, and when caught it was as stated

202 inches. The points indicated by the lines drawn on text-

fig. 104 are calculated from the averages of the 20 scales, and

they appear to coincide absolutely with the points I should have

marked from examination of this scale alone.

Text-fig. 106.

aie ice a A

Scale of Sockeye (O. nerka), $.7 lbs.. 26% inches long; girth 144 inches.

e 5 ,

A=migration to sea at end of Ist year. Bend of 2nd year. C=end of 3rd year.

The next scale, text-fig. 105, is from a Sockeye caught at the

same place and time. In the illustration the parr or fingerling

scale is not very clear, but the examination of 20 scales enables

me to put the lengths of the fish at the various points and times

indicated by the lines drawn on the picture as follows :—1st year

2% inches, entered the sea during the second year when 3 inches

long, was already 9 inches long at the end of the 2nd year,

193 inches long at the end of the 3rd year, and, as was known,

HISTORY OF THE PAGIFIC SALMON. 591

243 inches at the time of capture near the end of the fourth

summer,

The scale of a Sockeye, also taken at New Westminster in

August, is shown in text-fig. 106. In the case of this fish, the

entry into the sea seems to have coincided with the end of the

first year’s growth. That is to say, it was some time during the

winter ee 10. The readings from 20 scales show it to have

been then 2? inches long. At the end of its second year it was

113 inches long, at the end of its third year 217 inches, and when

coq ed 264 inches.

The great difference in growth of the three fish in the second

year 1s very noticeable, but it seems quite natural, and goes far

to confirm the accuracy of my reading of the parr scale, when it

is observed that the fish which appears to have spent the whole

of its second year in the sea has grown most, and that the one

which spent the longest part of it in fresh water has made the

least progress.

It is further to be noticed that these fish all returned to spawn

when of the same age, namely at the end of their fourth year. I

have the scales of nine Sockeyes from the Fraser River. They

varied in weight from 37 to 84 lbs., but all are in their fourth

year. In British Co imeealaken the Sockeyes weigh from 3 to 10 lbs.

Fish up to 17 lbs. have been caught, but over 10 lbs, they are very

rare. I therefore seem to have got specimens of all the average

weights. That they are all of the same age may be an accident,

but much more probably it is not, and if it is not an accident it

is a fact of the utmost importance to the Fraser River District.

In the Fraser River the run of Sockeyes in every fourth year,

the year after leap year, is almost six times as large as in any of

the intervening periods of three years, and this has been the case

as long as records go back. Whether the fourth years have shot

ahead of the others, or whether all years were once equal to them

will never be known, but most likely adverse breeding seasons in

the intermediate years have gradually diminished the stock. If

it proves upon further examination of scales that every Sockeye

returns to breed in its fourth year im this district, it might well

be centuries before the breeding stock, once diminished, increased

to its original numbers. British Columbia has now only one

season out of four up to what might be the mark; and, if I am

right, the inference is obvious that the efforts of the hatcheries

should be devoted chiefly to the collection of ova in the lean years

even if they have to import them from other districts. In the

springs of 1906 and 1910, the years following the last two bi

years, 100,479,000 and 105,312,500 Sockeyes were liberated from

British Columbian hatcheries. In the intermediate years the

numbers were only 36,965,900; 51,855,200; and 41,909,500.

My contention is, that if it is humanly possible the numbers

hatched out in the seasons following the lean years should be

made even greater than after the fat years. Some people may

say, “ But how do we know that if we turn out fry in the Fraser

592 MR. J. A. MILNE ON THE

River or its tributaries, they will return there when they grow

up?” The very fact that it is in that district alone that three

years out of every four are lean years shows that the fish return

to their own river. If it were not so, the shortage in these years

would have gradually spread itself over a wider area. It seems all

the more wonderful that it has not done so, when it is remembered

that the big shoal which comes in in the summer is by no means

solely composed of fish making for the Fraser. The facts further

appear to show that not only do the fish return to the Fraser,

Text-fig. 107.

Scale of Sockeye (O. nerka) removed after spawning.

but to the very creek in which they were hatched, for how

otherwise can one account for many of the creeks which are full

of spawning fish in the big years being always absolutely un-

tenanted in the years between them. Salmon marking in our

own country has now shown almost conclusively that when the

access to a river is unimpeded the salmon born in it invariably

return to it. The fish that have been marked in one river and

subsequently recaptured in another have all been-from some

HISTORY OF THE PA@IFIC SALMON. 593

comparatively small stream, or from some river with a bad mouth

which they might well find difficulty in entering again when

they wished to do so.

The value of hatcheries is a much-disputed point on which I

will not express any opinion. For the moment I am content

to assume that they are of some value on the Fraser, and that

being so, to point out how they can best be utilized. But I will

say that if the falling off in the catches which has been apparent

since about 1900, when each year is compared with the fourth year

before it, proves to be more than a mere temporary fluctuation,

hatcheries alone will certainly not remedy the evil, and it will

be necessary for a time at any rate to restrict the number of

fish that are permitted to be caught.

Text-fig. 108.

Piece of skin taken from a Sockeye (O. nerka) after spawning.

I should mention that I have scales from Sockeyes in their

fifth year, but they came from the State of Washington, not

from the Fraser.

In text-fig. 107 is shown the scale of a Sockeye which weighed

6; lbs. and was caught at Morris Creek near Lake Shuswap,

about 300 miles from the coast. So far as I can judge it is quite

typical, and its interest is in showing, from the worn condition

of its edge, that if a Sockeye ever returned to the sea after

spawning and was caught again, a clear spawning mark should be

apparent on its scales. The scales le so far apart, and are so

594. MR. J. A. MILNE ON THE

flexible, that unless one saw how much they may become worn

one would hardly believe it possible.

It will, J think, be news to most of those employed on the

hatcheries that the spawning Sockeye possesses scales at all.

Last year when I asked for scales from a hatchery, I was told

that it was well known that the Sockeyes absorbed all their

scales as food in the course of their run up the river and never

had any when they reached the spawning beds. IJ argued the

point, and the hatchery people ultimately sent me the piece of

skin shown in text-fig. 108 taken from a Sockeye that had

spawned, to convince me that I was wrong. The scales were

there, but are by no means so clear in reality as they are in

the photograph. They are very deeply imbedded in the skin,

and, in order to remove some of them for examination, it was

necessary first to soak it well and then to pull and stretch it so as

to open the skin-pockets. After that the scales were still invisible

and adhered closely to the skin which formed the top of the

pockets. It was possible, however, to remove them with a pair

of foreeps without much further difficulty. So even those who

have handled spawning Sockeyes for years may be excused for

thinking them to be without scales.—How many fishermen in

this country know that eels are covered with scales @

THe Quiynat (Oncorhynchus tschawytscha).

The known range of the Quinnat on the American Coast is from

the Ventura River in California to Norton Sound, Alaska. It

probably really extends into Aretic regions. ‘The Quinnat is also

known as the Chinook Salmon, the King Salmon, the Tyee, the

Red or the White Spring Salmon, and the Black Salmon. The

flesh is generally of a deep salmon-red colour, but in the south of

Alaska and down to Puget Sound, sometimes as many as one-

third of these fish have white flesh. Sometimes one half of the

body is red and the other white, and sometimes the flesh is

mottled. The white-fleshed fish are of little use. All the scales

sent me and marked as from either Red or White Spring Salmon

are from fish which have spent three winters, and a considerable

part of the feeding season following the last winter, in the sea.

Analogy from the true salmon would lead one to suppose that

the white and mottled fleshed Quinnats might be fish that had

spawned, but I can find no trace of a spawning mark on any of

their scales. It has been suggested that the Red and White

Spring Salmon are distinct varieties and that the specimens

with mottled flesh are crosses, but upon this point I can offer no

opinion. Differences in food might account for much, as in the

case of the Brown Trout (Salmo fario).

The name Black Salmon arises from the colour assumed by the

Quinnat at spawning time.

In 1909 the weight of Quinnats canned on the Pacific Coast

was 12,640,344 Ibs.

HISTORY OF THE PACIFIC SALMON, 595

As a rule Quinnats weigh from 18 to 30 lbs. in British

Columbia, but much larger specimens are taken, and in Alaskan

waters they have been caught over 100 lbs. in weight, and average

23 lbs. in some seasons.

Quinnats are very powerful swimmers and ascend fast-running

streams in preference to others, making for the head waters, and

sometimes running up for many hundreds of miles.

Text-fig. 109.

Seale of Quinnat (Oncorhynchus tschawytscha). 15 lbs. Length 314 inches ;

girth 193 inches. New Westminster, August 1912. (For references see text, p. 596.)

Most of the principal rivers have spring and autumn runs,

and some of them a summer run also, so Quinnats may be said

to run from January until October.

The scales I have examined show the running Quinnats to be

in their fourth and fifth years, those over about 30 lbs. in weight

being a year older than the others. Numbers of them are

Proc. Zoou. Soc.—1913, No. XL. 40

596 MR. J. A. MILNE ON THE

spawned in the hatcheries, from which it was estimated that

90,740,472 were liberated in 1910.

I have already shown photographs from scales of Quinnats

which eee 333 lbs. and 623 lbs. (see text-figs. 95 and 96,

pp. 574 & 575) and 1 now append another’ (text-fig. 109) from a

fish which weighed 15 lbs. The parr or fingerling ; stage of each

scale shows that the fish spent a year (A) and the greater part of

a second year (B) in fresh water.

I have examined scales from eighteen Quinnats varying in

weight from 10 to 623 lbs., and all show about the same duration

of the freshwater stage.

When the little fish enter the sea they seem to be already

larger than the Sockeyes, and ae measure from 4 to 6 inches in

length. The lengths of the 623 Ib. fish at various ages, deduced

from the qneAS SMAI: of olen seales, were as follows :—At the

end of the first year 22 inches, when it entered the sea in its

second year 44 inches, at the end of the second year 133 inches,

at the end of the third year 25 inches, at the end of the fourth

year 39 inches, and when captured it measured 503 inches. The

scale illustrated (text-fig. 96) agrees exactly with these averages

up to the end of the third year, but the position of the fourth

winter band on it would show the fish to have been about

40 inches long at that time instead of 39 inches, if that scale

alone were to be depended upon.

The lengths of the 15 1b. fish,as shown from 22 scales, were, at

the end of its first year 34 inches, when it entered the sea

5 inches, at the end of its second year 13 ; inches, at the end of its

third year 26 inches, and when captured it measured 312 inches.

Text-fig. 109 shows the points from which the measurements were

taken, the scale being exactly an average one.

The measurements of the 337 Ib. fish, averaged from 16 scales,

were, at one year 37 inches, when it entered alae sea 43 inches, at

the end of the second year 127 inches, at 3 years 213 inches, at

4 years 31 inches, and when captured 412 } inches.

The above seem to be about the average lengths of Quinnats at

the ages given. None of the twelve specimens I have measured

depart far from them.*

The Spawning Mark.

On the scale of the 33} lb. Quinnat (text-fig. 95, p. 574),

check in growth (S) may be observed beyond the fourth winter

band (KE). It is equally clear on every scale of this fish, and

IT think it must be a spawning mark. One might expect

* Note:—Since this paper was written I have ascertained that in the Natural

History Museum, South Kensington, there are specimens of Quinnat parr, about

4 inches in leneth, which came from a lake near the head weters of the Fraser River.

I have had the priv elege of examining the scales of one of these specimens which

measured about 44 inches. The first winter band is quite clear, and five lines

formed in the second summer are apparent.

HISTORY OF THE PACIFIC SALMON, 597

a better defined scar, but Quinnat scales are very thin and

flexible, and besides are well protected by a great thickening of

the outer skin at spawning time, so they may never become much

worn at the edges. ‘That they may not have become much worn

up to the actual time of spawning is shown by text-fig. 110, a

photograph of a scale removed from a 12 1b. Quinnat after it had

spawned in the South Thomson River, about 300 miles from the

sea. That scale not only sustained the wear from the shrinkage

of the fish owing to lost condition between the time at which it

left off feeding, which the lines following the last winter band

Text-fig. 110.

1g” naar . WAX

Liat pak

Midis GC 2)

SP 4

<x heey

SS e

A rt Sl

Scale of 12 1b. Quinnat (O. tschawytscha) from South Thomson River.

Fish speared after spawning, 13th October, 1911.

show to have been quite early in the year, until it was speared

for me under the supervision of Mr. John Brown of Celista, B.C.,

on the night of 13th October, 1911; but the skin was sent to me

dried and folded, and I removed the scales from it myself. As this

scale shows so little sign of wear at the edge, it seems to me quite

conceivable that the mark S on text-fig. 95 is a spawning mark.

Tf it is not, Tam unable to suggest any other explanation of it.

Compare text-fig. 95 with text-fig. 109. These scales were

taken from fish caught at the same place at the same time. There

are more lines after the last winter ring on the former than on

40%

598 MR. J. A. MILNE ON THE

the latter, and that in spite of what is a very obvious check in

growth. ‘The check clearly lasted long enough for the edges of the

scales to become somewhat worn, for the lines following it do not

at all points exactly follow the contours of those immediately

preceding them. ‘This fish cannot in these circumstances have

grown faster than its fellow, which was feeding all the time,

and yet there are 20 lines as against 12. If, however, it

spawned it added the twelve lines between the winter band and

the scar in the early part of the year 1911, and the eight lines

outside the scar in 1912. It would have spent the early part of

1911 in the sea, then come in to fresh water to spawn, and

remained there after spawning for some little time. It would

then have returned to the sea in poor condition and possibly not

very early in 1912, so that by the time it had made up condition

and started to grow again it would add but another 8 lines or so

to its scales before the spawning instinct again induced it to seek

fresh water, and thus brought it to the place of its capture.

Text-fig. 111.

Quinnat Skin

10 lbs. Nat. Size

Piece of skin taken from a Quinnat (O. tschawytscha) after spawning.

Compare this mark also with the outer spawning mark on

text-fig. 101, and note how like they are.

A Cutemet of about 30 lbs. is just the size on which I think a

spawning mark is most likely to be found. The larger fish, like

the 623 pounder, a scale of which is shown in text- fig. 96, Tere

most probably attained their size on account of ane feeding

HISTORY OF THE PACIFIC SALMON, 599

instinct being more pronounced than the sexual. Scale reading,

so far as it has gone, has shown that our own very large salmon

are all maiden fish.

Text-fig. 111 shows a piece of Quinnat skin taken from a fish

that had lately spawned.

Tur Conor (Oncorhynchus kisutch).

The salmon with which J shall next deal is the Cohoe, also

known as the Silver Salmon and the Fall Salmon. The photo-

graph of its skin (text-fig. 112) shows that the scales overlap

much more than those of either of the species already described.

A spawning mark should therefore be better defined if one were

to be found.

Text-fig. 112.

Cohoe Salmon |

Piece of skin of the Cohoe (Oncorhynchus kisutch).

The Cohoe Salmon gets the name of ‘ silver” salmon from its

appearance when it first comes in from the sea. It is then a

most brilliant silver with a greenish tint on the back. As

spawning time approaches it becomes a dirty red. The Cohoe is

found in almost all the Pacific Coast streams from Monterey Bay

northwards. The run in the Fraser River is in September and

October, but the Cohoes are on the coast from July to November.

The weight of Cohoes canned in 1909 was 17,789,890 lbs., but

large numbers, 1,152,452 lbs. in United States waters alone, were

dealt with in other ways. The Cohoes in British Columbia

- weigh usually from 3 to 8 lbs., but larger specimens are by no

600 MR. J. A. MILNE ON THE

means uncommon and may weigh as much as 30 lbs. More

Cohoes would probably be canned if the run did not occur so late

in the year, at a time when most of the canning stations are

closed for the winter.

The number liberated from the hatcheries in 1910 was

50,424,586, but in some years the numbers have been much

greater.

Text-fig. 113.

Scale of Cohoe (O. kisutch), 2. 4 lbs. 60z. 17th November, 1911.

I find the scales of the Cohoes the most difficult of all to read.

I have specimens of scales from eighteen fish from Puget Sound

and from Shuswap, on the Fraser River. These fish varied in

weight from 3 to 18 lbs. What strikes one as most curious is

that, with the possible exception of one fish of 5 Ibs. 6 oz.

(text-fig. 113) in which a few close lines between B and C, which

I think due to a slight check in feeding, may really be the second

winter band, their scales show them all to have been of the same

age, namely nearing the end of their fourth year.

I am satisfied that the Cohoes spend the whole of their first

year in fresh water. At the end of it they measure, according

to their scales, from 23 to 3 inches. One rapid grower, which

weighed 15 lbs. near the end of its fourth year, measured as

much as 4 inches at the time of migration. The growth in the

HISTORY OF THE PACIFIC SALMON. 601

second year is very difficult to make out. I feel convinced that

the greater part of it (I mean as to time) has taken place in fresh

water in nearly every instance, and in no instance am I quite

convinced that the whole of it did not doso. If the whole of the

second year’s growth took place in fresh water, the fish are a year

older than I have stated above. In all cases (I speak, of course,

only of the fish I have examined) the first entry into the sea

appears to have taken place between 18 months and 2 years

after hatching. Further certainty is added to this statement

by the measurements of the scales, which show the second year’s

growth to have been comparatively small. In three instances in

which, from the arrangement of the rings, I think the fingerlings

entered the sex when about 18 months old, the little fish were

three times as long at the end of their second year as at the end

of their first. None of the others even quite doubled their

length. That is what one would expect at that stage from fresh-

water feeding, but not from seafood. In the third year growth is

very rapid, and hardly diminishes even as winter comes on.

Text-fig. 114.

Scale of Cohoe (O. kisutch), g. 14 1bs. Puget Sound, 18th October, 1911.

Text-figs. 114 and 115 are typical Cohoe scales.

All the scales from Puget Sound are from fish captured on

18th October, 1911, and all show at their edges the lines coming

clese together to form the next winter band, see text-fig. 114.

A similar formation may be noted on text-fig. 113, which reached

602 MR. J. A. MILNE ON THE

its spawning ground a month later. But compare these scales

with text-fig. 115. This fish was forwarded to me from Toronto

on October 17th, so that it must have been procured at Lake

Shuswap some days earlier.

Text-fig. 115.

acvAN

Scale of Cohoe (O. kisutch), 9. 34 lbs. Shuswap Lake.

Weighed after spawning. °

According to Dr. Greene (Migration of Salmon in Columbia

River, Bulletin of U.S. Bureau of Fisheries, vol. xxix. 1909), a

Silver Salmon travels about 7 miles per diem. This fish, then,

to reach Shuswap by, say, October 12th, must have ceased feeding

in the sea not later than 23rd August. It will be noticed that

the outer lines of the scale are still widely spaced, showing that

summer feeding and growth were still in progress up to the time

at which the fish left the sea for the river.

THe Houmppack Saumon (Oncorhynchus gorbuscha).

I have but little to say at present about the Humpback, the

Dog Salmon, and the Steelhead Trout.

The Humpback has flesh of a pale pink colour. It was not

used for canning purposes until quite lately, but now there is

a good trade in it with China and Japan, where it is much

esteemed. The Chinese, not being accustomed to red-fleshed

fish, were very shy of the canned Quinnats and Sockeyes when I

was in China in 1893, and this may account for their partiality

to the Humpback.

The total weight of Humpbacks canned on the Pacific Coast in

1909 was 100,326,144 Ibs., and over another 3,000,000 lbs. weight.

was used in other ways.

HISTORY OF THE PACIFIC SALMON. 603

Humpbacks were first dealt with in the hatcheries in 1904, and

since then have been turned out from them intermittently, but

never in very large numbers, comparatively speaking.

They take their name from a peculiar hump which appears

behind the heads of the males at spawning time. They are from

3 to 11 lbs. in weight, and have very small scales.

The main run is in Alaska. South of Puget Sound the

Humpback is almost unknown. In the Fraser River the runs

in the big Sockeye years and in the years next but one after them

are much larger than in the other years. The fish, however,

Text-fig. 116.

Scale of Humpback Salmon (O. gorbuscha), from Puget Sound.

(For reference see text, p. 604.)

seem to come in at various ages, so this state of affairs] will

probably gradually right itself again if the netting is not too

severe. J am judging from possibly but a few specimens. A

large number of Humpback scales have been sent me, but,

unfortunately, with no further information than that they were

taken from Humpbacks in Puget Sound on their way to the

Fraser River. I append a photograph (text-fig. 116) of one of

these scales. The centre is only moderately well defined, but in

all the others it is even less clear, so that I have not been able to

604 MR. J. A. MILNE ON THE

form any definite opinion as to the time spent in fresh water.

This scale shows clearly one winter band formed in the sea (AA).

I presume that it represents the growth of the second winter.

All the scales show a similar hema and many of them another

evidently formed a year later. ’

I can give no measurements as I do not know the length, or

even the weight, of any of the fish when caught.

THe Doe Saumon (Oncorhynchus keta).

The Dog Salmon frequents the Pacific Coast all the way from

San Francisco to the Arctic Circle, but is most plentiful between

Puget Sound and South-East Alaska. ‘This fish is also called the

Chum Salmon, and is known in Japan as Sake, and in Siberia as

Kita.

I have not before referred to the prevalence of any of the

species on the other side of the Pacific, and only do so now

because the Dog Salmon has long been a staple food of Japan,

and it is because this fish is white-fleshed, or almost white, that

the red colour of the other varieties has caused them to be

viewed with suspicion when exported to the East. This fact

was ascertained by me when I was in China and Japan in 1893.

The Dog Salmon gets that name from the distorted appearance

of the mouth of the males at spawning time. They then look

not unlike snarling dogs. The average weight of the Dog Salmon

is about 8 lbs. In British Colombia it is higher, from 10 to

12 lbs. 16 lbs. would be about the maximum w eight.

The weight canned in 1909 was 25,660.845 lbs., and over

4,000,000 Ibs. weight was otherwise dealt with for export.

It is only in Washington State that these fish have been

spawned in the hatcheries. There an average of about 7,000,000

fry have been turned out in recent years beginning with the

year 1900.

Asa general rule the Dog Salmon comes in late in the year,

September to November. But in Alaska the run begins in June,

and in the Fraser River in the middle of August.

In British Columbia they spawn close to the sea. This I had

gathered from the appearance of the scales before I found out

that it was known to be the case. I have received scales from

two specimens weighing 114 and 12 lbs. respectively, both caught

in Puget Sound on their w ay to the Fraser River. They show

no indication of fresh water feeding at all. The fry evidently

descend to the sea as soon as hatched out. The measurements of

10 scales of the 114 lbs. fish, one of which is shown in text-

fig. 117, enable me to give the rate of growth as follows :—At

the end of the first year 71 inches, at two years 13 inches, at

three years 22 inches, at four years 274 inches, and when caught

in its fifth year it measured 31 inches. Its capture was evidently

HISTORY OF THE PACIFIC SALMON. 605

an early one, as the summer growth is only just complete. Un-

fortunately I have not got the date. The 12 lbs. fish was caught

at the same time and place. It was shorter, 302 inches, but

thicker, 17 inches. Its length at the end of each winter was

54, 11, 203, and 263 inches. From a smaller beginning it was

steadily overtaking the other fish in length, and had already done

so in weight when caught.

Text-fig. 117.

Scale of Dog Salmon (Oncorhynchus keta). 9. 11% lbs.

Length 31 inches; girth 16? inches.

THE STEELHEAD Trout (Salmo gairdneri).

The last of the Pacific Coast salmon with which I am dealing

is the Steelhead, which is there called a trout, but is the only

Pacific member of the sub-genus Salmo, to which the true salmon

belongs. It is believed to be a migratory form of the Rainbow

606 MR. J. A. MILNE ON THE

Trout. The flesh is pale-coloured, and is not much used for

canning.

The Steelhead is found from Carmel River, California, north-

wards to Central Alaska. In California it may be fished for only

with hook and line. In 1909 4,229,704 lbs. weight was canned

and 1,960,000 lbs. weight was otherwise used in the United

States fisheries, without counting those frozen or salted in British

Columbia.

Text-fig. 118.

Scale of Steelhead Trout (Salmo gairdneri), 8. 122 lbs. Length 33 inches ;

girth 163 inches. New Westminster. Autumn,1912. (For references see text, p. 607.)

In 1910 there were liberated by the U.S. hatcheries 12,023,646,

but none was turned out in Canada.

The spawning season is in the spring, but Steelheads are to be

HISTORY OF THE PACIFIC SALMON. 607

found in fresh water at most seasons of the year. They weigh, on

an average, at different places from 8 to 15 Ibs., witha maximum

of about AD lbs.

The scales, of which I have specimens from several fish, but

from only two that had been weighed and measured, show that

life in fresh water before the first migration to the sea may be of

considerable duration. In this species, as in the case of our own

salmon, no doubt is possible as to the limits of the part of the

scale formed in fresh water. From examination of a large

number of scales from my two measured fish, I judge them to be

about 7 inches long when they enter the sea. As spawning takes

place in the spring, the number of lines on the scale at the end of

the first winter is naturally very small. The scale illustrated in

text-fig. 118 shows five up to the point A, and that seems the

usual number, but the first band is not always visible on all the

scales, which is not to be wondered at seeing how minute they

are, =; inch long at the most, at that time.

A second winter in fresh water is shown in text- fig. 118 by the

band B, and a third by the band C. Outside of i are two or

three more lines added to the parr scale in the fish’s fourth

year, before it entered the sea. All the Steelhead scales which T

possess show similar lines and bands, so I suppose one may take it

that these fish remain in fresh water until well on in their fourth

spring, counting that in which they were spawned as one. They

would then be just three years old.

When they get to the sea they grow very rapidly. The first

band completed in the sea within a year of migration shows my

two measured fish to have been then already from 194 to 202 inches

long (see text-fig. 118, E), and the second winter band shows a

length of from 29 to 301 inches (text-fig. 118, F), but I am not

sure that I have identified this band cor rectly on the seale

illustrated. These two fish were both caught near the mouth of

the Fraser River in the autumn of 1912, when they measured

312 and 33 inches respectiv ely and weighed 12? and 13 lbs.,

the shorter and thicker fish being a female. They Tad then done

a considerable amount of feeding i in their sixth year.

This completes for the present my review of the salmon of the

Pacific Coast of North America.

It must always be borne in mind that my observations have

been made on the scales of fish from the Fraser River district

alone, and that readings from the scales of salmon caught else-

where might show considerable differences. Some dive gence is

almost cer Seat in the duration of freshwater life, for observations

of the scales of the true salmon (Salmo salar) hee shown that

the further north one goes the longer does the young salmon

remain in fresh water, Band mtn as “aeh at all unlikely that the

Pacific salmon are influenced in the same way by climatic

conditions.

608 MR. J. A. MILNE ON THE

The strongest argument which I have heard in favour of the

prevailing idea that Pacific salmon migrate to the sea as soon as

they can swim, is that many of the rivers which they frequent

are small, and, at times, are either dried up or frozen hard. This

argument was put before me by a gentleman who is largely

interested in the Pacific Coast fisheries and generally recognized

as an authority thereon. It therefore seems to demand an

answer. My answer is that I have examined only the scales of

fish hatched in the large watershed of the Fraser River, and that

possibly the fry of the Pacific salmon can adapt themselves to

varying conditions. It is known that our own salmon, which

generally migrate to the sea at two years of age, may enter it at

one year without hurt, or may remain in fresh water for three,

four, or even five years where the circumstances, as in Norway,

make it desirable for them to do so.

But, even supposing that they can adapt themselves to a

certain extent, I fail to see how any salmon can perpetuate its

species in a river which is frozen absolutely solid in winter, or

which is bone-dry in summer. In fact, I believe that the fish

that run into small rivers where such conditions prevail have not

been hatched in them at all, but are really natives of one or other

of the large river systems, and that, when they get into one of

these smail rivers, they become at once of no further use to their

own species, and might as well all be promptly utilized as food

for the good of ours.

For a fact which supports this belief I have only to refer to

what Mr. W. L. Calderwood says in a communication to the

‘Salmon and Trout Magazine’ for December 1912, page 24. He

there describes Capt. Callbreath’s hatchery in Alaska. Iam not

now concerned with the hatchery itself, but with the fact that it

is on a river, the Jadeska, only half a mile long, which runs out

of a small lake. A dam has been thrown across the river, only

100 yards from the mouth, so as to intercept all fish, and enable

the operators to select the Sockeyes and refuse the other salmon.

In addition to this dam a fence of racks has been erected having

at one place a trap. Here the Sockeyes have been taken out and

lifted over the dam, if not required for the hatchery, while the

other species have been left below. This selection has gone on for

eighteen years, and still continues, although the Sockeye hatchery

was a failure and hatching operations were discontinued in 1906.

Mr. Calderwood says :—‘ It is an interesting point that, although

the natural and artificial propagation of the Humpback and Dog

Salmon were entirely discouraged, the number of these fish

continued apparently undiminished.” He adds, and this is my

point, ‘‘ The moral may be that the fish in this river, like the

fish in other small rivers, are largely drawn from other sources.”

But, in this case, as none but Sockeyes have been admitted

to breed, it would seem proper to substitute the word “ entirely ”

for the word ‘“ largely,” and I submit that what applies to one

small river may equally well apply to them all.

HISTORY OF THE PACIFIC SALMON. 609

If the perusal of what I have written induces others to pursue

this fascinating study of scales upon the lines I have suggested,

my work will have served its chief purpose.

That the study is a most important one cannot be denied. The

salmon fisheries of the Pacific Coast are a source of vast wealth to

the countries so fortunate as to share in them. But that source

of wealth, like our own salmon fisheries, already shows signs of

having been too lavishly drawn upon. It is easy to kill the goose

that lays the golden egg. By legislation much may be done to

prolong its life, but legislation can effect its purpose only if based

upon thorough knowledge. In this country, if salmon scales had

been studied before the Acts which regulate our fisheries had been

passed, these Acts would have been very different from what they

are. The study of scales has so revolutionized our knowledge

that, of the eleven facts in the life of the salmon, mentioned by

Mr. Cholmondeley Pennell in 1886 as absolutely proved, one,

the most important, has now been shown to be absolutely

wrong, and two others, hardly less important, require much

modification.

I have not made myself familiar with the varicus fishery laws

and regulations of the Pacific Coast, but I know that many

experts think them insufficient. If the regulations as to netting

are to be made more stringent, let it be done with as complete

knowledge of the various salmon as possible ; and in order that

that knowledge may be obtained let their scales be systematically

studied. For months, or years, the fish are hidden from our eyes,

but from their scales we can fill in the gaps while they are out

of view, for it has been well said that every salmon carries a

record of its life on each of its scales. Whether I have read the

records written on the scales of the Fraser River salmon correctly

or not, I can at least claim to have proved that the records are

there.

In conclusion, I have to thank Mr. W. J. Sim of Northville,

Kings Co., Nova Scotia, for much valuable information, and for

having procured me numerous specimens through his friends

Mr. Frank Parry, now also of Northville and formerly of Granite

Creek, Mr. J. Brown of Celista, B.C., and Mr, Josephson of

Bellingham, Washington, U.S.A., and Mr. J. Kirkpatrick of

Vancouver. The various statistics I have given have been

compiled mainly from “The Salmon Fisheries of the Pacific

Coast,” by J. N. Cobb, Bureau of Fisheries, Document 751,

Washington, 1911, and partly also from the Report of the

Commission of Conservation on ‘ Lands, Fisheries and Game,

and Minerals,” Ottawa, 1911.

The following table may y;rove of interest :—

ON THE HISTORY OF THE PACIFIC SALMON.

| |

sot S = josh

i~ | . PS =| | O&

. aS s as PS S -S

Oe Geen a Hea bee iss

ae Bs |} Sos aS | CaS ot

jz 2 SS | 22 Bars | pe | oS

os SS | os sl p} o's

= Ss | Oo BS ey =a

RO os | S =. iS) a)

| ai We aes mlcesie

| a

it Pane Ee eee

Ome | Number of gill-rakers)

Ze) Ss | onthe anteriorarch ...) 32-40 | 20-28 | 20-25 | 28-36 | 20-25

on | |

BESS | Saar | *

: a | d

3 B sae | Number of well-deve-,

I San | loped rays in anal fin.) 14-15 | 15-18 | 12-14 15-16 | 13-14 10-12

3 out e —= ae |

fs oF — | | |

° ABS ‘Number of scales on

fe »o Me | dJateral line............... 125-185 | 140-1655 | 125-135 150-170 135-145

Aaa |

| fa, “Se =

io = Q2s). |||

| ear 4s : -

sa» Number of tmes of scales | |

S845 between lateral line’ |

q |e antd dowsalefiney ee. 20 25 nO BO BS

| |

Number of pounds canned, pickled,

frozen, or salted on the Pacific Coast

in 1909, in millions of pounds—nearest

snillion zie) 13 iQ | 1108 | a0). | °

F Daan

Number of fish caught in 1909, calculated |

from total weight as given above, and | |

general average weight in millions, | |

after allowing one-fifth for waste ...... 54 3 | 5 Bt z

eda De | | | u

Number of fish liberated from the) |

hatcheries in 1910 in millions—nearest |

million 316 Oi | sO 2 " 12

Average weight of fish, as caught, in) |

pounds 3-10 18-30 3-8 A Sales alo

Maximum weight attaimed, in pounds. |

aboubie erence cana enbacce ce sautiea decisis 17 100 | 30 11 16 45

Approximate time spent in fresh water!

before migration to the sea, in Fraser

River district, in months 12-15 18 18-24 ? 3orless, 36

ee | ary

Approximate length at time of migration

to the sea, in Fraser River district,

in inches 3-33 4-5} 23-5 P jlkorless 7

s a . diss

Age at time of return to fresh water, m)

Fraser River district, in years; as)

33 33-42 33 25-38 43 55

| shown by my specimens. ..................

MISS KATHLEEN HADDON ON PERIPATOIDES WOODWARDI. 611

38. Notes on Peripatoides woodwardt Bouvier.

By Karaieen Happon *.

[Received April 11, 1913: Read May 6, 1913.]

This species has been described by Prof. Bouvier t. It may

readily be distinguished from other Australian species by the

fact that it has 16 pairs of legs; in this respect this species and

P. sutert from New Zealand are unique among all the Australasian

Peripatoides. .

The material was obtained through Prof. J. Stanley Gardiner,

from Western Australia.

There are twenty specimens, well preserved and extended, but

they tend to be rather bent, thus making exact measurement

difficult. The approximate measurements (in millimetres) are as

follows :—

Females ... 46, 43, 43, 41, 37, 37, 33, 23, 21, 17.

Males ...... 31, 30, 30, 29, 27, 24, 24, 24,23, 29.

It will be seen, therefore, that these specimens are considerably

larger than those described by Prof. Bouvier {, the greater

number of which measured from 10°5 to 17 mm., only one

reaching the length of 21 mm.

Bouvier recognises the following types of coloration :—

1. Uniform, with predominance of blue-green pigment,

yellowish pigment being strewn in spots equally dis-

tributed over the dorsal surface.

2. Striped, with the yellow spots more numerous, sometimes

becoming predominant, and forming a longitudinal

band, with or without dark papille, on each flank above

the base of the legs.

A variation of the first type had the ground colour light

reddish yellow, almost all the papillae being darker, at any rate

at the summit; some were quite black and arranged in short

transverse rows. Thus the general tone was a dingy grey-black

with small spots of yellowish pigment round some papille.

A variation of the second type showed a yellow-red ground

colour, with stripes of dark papille in transverse rows in the

dorsal region and no mixture of dark pigment in the lateral

stripe.

All his specimens showed two large light-coloured papille on

the flanks between each pair of legs, arranged parallel to the

long axis of the animal.

These types are all represented in my specimens and also all

* Communicated by Prof. J. Srantey Garpiner, M.A., F.RS., F.Z.S.

+ ‘Die Fauna Siidwest-Australiens,’ Bd. ii. No. 18, Onychophora. M. E.-L.

Bouvier. Jena, 1909.

i Doc. cié. ps 315.

Proc. Zoou. Soc.—1913, No. XLI, 4]

612 MISS KATHLEEN HADDON ON PERIPATOIDES WOODWARDI.

the intermediate stages between them. Those belonging to the

first type show in addition a well-marked lateral stripe of a

greenish-blue colour. The antenne are also of a greenish-blue,

thus differmg from those of Prof. Bouvier’s specimens which he

describes as always being black. Four of the animals are

very dark in colour, one being practically black, but all have a

brownish-green tinge : the ventral surface is lighter, but even

then darker than the backs of the other specimens. The darkest

specimen, which is a male, differs from all the others in having

the crural papille black instead of white. All the animals show

a patch of green pigment on the inner side of each leg which is

extended on to the ventral surface of the body. In most cases

the ventral and pre-ventral organs are not well-marked, but some

of the smaller specimens show the ventral organs distinctly.

The two lght coloured papille described by Prof. Bouvier as

situated between each pair of legs are not apparent in these

specimens.

The coloration of these Peripatus agrees fairly closely with

that of Peripatoides orientalis *, except that the dark variety is

by far the most common in the last-named species (772 per cent.

according to Steel t), whereas in P. woodwardi they are in the

minority, as far as our present material goes.

With regard to the external features, these specimens of

P. woodwardi agree with those described by Prof. Bouvier ¢ in

every respect except for the number of crural papille in the male.

These are very prominent and capable of retraction within the

leg, and he describes them as present on all the legs with

the possible exception of the 15th and 16th, that is the last two

pairs, while he found coxal glands in all the legs except possibly

the 15th pair.

I have examined all the males in my material, and although

the papille show extremely clearly in most cases, I can find no

traces of them on the 4th or 5th legs; they appear to be entirely

absent from these legs, as well as from the 15th; the last pair

of legs may show a small papilla. In the male that I dissected

the coxal glands showed very clearly in the other legs, but no

trace appeared in the 4th, 5th, or 15th pairs. The absence of

these glands in the 4th and 5th pairs of legs seems to be correlated

with the great development of their nephridia, which open by a

urinary papilla on the proximal of the three spinous pads of the

foot.

As deseribed by Prof. Bouvier, the first pair of crural glands

are extremely long and lie in the Jateral sinus with the nephridia ;

the last pair are elongated and iridescent and lie in the lateral

sinus or in the general cavity of the body, while the remaining

* “Monographie des Onychophores.” E.-L. Bouvier. Ann. des Sc. Nat. vol. v.

9th series, 1907, p. 233.

+ “Observations on Per ipatus,” Proc. Linn. Soc. N. S. Wales (2) vol. xxi. 1896,

p- 94.

{ ‘Die Fauna Stidwest-Australiens,’ p. 320.

ON THE ENEMIES OF BUTTERFLIES. 613

ones are of the same type as those of P. orientalis, a vesicular

gland with a narrow duct lying in the leg.

The only other particular in which my specimens differ from

those of Prof. Bouvier is in respect to the female generative

organs. In the two examples that he dissected there was little

or no trace of a receptaculum seminis, but he points out that the

ovaries were very large, thus making the dissection difficult.

In both the females that I dissected it was easy to trace the

oviducts for the whole of their course, and shortly after their

emergence from the ovarian sac, where they at once diverge from

one another, there was a well-marked receptaculum seminis on

each side, full of sperm. This is another point of resemblance

between this species and P. orientalis, which seems on the whole

to be the most nearly related form.

39. Field-Observations on the Enemies of Butterflies in

Ceylon. By J.C. F. Fryer, M.A., F.E.S.; Fellow of

Gonville and Caius College, Balfour Student in the

University of Cambridge. * ,

[Received April 22, 1913; Read May 6, 1913.]

Ermonocy (Birds AND BUTTERFLIES).

The subject of Birds and Butterflies in Ceylon has already been

admirably dealt with by Col. N. Manders in his paper entitled

“« An Investigation into the Validity of Mullerian and other forms

of Mimicry, with special reference to the Islands of Bourbon,

Mauritius, and Ceylon”. In spite of this, however, I venture

to place on record a few further notes from the island, partly

because they deal more fully with the butterfly-eating habits of a

most interesting bird, the Wood-Swallow (Artamus fuscus Vieill.),

but also because the subject, with its direct bearing on the various

theories of mimicry, is essentially one which requires as many

observations as possible before any fair generalisations can be

mace.

A residence of a year and nine months, largely devoted to

entomological research, gave an opportunity for keeping a con-

tinual watch on certain of the more common insectivorous birds ;

and the following notes comprise every observation made during

that period. The locality in each case, unless otherwise stated,

is the celebrated Botanical Garden at Peradeniyat. Other

localities mentioned are Vavonia-Valunkulam, Maha I]lupalama,

and Trincomalee—all places in the hot dry northern country,

which is characterised by the large area of jungle it contains,

* Communicated by Prof. J. SranntEy GarpiIner, M.A., F.R.S., F.Z.S.

+ Proc. Zool. Soc. 1911, p. 696.

t Elevation about 1600 feet.

Ase:

614 MR. J. C. F. FRYER ON THE

and also, in my experience, by a greater quantity of butterflies

and other insect life than can usually be found in the hills.

The notes are given in chronological order, and are extracted

from my journal practically without alteration, as this method,

though possibly wearisome, places a check on the personal pre-

judices of the observer, the introduction of which it is difficult to

avoid in a mere summary of results.

PERADENIYA.

1911. April 7th. A drongo* took a Papilio polytes Linn. 3,

which could hardly fly, as it had only just emerged ; it then

caught a Melanitis ismene Cram., also released soon after it had

left the pupa.

April 2nd-9th. General note that “drongos were only observed

to take small butterflies, chiefly Yphthima ceylonica Hewit.,

Lyceenids, and occasionally Precis iphita Cram.” ‘ Of the various

attempts quite one out of two missed their object, while in many

cases the swoop at a butterfly seemed to be taken more for amuse-

ment than to gain food.” ‘“ Paradise fly-catchers (Terpsephone

paradisi Linn.) being more retiring, were harder to observe ; they

certainly took Yphthima ceylonica and small Lycenids, but were

not observed to attack anything large.” [This week, April 2nd—

9th, was spent mainly in bird observations, and the above is the

result. |

April 15th. A drongo was seen to attack Catopsilia pyranthe

Linn., but apparently without success.

May 9th. A drongo was seen to take an Yphthima ceylonica in

the garden. ... Three drongos chased a Papilio parinda Moore,

but did not catch it.... Two bulbuls (A/olpastes hemorrhous Gm.)

chased an orange-coloured moth, probably Argina sp., but lest it.

June 25th. A Paradise fly-catcher swooped at a Limenitis

calidasa Moore, which escaped.

July 9th. A magpie-robin (Copsychus saularis Linn.) took a

specimen of the Chalcosiid moth, Heterusia cingala Moore +, and

flew away with it.

October 16th. A drongo chased a Papilio parinda, but lost it.

October 19th. A drongo caught a Papilio polytes 3, and ate it.

October 21st. A scanty P.-polytes migration for the last three

days... . A drongo swooped at a P. polytes 2 form polytest, but

missed it. Many females of the romulus form § with the entire

hinder portion of the hind wings missing.

November \st-15th (at Vavonia-Valunkulam). A continual

migration of Appias (A. paulina Cram. and A. albina Boisd.),

* Throughout, the word ‘“‘drongo” is used for either Dicrurus leucopygialis

Blyth, or D. cerulescens Linn. The former was more common; but the species

in wost cases was not ascertained.

+ This should undoubtedly be an “inedible” species, as it has all the Chalcosiid’s

nauseous characters to a marked degree.

“ + P. polytes 2 polytes resembles Papilio aristolochie Fabr.

§ P. polytes 2 romulus resembles Papilio hector Linn. Both P. aristolochie

and P. hector are common Ceylon insects.

ENEMIES OF BUTTERFLIES. 615

Catopsilia (C. crocale Cram. with a few C. pyranthe Linn.),

Papilio polytes, P. jason Linn., P. nomius Esper, and P. hector

Linn. Bee-eaters (Merops sp.?) were common, but were only

observed to chase butterflies (P. jason, Appias sp.) on three

occasions, each attempt being unsuccessful. Wings of all the

above butterflies were found occasionally along the roads, but not

in any number.

November 25th (at Peradeniya). A drongo seen to take a

Papilio polytes 2 romulus, which was flying badly.

November 29th. A number of wings of Huplaa and Danais

found on the ground in the garden. ‘The only birds seen near

were wood-swallows (Artamus fuscus Vieill.). The wings iden-

tified were :—Huploaa core Cram., fore wings 6; #. kollari Feld.,

fore wings 2; HL. coreta God., fore wing 1; Danais septentrionis

Butl., fore wings 5, hind wings 3.

November 30th. Spent an hour, 11 A.m.—12, watching the wood-

swallows hawking butterflies, which appeared to be forming their

sole food. The birds sat on the top branches of a leafless tree,

and swooped out after their prey, usually effecting a capture.

‘The successful attempts were estimated at two out of three. The

butterfly was carried to the perch and the wings there discarded.

Butterflies on migration were extremely abundant, and indi-

viduals of the genus Catopsilia were perhaps most numerous,

though Eupleas, Danaids, Papilio polytes and P. demoleus Linn.

were all quite common. ‘The wood-swallows limited themselves

almost entirely to Eupleas and Danaids, only three swoops at

Catopsilia and one at Papilio demoleus being observed, and these

were all failures. The majority of the discarded wings were

blown into the river, but the following were collected, practically

all of which must have fallen during the morning.

Result :—Danais septentrionalis: fore wings 61, hind wings 9.

Huplea core: fore wings 48.

EL. coreta: ss BAe AS)

E. kollari : bs , 6.

Kupleine hind wings 9, fragments 4.

Papilio demoleus: fragments 2.

P. polytes 3 ov 2 cyrus-form*: fragments 7

(2 fore wings).

P. polytes 2 romulus: fragments 5 (3 fore wings).

P. polytes 2 polytes: hind wing 1.

The predominance of fore wings is explained by their greater

weight, hind wings being blown further.

A pair of bee-eeters were feeding near the wood-swallows.

Most of their attacks were on small insects; but four attempts

on Catopsilia and two on Papilio demoleus were noted, all

of which failed. These bee-eaters were either unskilful or

only amused themselves with butterflies. A young Paradise

* Papilio polytes 2 eyrus-form resembles the g.

616 MR. J. C. F. FRYER ON THE

flycatcher and a drongo confined themselves to small insects,

in spite of the swarms of butterflies passing.

December \st. Again visited the wood-swallows, but they left off

feeding early ; the butterfly migration was larger, with a higher

percentage of Huploea. It was distinctly observed in one case that

the bird carried the butterfly, a Danais septentrionis, to its perch

and then held it with its feet while it pulled the wings off with its

beak *. Watched the bee-eaters again, and saw one attack first

a Papilio agamemnon Linn. and then a Huplaa, but it missed

each time. They were continually hawking small insects, which

they presumably caught. No count could be made of discarded

wings, as all had been blown by a high wind into the river.

December 2nd. Again watched the wood-swallows hunting.

The butterfly was always carried back to the perch before the

wings were removed, and it was noticed that portions of the wings

were often left on onal eaten.... After failing in a stoop, the birds

varely made another at the same insect. ... Butterflies at some

distance were usually taken, perhaps to allow the bird to attain

greater pace... . If a butterfly noticed that it was being pursued,

it at once dodged and, flying rapidly downwards, frequently

escaped. Wings were again collected :—

Danais septentrionis: fore wings 35.

D. Vimniace : s Kage

Huplea core: é

EF. coreta: one whole insect and fore wings 10.

i. kollari: fore wings 4.

Papilio polytes 2 romulus: fore wing I.

December 7th. Wood-swallows hunting in the afternoon; they

usually stop before midday.

December 10th. A young Paradise flycatcher swooped at two

Papilio polytes,a 3 anda 2 polytes form, which were flying round

inside a large breeding-cage, but of course could not get at them.

December 16th. A drongo was seen to attack a Papilio aga-

memnon and snap off a portion of the hind wings. The butterfly

escaped.

1912. January \st-6th (at Maha Illupalama). Bee-eaters

were very common and butterflies, notably of the genera Cato-

psilia and Appias, were abundant. The only attack observed was

that of a bee-eater on a Leptosia xiphia Fabr., which was caught

and eaten.

January 10th (at Peradeniya). A drongo caught a Papilio

polytes Q eyrus-form, which had just emerged and had escaped

from a breeding-cage.

February 2 Ond. A Papilio polytes 2 cyrus escaped from a cage

and was eaten by a drongo; it was flying Heay) weakly, as one

wing was broken halfway down the costa.

* Legge, ‘Birds of Ceylon,’ vol. 11. p. 668, states that the wood-swallow beats off

the wings of its prey.

ENEMIES OF BUTTERFLIES. 617

March 6th. Noticed three wood-swallows sitting on the

telephone-wires crossing the River Maha-Weliganga. They were

hawking butterflies continually, and during the time they were

watched selected species of Huplea only from a passing migration

which consisted largely of Appias paulina. A search for wings

failed, as the birds confined themselves entirely to the river. ‘The

reason for this seemed to be that the butterflies were only caught

with ease when away from cover; among vegetation they dodged

their pursuers nine times out of ten. This fact was specially

evident during this migration, as species of Huplea and Danais

were uncommon, and the birds were often forced to swoop at

butterflies travelling along the river-bank instead of waiting for

individuals to come out into the open.

March 8th. The migration of butterflies almost ended; but to

obtain confirmation, by a fresh witness, of the fact that wood-

swallows attack esgectlly members of the genera Huplea and

Danais, Alexander * and I watched three wood-swallows for an

hour, and we both plainly observed them eat over a dozen Hunlea

and one Danais. We also saw one bird with a Papilio, either

P. polyies 2 polytes or P. aristolochie, but were unable to

ascertain which. As before, the birds usually carried the butter-

fly in the beak, though occasionally it was transferred to the

claws when the distance back to the perch was great.

March 15th. Watched the wood-swallows for three-quarters of

an hour; Appias spp. common, Luplea and Danais scarce. There

were only two birds feeding, one probably young, for when the

other caught a Danais aglea Cram. the young one opened its

beak and fluttered its wings as if asking for food, though it

gained nothing by this proceeding. The old bird then ceased

feeding, but the other, possibly encouraged by the sight of the

Dearne, worked steadily, soaring round in circles over the river.

It ca ught five Eupleas- out of seven attempts.

March 16th. Watched a wood-swallow for a quarter of an hour

and saw it take two Kupleeas.

March 17th. Watched a wood-swallow from 11.30-12. It took

two Euploeas, but then seemed to have no room for more, as it sat

for a long time with the abdomen of the last victim protr uding

from its mouth like a cigar.

March 18th. In five minutes saw two wood-swallows take three

Eupleas and a Papilio, either P. polytes 2 polytes or P. aristo-

lochie, almost certainly the former.

March 22nd. Species of Huplea and Danais abundant on

migration. A wood-swallow picked out two Danaids (D. limniace

and D. septentrionis) and then a Huplaa, after which it stopped

feeding.

Mar ch 23rd. Watched the wood-swallows from 11.40 a.m.—12;

two were under observation most of the time, occasionally four.

They took six specimens of Huplaa and two D. septentrionis.

* Mr. Alexander, Assistant Curator of the Perth Museum, W.A., to whom I am

indebted for much critical assistance during his brief visit to ‘the isl ind,

618 ON THE ENEMIES OF BUTYERFLIES.

April 23rd. A drongo swooped at and probably took either a

Papilio polytes 2 polytes or a P. aristolochic.

April 24th. Mr. Pole of Maskeliya* wrote that he saw two

large dragonflies take two Appias sp. ? and one P. polytes 3.

July Tth (at Maha Illupalama). Noticed the remains of Danais

septentrionis, Papilio jason, Appias paulina and A. albina, killed

probably by Asilid flies (Scleropogon piceus and other species), as

the bodies were intact though empty. Asilid flies were twice

seen carrying Appias sp.? 3.

August 8th (at Maha Illupalama). Caught an Asilid fly with a

medium-sized dragonfly +, which it was sucking through the eye,

and later saw another Asilid killing a large Cicada $ which was

flying madly about, shrieking (stridulating) all the time. A

slight bite from an Asilid was most painful, and felt as if

poisonous matter had been injected.

August 13th (at Maha Illupalama). Noticed Asilid flies kill

Appias sp.? 3 and Papilio jason.

August 28th (at Habarana, Maha Illupalama District). An

Asilid attempted to kill the formidable Ceylon hornet, Vespa

cincta, which, however, escaped.

August 31st (at Trincomalee). A few Euplea wings were found

scattered about Fort Ostenburg ; they were probably dropped by

wood-swallows, which were hawking round the inner harbour.

September 1st (Trincomalee). On the Nilavelli-Kuchivelli road

bee-eaters were numerous. One was seen to catch an Appias sp.,

and wings of Huplea, P. jason, and Appias spp. were occasionally

found on the ground.

September 7th (at Peradeniya). Released an Actias selene 3.

It flew fairly well, but was soon snapped up by a drongo, which,

before eating it, first pulled off the “ tails” and then the rest of

the wings.

The observations having now been given at length, it is

perhaps permissible to summarise the impressions gained in their

making :—

i. Butterflies do not form any large percentage of the food

of the more common insectivorous birds in Ceylon.

il. With the exception of the Wood-Swallow, birds are by no

means clever in capturing butterflies.

ili, The Wood-Swallow is the only bird which actually lives on

butterflies, and it almost always chooses butterflies of the

so-called nauseous genera Danais and Huplea; it seemed,

however, that this preference was due to the difficulty of

catching faster-flying butterflies, and not to the superior

flavour of the Danais or Huplea.

* Hill district, elevation about 4000 feet.

+ This and one or two subsequent notes, though not dealing with butterflies, are

included, as they seem worthy of record and yet are insufficient for a separate

contribution.

Pola ch Sie, Pl LQ,

C.D. photo.

EUPARKERIA CAPENSIS

1913. Pl. LXXVL

28)

University Press, Cambridge.

R-Broom ,del.

EUPARKERIA CAPENSIS.

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R.Broom,del. University Press, Cambridge.

9,10. ORNITHOSUCHUS WOODWARDI. 11. 0. TAYLORI.

P.Z.S. 1913. Pl. LXXVIUL

G en Oeononome

R. Broom ,del.

University Press, Cambridge.

MESOSUCHUS BROWNI AND OTHERS.

P.Z.S. 1918. Pl. LXXIK.

R.Broom,del. University Press, Cambridge.

BROWNIELLA AFRICANA AND OTHERS.

ON FOSSIL REPTILES. 619

iv. The various species of Bee-eaters, when attacking butter-

flies, usually choose members of the Papilionide and

Pieridee.

v. The Asilid diptera are formidable enemies to all other

living insects. They are, however, present in large

numbers only in the northern country, and then only for

a limited period of the year.

vi. In Ceylon a resemblance to the genera Danais and Huplea

is doubtfully of value ; in fact, in the neighbourhood of

Wood-Swallows it is a distinct danger.

vii. The mimetic females of Papilio polytes are not obviously

protected by their mimicry, and as a whole probably

suffer about as much, or as little, from the attacks

of birds as any other fast-flying butterfly in Ceylon.

40. On the South-African Pseudosuchian Kuparkeria and

Allied Genera. By R. Broom, M.D., D.Sec., C.M.Z.S.

[Received and Read May 20, 1913.]

(Plates LXXV.-LXXIX.*).

InprEx. 4 Page

JD OG DOTIRAPIGL CHYDCISOS VEXTIOIN, oo nao ococboone a. 5senad ade coe no8eo0K0qbC 619

Ornithosuchus woodwardi Newton .........0.. cc. ccc cee vev eee eee eee 624,

Onmthoswenusiianloniasp aeeeerere eae eee eee ee OLD)

Herpetosuchus granti Newton .......2......:c0se000.se0ese00sss0200 626

Browniellanafricand, See CbiSpsMe 5: assess -eeteasseeeaee eee OT

TESOS WEDS WIRDUTOR NN ENBIOD aoe sao coo tosenseceansene sAseseoasosannosns (Ae

Scleromochlus taylori Smith-Woodward ........................... 629

Affinities of the Pseudosuchia.................20.c..csseeese0eeeeeeeeee 629

KUPARKERIA CAPENSIS Broom. (Pls. LXXV., LXXVL.,

LXXVIIT., LX XTX. figs. 1-8, 17-20, 22, 24.)

In the collection of Mr. Alfred Brown of Aliwal North are

a number of imperfect skeletons of a small Rhynchocephaloid

reptile of very great interest, which he discovered in Upper

Triassic beds about four yearsago. Mr. D. M.S. Watson examined

them a year ago, but unfortunately he mistakenly regarded them

as belonging to the little acrodont-toothed reptile named by

him Mesosuchus brown. Though the two forms were met with

together and are of about the same size, they certainly belong to

different families, if not to different suborders. In a short note

commmunicated to the ‘ Records of the Albany Museum’ (6), I

have called attention to the confusion and proposed the name

Huparkeria capensis for the Thecodont form.

As the type, I take a specimen showing a beautiful skull

with most of the skeleton. Though this specimen was in the

* For explanation of the Plates see p. 633,

620 DR. R. BROOM ON

collection as seen by Watson, it was scarcely at all developed,

and he did not recognise that the block of stone contained an

almost perfect skull which, had he seen it, would have prevented

the confusion of the two animals.

In the type specimen is preserved the skull almost complete

but shghtly crushed, and with the bones in beautiful condition and

showing nearly every detail of structure. A number of cervical

and densa vertebre are shown, both coracoids, the interclavicle,

the right clavicle and scapula, and the right marae on radius, and

ulna. There are also seen in the specimen the whole of the

abdominal ribs in perfect condition, the pelvic bones in good con-

dition, but displaced, and most of the left hind limb. Other

specimens show the pelvic bones in position and most of the tail.

One specimen shows a dentary with a perfect right pes.

The skull is very similar to that of Or “ailioenieaus woodward,

and there can be little doubt but that Hauparkeria belongs to the

same suborder and to the same family, though the two genera

must be regarded as distinct. From the snout to the occipital

condyle the skull measures about 83 mm. From the snout to

the front of the orbit is about 45 mm. The orbit is almost round

and has an antero-posterior diameter of 23 mm. There is a

large antorbital vacuity about 20 mm. in length. The infra-

temporal opening measures 18 mm. in height and its greatest

antero-posterior length at its lower part is 17 mm. The supra-

temporal fenestra is small, measuring 13 mm. by 10mm. The

width across the frontals between the orbits is 14 mm., and the

width across the squamosals where they form the temporal arch

is 34 mm.

The front part of the premaxilla is missing from the type, but

it was probably somewhat similar to that in Ornithosuchus and

other allied types. It forms the lower half of the posterior

margin of the rather large nostril. It supports at least two and

most probably three flattened pointed thecodont teeth.

There is no evidence of a septo-maxillary on the face.

The maxilla is a long slender bone, which forms the lower

and anterior borders of the antorbital vacuity. The anterior

ascending process passes up behind the premaxilla and the

descending anterior portion of the nasal, and meets the anterior

end of the large lacrymal. The posterior horizontal portion

passes back to below the middle of the orbit and meets the jugal.

It supports apparently 13 thecodont teeth, of which 5 are

preserved in the specimen. These are pointed flattened teeth,

very similar in general shape to those of carnivorous Dinosaurs.

They are feebly serrated behind and probably also in front.

The nasal is rather peculiar in shape. When viewed from

above, it appears as a long narrow bone about twice as wide

behind, where -it meets the frontal, as in front. In reality the

front is as wide as the back part, as it forms a curious downward

process behind the nostril to meet the premaxilla. The peculiar

shape will best be understood from the figures.

FOSSIL REP'LILES, §21

The lacrymal is an unusually large bone. It forms most of the

anterior orbital margin. Above, it has a long articulation with

the prefrontal, and below, a short one with the jugal. Anteriorly,

it has a large development which forms the upper margin of the

antorbital vacuity meeting the ascending process of the maxilla.

Of the anterior process much is below the level of the general

surface of the face, suggestive of the antorbital vacuity having

lodged a large gland.

‘Lhe prefrontal is a small narrow bone which forms about half

of the upper orbital margin. It is bounded above by the frontal

and nasal and below by the lacirymal.

The frontal is a long narrow bone. Behind, it articulates with

the parietal and laterally with the postfrontal, and in front

with the nasal and laterally with the prefrontal. It only forms

a small part of the orbital margin.

The postfrontal is a small triangular bone which articulates

with the frontal, parietal, and postorbital, and forms a small part

of the upper orbital margin.

The postorbital is a tiivadiating bone. The upper process

passes upwards behind the postfrontal and meets the parietal.

The inferior process passes downwards and articulates with the

jugal, partly lying in front of it and forming with it the post-

orbital arch. The posterior process is shorf. It meets the

squamosa! and forms with it the temporal arch.

The jugal is also a triradiating bone. The anterior process is

the strongest. It forms most of the infraorbital arch, forming a

long suture with the maxilla and meeting the lacrymal. ‘The

upper process meets the postorbital and forms with it the post-

orbital arch. The posterior process is long and slender and

forms the zygomatic arch meeting the quadrato-jugal.

The quadrato-jugal is an angular bone which supports the

quadrate and the jugal and binds them together. Above, it

meets the squamosal. A large foramen is present between the

quadrato-jugal and the quadrate near the lower part of the

bone.

The squamosal is small but fairly strong. It may be regaraed

as a clasping bone which holds together the quadrate, quadrato-

jugal, postorbital, parietal, and opisthotic. A small upper

process lies in front of the lateral process of the parietal. The

relations to the quadrato-jugal and quadrate will be better

understood by the figures given.

The quadrate is a long, well-developed bone. Its upper end

is firmly articulated with the squamosal and possibly also with

the opisthotic, and also meets the quadrato-jugal. The lower

end forms the articulation. There is a large opening between

the quadrate and quadrato-jugal. The shape of the bone is seen

in the drawings given.

The parietal is smaller than the frontal. It forms the back

part of the upper cranial wall and has a postero-lateral process

which forms much of the posterior wall of the upper temporal

622 DR. R. BROOM ON

fossa, and at its outer end meets the squamosal. There is no

trace of a pineal foramen.

Behind the parietals and partly wedged between them is a small

interparietal, It forms the upper part of the occiput, and partly

divides the parietal from the supraoccipital.

The supraoccipital forms the middle part of the occiput. It

articulates with the exoccipitals, the interparietal, and the

parietals.

The exoccipital forms part of the occipital condyle and passes

outwards, fusing with the opisthotic.

Only a small part of the basioccipital shows in the specimen

as the middle part of the condyle.

The lower jaw is well preserved. The dentary forms the

anterior half, and the angular and surangular the greater part

of the posterior half. A lar ge oval opening is seen on the outer

side of the jaw between the angular and surangular.

A pair of long rib-like bones represent portions of the hyoid

apparatus. These are evidently the ceratobranchials, and indicate

that Huparkeria had a birdlike tongue.

There are well-developed sclerotic plates in the eye, which are

curved as in the bird.

Two slightly displaced bones are probably the proatlas and

portion of the atlas. There are probably about 9 or 10 cervical

vertebrae, of which the upper 3 or 4 are hidden by matrix. The

lower cervicals have comparatively short centra. They are

practically amphiplatyan or incipiently procelous. The ribs are

double-headed and have small uncinates. There are 2 sacral

vertebra, and apparently 26 presacral. The tail is very long and

has powerful chevrons,

The shoulder-girdle is well preserved. ‘There is a long slender

interclavicle which is apparently narrow even at its anterior end.

The clavicle is also long and slender. The coracoid* is large,

measuring 24 mm. in antero-posterior diameter and 16 mm. in

its transverse diameter. ‘There is a large oval foramen near the

scapular articulation and a little in front of the median plane of

the bone. The scapula is long and slender. It measures in

greatest length 38 mm., and its lower end is 15 mm. across and

the upper end 12 mm. in width. There is no distinct acromion

process, and the clavicle has been probably rather loosely attached

to the front of the lower third of the bone.

The humerus is very slender. Its length is 36 mm. Only the

outer aspect is displayed, and it cannot be seen whether there

is an entepicondylar foramen. There is no indication of an

ectepicondylar foramen. The deltopectoral ridge is very short.

The radius and ulna are slender straight bones. The radius

measures 32 mm. in length and the ula) is probably slightly

longer. The manus is lost from the specimen.

* Though to avoid any confusion I have used the universally accepted term

“* coracoid,” as I have elsewhere recently shown the bone ought more properly to be

called the“ precoracoid.”

FOSSIL REPTILES. 623

There is a large broad plastron of abdominal ribs which

occupies the whole of the front of the abdomen, from a little

behind the coracoid to the front of the pubis. The riblets are

all slender and arranged in series of threes—a long outer riblet,

a shorter middle one, and a still shorter inner one. In front the

innermost series remain free, but on passing back they become

anchylosed into united groups of two, three, and four. There

are probably four series of riblets to each vertebra, Altogether

there are 31 groups of ribs.

The pelvis is preserved in two specimens besides the type, and

every detail of its structure is known. The ilium is similar to

the type found in Belodon and Hrythrosuchus. ‘The acetabulum

is large, and closed, and the ilium has a well-marked supra-

acetabular ridge. The iliac crest is developed much more back-

wards than forwards, and the whole crest measures 28 mm. in

length. The pubis is comparatively short and broad. It forms

a complete symphysis with its neighbour, and the outer border,

which is considerably thickened, passes almost directly down-

wards. A very remarkable feature of the bone is the presence of

two pubic foramina. The upper one, which is apparently the

homologue of the normal pubic foramen, is the smaller of the two

and is fairly close to the acetabulum. The other is close to the

symphysis. In the type specimen both pubes are well preserved

and the foramina are exactly similar on both sides. The ischium

is a variety of the plate-like form, but long and slender. It has

a complete symphysis with the other ischium.

The femur measures in the type 58 mm.: in a second specimen

55mm. It has a slight double curve as in most reptilian femora.

The ends have been largely cartilaginous. On the inner and

posterior side, near the union of the upper and middle third, there

is a well-marked small trochanter.

The tibia and fibula are not perfectly preserved in any of the

specimens, but in one or other both upper and lower parts are

shown. They are apparently a little shorter than the femur and

considerably more slender,

The tarsus consists of two proximal elements, and probably

four distal tarsals.. The astragalus is irregularly cubical, and the

caleaneum considerably broader than long. There appears to be

little in the way of a heel. Of the distal tarsals the 4th is the

only large one.

The metatarsals are all well developed, and the 5th has the

peculiar Rhynchocephalian development. Of the others the 3rd is

the longest. The 4th is slightly longer than the 2nd, and the 2nd

considerably longer than the Ist. The Ist, 2nd, and 5rd digits

have well-developed claws, but the 4th has the phalanges rather

weak and the claw, if present, was very small. The 5th toe

also has a small claw, though larger than that of the 4th. The

digital formula is 2, 3, 4, 5, 3.

Dermal ossifications are present along the back from the head

to at least well down the tail. These are arranged in pairs, one

624 DR. R. BROOM ON

on each side of each vertebral spine. All the best preserved

scutes are about twice as long as broad and have the long axis

lying antero-posteriorly,

Some at least of the ribs of the middle region of the body

appear to have uncinate processes. These are little ossifications

about 3 or 4 mm. in length and about 1 mm. in width. They

are firmly attached to the posterior side of the ribs but not

anchylosed. Just possibly, though much less probably, they are

smal] lateral dermal scutes.

Before discussing the affinities of Huparkeria and the Pseudo-

suchians generally, it will be well to consider some points in the

structure of the allied genera from Elgin. These very interesting

specimens, preserved in the British Museum, reveal a few points

in the structure of the group not seen in the specimens of

Euparkeria, and afford a very thorough knowledge of the sub-

order. Aétosaurus ferratus, on which the subondies Pseudosuchia

was founded, has never been very thoroughly described, but

Mr. Watson assures me that there is no doubt that it has two

temporal vacuities like those of Huparkeria and Ornithosuchus,

and not one as described and figured by Fraas. Though there

is thus httle doubt that Aétosaurus belongs to the same embondler,

it differs in a good many points from the Elgin and South-African

forms.

ORNITHOSUCHUS woopwARDI Newton. (Pls. LXXVII.-

NONE XG ois Oe OG 525).))

The type of this Pseudosuchian is a fairly complete skeleton from

Elgin, described in 1894 by Mr. E. T. Newton. The specimen is

preserved in the British Museum, and through the kindness of

Dr. Smith Woodward I have been enabled to make a fresh study

of it in the light of the new knowledge obtained from the allied

South-African form. Newton’s study of the type has been so

thoroughly and carefully done that there is very little in the

specimen he has failed to observe, and the points where I incline

to differ from him are very few in number. Fortunately the

British Museum has recently obtained a second specimen which

supplies a number of blanks in our knowledge.

The skull, on the whole, resembles that of Huparkeria in all

essentials, so far as can be seen, though the cranial sutures are

less easily made out in Ornithosuchus and the palate is unknown

in Huparkeria. The skull of Ornithosuchus woodwardi is more

slenderly built, and it differs in having apparently no inter-

parietal, in the shape of the jugal, in “having a much larger

antorbital vacuity, and in having the teeth differently arranged,

In the restored side view of the skull which I give, and which

differs only slightly from that given by Newton, the shape of

the various openings and bones can readily be seen. The dental

formula of Ornithosuchus appears to be 1.3, m.9 as against

i.3, m.12 in Huparkeria; and there is the further difference

that, while in the South African genus only one mandibular tooth

overlaps the upper jaw-border, in Oriithosuchus there are two

FOSSIL REPTILES. 625

teeth which pass to the outside between the premaxillary and

maxillary teeth.

The palate of Ornithosuchus is well preserved but not com-

pletely displayed in front. The restoration of it which I give is

hypothetical only in the prevomerine region and in the basi-

occipital. The pterygoids are large, as are also the palatines

and ectopterygoids, and these three “bones of the two sides forma

large vaulted bony roof to the mouth. On each side there is a

pair of oval openings—one between the palatine, the ectopterygoid,

and the pterygoid, and the other between the pterygoid and the

palatine. This latter is regarded by Newton as the posterior

nares. In this, I think, he is in error. The opening has the

borders quite flat, and was probably covered by membrane in life

and did not transmit any structure of importance. The ptery-

goid sends forward a long slender process along the inner side of

the palatine which doubtless meets the prevomer as suggested

in the figure.

The shoulder-girdle and anterior limb, though lost from the

type, are fairly well preserved in the second specimen. The

scapula differs from that of Huparkeria in being very narrow in

the middle and much more expanded at the base. It is much

more Dinosaurian in appearance. The coracoid has a rather

small foramen, and the posterior border below the glenoid cavity

is deeply notched. The interelavicle is narrow and: not expanded

in front apparently. The clavicles are also slender.

The humerus measures 57°5 mm. in length. There is a well-

developed deltopectoral crest, and the general appearance of the

bone suggests a comparison with that of the Theropoda. The

radius and ulna are not very well preserved. The former

measures 47 mm. and the latter 49 mm. Only afew bones of the

manus are preserved, and these are apparently displaced. A

cast of a specimen obtained by Mr. Watson shows rather more

of the manus. There are pretty certainly three well-developed

digits, and, I think, evidence of a fourth. The carpus is badly

preserved: possibly it was imperfectly ossified.

Ihave given a restoration of the pelvis for most of which,

I think, there is clear evidence in the specimen. The pubis is

essentially similar to that of Huparkeria, but much more elongated.

It has a large single pubic foramen. There is evidently a com-

plete symphysis.

The bones of the hind foot of the type, though much displaced,

have been separated out and identified by Newton with marvellous

care, and the foot is seen to resemble that of Huparkeria pretty

closely, the main difference being that the fifth metatarsal shows

less of the peculiar Rhynchocephalian development, and the

phalanges of this, too, are feebler.

ORNITHOSUCHUS TAYLORI, sp.n. (PI. LX XVII. fig. 11.)

In 1904, Mr. G. A. Boulenger described some rather badly

preserved ‘reptilian remains from Hligin as a large example

of Ornithosuchus woodwardi. Myr. Taylor, when sending the

626 DR. R. BROOM ON

specimen to the British Museum, recognised that it was a near ally

of Ornithosuchus woodwardi, though about 24 times as large.

Mr. Boulenger discusses at some length the question whether the

specimen is merely an older example of O. woodwardi or a new

species, and comes to the conclusion that there are no other

differences than those that might be accounted for by difference

of age. My comparison of the specimens has led me to con-

clude that the animals, though allied, are at least distinct

species, and I have therefore much pleasure in naming the form

after the discoverer.

The specimen shows most of the skull. The jugal is probably

perfect but is partly hidden by matrix, and the quadrato-jugal

is also nearly perfect. The squamosal and postorbital are much

crushed, but the side view of the back half of the skull can be

restored with moderate accuracy. Most of the upper side of the

skull is preserved, but partly broken and not well displayed. A

fairly complete snout with maxilla and premaxilla of what may

perhaps be a second individual isalso preserved. And as we have

the remains of the last maxillary tooth in the first specimen, and

the corresponding tooth in the snout-fragment, we can make a

fairly complete restoration of the skull. When this is done, the

very marked differences between it and the skull of Ornithosuchus

woodwardi are apparent, ‘The arches are seen to be massive

‘instead of slender bars, and the snout relatively much more

powerful. The fact of the two animals being distinct is further

confirmed by the fact that the British Museum has recently

obtained from Elgin two new specimens—the one exactly

corresponding in size to the type of Ornithasuchus woodwardi,

and the other, so far as can be made out, agreeing in size

with the large form. In the new specimen of O. taylori part

of the palate is preserved, and it agrees essentially with that of

O. woodwardi. There are two similar openings in the back part,

and the pterygoid sends forward a similar but relatively larger

anterior process.

HERPETOSUCHUS GRANTI Newton.

This small Pseudosuchian from Elgin is less satisfactorily pre-

served than Ornithosuchus woodwardi, but there is sufficient to

show that it is a fairly near ally. The skull differs considerably

in its proportions, but probably the essential structure is similar.

The palate is narrower, and the anterior process of the pterygoid

is relatively broader, and there is only a single opening in the

pterygoid region. The shoulder-girdle and anterior limb are

differently proportioned, being much more slender, and indicate

that the habits of the two genera were different. An interesting

point revealed by this specimen is the fact that the manus has

five well-developed digits. In the carpus there are at least three

elements. The five metacarpals are well preserved, but most of

the phalanges are missing.

FOSSIL REPTILES. 627

BROWNIELLA AFRICANA, gen. et sp. n. ' (Pl. LX XIX. fig. 21.)

In Mr. Brown’s collection there are portions of at least two

skeletons of an animal rather larger than HLuparkeria capensis

but closely allied to it. The only parts well preserved are the

shoulder-girdle, pelvic girdle, and femur. These indicate an

animal nearly a half larger than the better known African

form.

The following measurements will illustrate the differences in

size of the two forms :—

Huparkeria . Browniella

capensis, africana.

F mim. mim.

Width of base of scapula ...... 15 20

enguhoh coracoidy ese eee 24 30

Pens theotsisehnumy eee. nee at ee 36 about 50

Wemethvolstomuiy ees. a aes 56 74

In general structure there is a close similarity between the

bones, but those of Browniella are more massive, and there are

numerous minor differences.

The ischium differs in shape in being constricted near its

middle, and the pubis, besides being much broader and stouter,

differs in having only a single pubic foramen. This difference

seems of sufficient importance to justify the placing of this

species In a separate genus, and I have much pleasure in naming

it after Mr. Alfred Brown, the veteran collector, to whom Science

owes such a deep debt.

MEsosucHUSs BROWNI Watson. (Pls. LXXVIII., LXXIX.

figs. 12-15, 23.)

Though Mesosuchus brown? differs considerably from Huparkeria

and is probably not a Pseudosuchian, it has nevertheless some

interesting affinities, and, both from its association with Ha-

parkeria and from the fact of the two forms having been confused,

it seems well to consider the one along with the other.

As already mentioned, Watson, in his description of MWesosuchus

browint, had regarded some of the imperfect skeletons associated

with it as belonging to the same form. As we now know that

most of these skeletons belong to the similar sized but distinctly

different Huparkeria capensis, it is necessary to redefine J/eso-

suchus to some extent.

The type specimen consist§ of a badly crushed skull showing

the premaxilla and maxilla of the left side in good condition,

with most of the lower jaw and most of the palate badly crushed.

Much of the rest of the skull is present, but so badly distorted

that a restoration is difficult. Of the rest of the skeleton of the

type individual almost the whole of the vertebral column from the

head to about the middle of the tail is preserved. Of the shoulder-

girdle there is practically nothing preserved, though much of the

Proc. Zoou. Soc.— 1913, No. XLII. A2

628 DR. R. BROOM ON

right arm is seen. The pelvis is badly preserved, but both ischia

ave well seen and most of the left hind leg.

Tie pcremaxilla is particularly interesting in having no inter-

nasal process, and in this resembling the bone in Rhynchosaurus

and Hyperodapedon. It has two well-developed acrodont teeth,

which ave round and blunt. The maxilla is long and narrow,

and has 13 irregularly arranged rounded blunt acrodont teeth.

A disarticulated quadrate, which is not that of Zuparkeria and

pretty certainly that of Mesosuchus, is nearly as broad as long,

and uch more massive than the quadrate of Huparkeria. here

are two moderately distinct condyles. Watson's description of the

palate cannot at present be added to. “ Pterygoid of remarkable

shape with a deep posterior ramus applied to the inner side of

the quadrate, small external ramus (not well exposed) and long

anterior ramus which bears a closely-set series of small pointed

teeth. Vomer apparently narrow, with a series of small poimted

teeth articulated with anterior end of pterygoid. Other bones

of palate not shown. Epipterygoid widened with a deep notch

for the optic nerve, touching the top of the deep posterior ramus

of the pterygoid. Parasphenoid very large and placed high up in

the skull.” Most of the postcranial skeleton described by Watson

are really bones of Huparkeria, while the supposed scapula is

really the ischium of Browniella africana.

The lower jaw is fairly well preserved in the type specimen.

Tt differs from the jaw of Huparkeria in having a relatively

small lateral opening and in having the part of the jaw behind

the opening larger than the dentary portion. The surangular

forms more than the upper half of the outside of the back of the

jaw. and the rest is mainly formed by the angular.

The vertebrz are not well preserved. They are of about the

same size as those of Huparkeria capensis. ‘The cervicals have

fairly long spines, and the whole neck 1s relatively longer than in

Eupurkeria—probably 11 vertebie may be cervical. Altogether

there a pest to be, as in Huparkeria, 26 pre-sacral vertebree, and

apparently 2 sacral.

The humerus, radius, and ulna are much more massive than in

Kuparkeria, but not very well preserved. The humerus measures

37 mm. in length.

The ilium differs considerabl y from that of Kuparkeria, and

resembles much more closely that of Howesia. hough imper fectly

preserved, the upper part of the ium is manifestly about twice

as deep as in Hupark ervd. ‘The ischium also differs markedly from .

that of HMuparkerva in being relatively much shorter, and having

only a short symphysis. The pubes are very badly pre esenveds

but have manifestly been much broader than i Kuparkeria,

though essentially similar in type. The illustrations given show

the specimen as preserved and the pelvis restored.

The femur, tibia, and fibula are not unlike those of Huparkeria.

The femur probably measures 49 mm. in length, and the tibia

47mm. The tarsus has the bones displaced, but 1s apparently

FOSSIL REPTILES. 629

better ossified than in Huparkeria. There are three large

elements and at least two small ones. ‘The rest of the pes is

probably of the Rhynchocephalian type.

There seem to have been no dermal ossifications in Mesosuchus.

SCLEROMOCHLUS TAYLORI Smith Woodward.

One of the most remarkable of the double-arched reptiles from

the Elgin sandstone is the little form named by Dr. Smith:

Woodward Scleromochlus taylori. Though there are preserved in

the British Museum the remains of five individuals, and though

something is known of most of the skeleton, the animal is too

small to be well preserved in the coarse sandstone, and hence our

knowledge of the detailed structure is very imperfect, Smith

Woodward’s description and restoration give practically every-

thing one can be sure about. The skull seems to be essentially

similar to that of Ornithosuchus. ;

Affinities of the Pseudosuchia.

Mr. EK. T. Newton, in describing Ornithosuchus and Herpeto-

suchus, discusses the affinities of the forms. He calls attention to

the marked resemblances of the forms to Aétosaurus, and even

thinks 1t possible —as we now know to be the fact—that détosaurus

has an infratemporal vacuity, and also discusses the relationships

with Stagonolepis and others of the Parasuchia. A comparison is

also made with the Dinosaurs Compsognathus, Anchisaurus, and

others, and with the skull resemblances in the Pterosaurian

Scyphognathus. In summing up the evidences he says:—‘ The

many points of resemblance between the Parasuchia and certain

of the forms usually included among the Dinosauria, have also

been noticed by other writers; and the difficulty of separating

the two groups is increased by a study of this new Elgin reptile,

which holds, as I think, a more intermediate position between

the two series than any form hitherto described, for although the

characters of its skull and teeth find their nearest counterpart

among the Dinosaurs, and the pelvis and limbs might belong to

either a Theropodous Dinosaur, or a’ Parasuchian, the form of

the free astragalus is more Crocodilian than Dinosaurian. While

acknowledging the ditliculty of assigning this new reptile to either

of these groups, it seems most in accordance with the facts to

lace it provisionally with the Dinosaurs.”

Mr. G. A. Boulenger, when describing the large specimen of

Ornithosuchus, criticises Newton’s conclusions. He says:—

“ Much as I admire Mr. Newton’s description of Ornithosuchus

....I1 cannot... . endorse his conclusions as to the systematic

position of the reptile.’ “ Dr. Smith Woodward was nearer the

truth when he compared it with Aéfosawrus.” Boulenger argues

that Ornithosuchus should be placed with Belodon, Stagonolepis,

and dAétosaurus in the order named by Owen, Thecodontia—a

495

630 DR. R. BROOM ON

group which agrees quite as much with the Rhynchocephalia and

the Carnivorous Dinosaurs as with the Crocodilians.

Though these two opinions seem at first sight to be at variance

they are really pretty similar. Practically, it amounts to this,

that in the Pseudosuchia we have a group of primitive reptiles

which, while they do not fit into any of the later specialised

orders, have affinities with quite a number of other groups.

There cannot, I think, be the shghtest doubt that the Pseudo-

suchia have close affinities with the Dinosaurs, or at least with

the Theropoda. This has been recognised by Marsh, v. Huene,

and others. In fact there seems to me little doubt that the

ancestral Dinosaur was a Pseudosuchian. The skulls of such types

as Huparkeria or Ornithosuchus are practically Dinosaurian even

in detail, and the skulls of the early Dinosaurs, Such as Anchi-

saurus, differ less from the skulls of Pseudosuchians than those

of the early Dinosaurs do from many of the later types. And

there is nothing in the post-cranial skeleton that is not just what

we should expect to find in the Dinosaur ancestor. The shoulder-

girdle is more primitive in retaining clavicles and interclavicle,

but these are elements which we know from the history of other

groups are very variable and readily lost. The pelvis is almost

Dinosaurian, and differs only in having the acetabulum closed.

This is an important character ; but when we consider the con-

dition in the two nearly allied Monotremes—the one with the

acetabulum closed, the other with it open—we see how easily even

this character may change. ‘The hind limb is almost Dinosaurian

in Huparkeria. The ankle is less specialised and the fifth toe is

still well developed and retains the Rhynchocephalian characters.

Huparkerva is ia my opinion potentially bipedal, and was probably

partly bipedal in its habits. The fourth toe of the hind foot is

more feebly developed than the third and the axis of the foot is

down the third toe, which would seem to indicate that the feet

were at least not so laterally placed as in lizards, and that the

animal possibly ran on its hind feet. The relative shortness of the

toes also seems to confirm this view, as well as the feebleness of

the fore limbs. I believe Huparkertia fed on some large forms of

insects like locusts, and captured them with its front feet.

Ornithosuchus was probably very similar in habit to Huparkeria

and was even a little better adapted for running on its hind feet ;

the large species O. taylor could hardly have had the same habits

as the small O. woodwardi. tis too massively built, and probably

had become largely carnivorous, adding to thelarger insects various

small vertebrates, and perhaps, like the vulture, the flesh of

animals too large for it to kill.

The aflinities of such small Pseudosuchians as Huparkeria,

Ornithosuchus, and Aétosaurus with the Belodonts through such

an intermediate form as Hrythrosuchus is very manifest. And as

Boulenger stated, the Pseudosuchians are about as near to the

Parasuchians as to the Dinosaurs. The series of dermal plates

down the back, though suggesting affinity is, however, not a

FOSSIL REPTILES. 631

character on which too much weight must be placed. As we

see in the Lacertilia, dermal ossifications are subject to great

variation in even allied forms.

Another group to which the Pseudosuchians seem to have

affinities as suggested by Newton, isthe Ornithosauria. In general

proportions the Pterodactyles differ very greatly, but the form

from which they arose must have been very much like that seen

in Pseudosuchians. ‘The Pterodactyl and Pseudosuchian skulls are

almost exactly similar in essentials. As pointed out by Newton,

theskullin Scyphognathus resembles pretty closely that of Ornitho-

suchus. The Pterodactyl manus is simply an ordinary reptilian

manus with the 5th digit lost and the 4th greatly specialised,

and there can be no doubt that the 5th digit was lost before the

wing-membrane was developed. The pelvis of the Pterodactyl

is not thoroughly known beyond doubt, but seems to be a modi-

‘fication of the ordinary plate-like type with the prepubis ossified.

Seleromochlus is a very lightly built Pseudosuchian evidently

adapted for taking leaps, and not for bipedal progression on the

ground. The limbs are long and slender, and the length and

slenderness of the fore limb suggest that possibly there was a

membrane stretched between the fore and hind limbs and perhaps

between the hind limbs and tail, which would enable the little

animal to take sustained leaps like Petauwrus.

Although Scleromochlus is already too specialised in the hind

foot structure to have been in any way ancestral to the Ptero-

dactyls, it may suggest how they have arisen, just as Galeopithecus

suggests how the bats arose.

There is still another group to which some Pseudosuchian has

probably been ancestral, namely, the Birds. For a time one or

other of the Dinosaurs was regarded as near the avian ancestor.

The resemblance of the hind limb and pelvis seemed to make this

extremely probable, and Huxley, Marsh, Cope, and others have

all favoured this view. Others, however, were more impressed by

the apparently avian characters in the skeleton of the Ptero-

dactyls, and especially in the striking avian appearances in the

brain, and have argued in favour of a close affinity between

the Birds and the Pterodactyls. Osborn, while recognising

the affinities to both groups, and especially to the Dinosaurs,

believed that the Birds and the Dinosaurs had a common ancestor,

probably in the Permian. Seven years ago, when describing the

skeletogenesis of the Ostrich, I argued that the bird had come

from a group immediately ancestral to the Theropodous Dinosaurs.

The Pseudosuchia, now that it is better known, proves to be just

such a group as is required. In those points where we find the

Dinosaur too specialised we see the Pseudosuchian still primitive

enough. The bird pelvis has probably developed from a type

like that of Ornithosuchus by the pubis turning further back and

the symphysis becoming lost. Whether the union of the meta-

tarsals is a primary or a secondary character is a debatable point.

The question is really whether the bird ancestor was a hopping

632 DR. R. BROOM ON

bipedal animal before it flew, or if it only hopped after the wing

had become specialised. Jam strongly of the opinion that it was a

hopping animal first, and that the metatarsus became strengthened

to support the weight of the body entirely borne by the hind feet.

It is easy to understand a hopping animal taking to an arboreal

life and ultimately developing a wing out of a four-toed hand,

while it seems unlikely that the hind foot could ever have

developed by arboreal habits. It is interesting to note that

while the ancestor of the Pterodactyls had four toes in the manus,

there is very clear evidence from the skeletogenesis of the bird

that the latter also had a four-toed ancestor.

A Pseudosuchian which through a bipedal habit had developed

a strengthened ankle-joint and a firm metatarsus, and had lost

the 5th “digit from the manus would meet all the requirements of

the avian ancestor.

We know at present too little to discuss the relationship of the

Pseudosuchians with Sphenodon and with Gnathodonts, nor can

we say whether Howesia and Jesosuchus should be placed with

the Pseudosuchians. ‘There is evidence of a generalised Permian

Rbynechocephaloid order which gave rise to the more specialised

‘Triassic groups, but at present we know too few forms and very

few even of these are well known, and until our knowledge has

much advanced it seems unwise to attempt any further classifi-

cation. In South Africa we can trace through the Lower Triassic

and Upper and Middle Permian beds forms that may be ancestral

to the Pseudosuchians, and when these are better known a

satisfactory classification will be possible.

Tam much indebted to Mr. EH. 8. C. Dyke for the photographs

of Kuparkeria.

References to Literature.

(1) E. T. Newron.—“ Reptiles from the Elgin Sandstone.

Description of two New Genera.” Phil. Trans. vol. 185,

1894, p. 573.

(2) G. A. Boutencer.—“ On Reptilian Remains from the Trias

of Elgin.” Phil. Trans. vol. 1968, 1903, p. 175

(3) A. 8. Woopwarp.—“‘ On a new Dinosaurian Reptile (Sclero-

mochlus taylori) from the Trias of Lossiemouth, Elgin.”

Owe Gasavolsixan, W907.) o. VAs

(4) F. v. Huzne.—“ Die Dinosaurier der Europiischen Tyrias-

formation.” Geol. u. Pal. Abhand., Jena, 1907.

(5) D. M. 8. Warson.—* Mesosuchus panne. gen. et spec. nov.”

Rec. Albany Mus. vol. il. pt. iv. p. 296.

(6) R. Broom.—‘ Note on Mesosuchus brownt Watson, and on

a new South-African Triassic Pseudosuchian (Huparkeria

capensis).” Ree. Albany Mus. vol. 11. pt. v. 1913, p. 394.

A full list of other papers will be found in y. Huene’s work.

FOSSIL REPTILES: 633

EXPLANATION OF THE PLATES.

Ang. Angular. B.o. Basioccipital. Cl. Clavicle. Co. Coracoid, D. Dentale,

Fr. Frontal. JZ.cl. Interclavicle. J.P. Interparietal. Ju. Jugal, 2. Lacryinal,

Mx. Maxilla. Wa. Nasal. Pa. Parietal. Pa.o. Paroccipital. Pma. Premaxilla.

Po.F. Posttrontal. Po.O. Postorbital. Pr.F. Prefrontal. Q. Quadrate. @./,

Quadrato-jugal. S.Ang. Surangular. Se. Scapula, 8,0, Supraoccipital. Sy,

Squamosal. Sé. Sternum.

Pyatr LXXV,

Kuparkeria capensis,

. Side view of type specimen. 4 nat. size.

Top of skull of type specimen. Nat, size,

. Left dentary of a second specimen. Nat. size.

Right hind foot of this second specimen, Nat. size,

Fic.

02 bo ju

PratE LXXVI.

Euparkeria capensis,

. Side view of skull, Nat. size.

. Upper view of skuil.

. The quadvate viewed obliquely from behind, showing the relations to the

adjoming bones. Simall portions of the squamosal and opisthotic are

broken off.

8. Shoulder-girdle, slightly restored and with the probable cartilaginous sternum

added in dots. Nat. size.

“TD on

Prave LXXVIT,

Fig. 9. Side view of skull of Ornithosuchus woodwardi. Nearly nat, size.

10. Under view of skull of Ornithosuchus woodwardi.. Nearly nat. size,

11. Side view of skull of Ornithosuchus taylori. About 2 nat. size. Restored

trom the type specimens.

Prate LXXVIII.

Fig. 12. Side view of imperfect skull of Mesosuchus browni. Nat, size.

13. lnpression of the teeth of the right maxilla of Mesosuchus browni. Nat,

size.

14, Fragment of ilium, and left hind limb of Wesosuchus browni, Nat. size.

15. Fragmentary pelvis of Wesosuchus browni. Nat. size.

16. Shoulder-girdle and fore lnnb of Ornithosuchus woodwardi. About + nat,

size. Slightly restored.

17. Top of lett ihum of Huparkeria capensis. Nat. size.

18. Acetabular portion of left illum of Huparkeria capensis. Nat, size,

19. Portions of three lower dorsal ribs of Huparkeria capensis showing unci-

nates. Nat. size,

20, Right temur, tibia, and fibula of Huparkeria capensis. Nat, size,

Pratt LXXTX,

. Right pubis, left ischium, portions of both ilia and of two dorsal vertebra

ot Browniella africana, Nat. size.

. Right pubis of Huparkeria capensis. Nat, size,

. Pelvis of Mesosuchus browni. Nat. size.

|. Pelvis of Huparkeria capensis. Nat. size.

. Pelvis of Ornithosuchus woodwardi. About > nat, size,

. Pelvis of Gryponyx africanus, Much reduced,

i>)

to bw br

Boe wr

634 MR. G. E. BULLEN ON

41. On some Cases of Blindness in Marine Fishes.

By G. H. Butien, the Hertfordshire Museum, St. Albans *.

[Received April 24, 1913: Read May 20, 1913. ]

Of recent years considerable attention has been given by

numerous ichthyopathologists and fish-culturists to the sporozoic

and bacterial diseases affecting those species of fish which in a

live state form an article of commerce.

The important work of Hofer+, and the Monograph now

appearing from the pen of de Drouin de Bouvillet, are perhaps

two of the best known contributions of a general character

upon the subject. There are, moreover, numerous more or less

important works upon those organisms giving rise to disease: the

Myxosporidia, for example, have received considerable notice

from such investigators as Balbiani§, Thelohan |), Doflem 4,

and Gurley **, to quote but a few of the authors upon the

subject.

Work which has been performed, even within the past decade,

upon the bacterial aspect of fish diseases is of too extensive a

character to be cited, however brietly, here. The same applies

to Saprophytic and other fungoid growths attendant upon

bacterial lesions.

The investigations of J. Hulme Patterson and certain others

upon Salmon disease are well known and important contributions

to economic science in this respect. But with the exception

of certain papers by G. H. Drew and Patterson’s findings, so

far as they apply to Salmon installed in sea-water, most of

the literature dealing with disease in marine fishes is confined

almost entirely to the cecology of pathogenic organisms. For,

whereas there is no inconsiderable amount of the literature

directed solely to the pathology of freshwater fishes, there is

at present a remarkable paucity dealing with those of marine

habitat.

The present paper, therefore, is put forward largely with a

view to associating the pathological aspect of certain forms of

eye diseases observed in marine fishes with those already

described, in more or less general terms, in the case of freshwater

species.

* Communicated by Joun Hopkinson, F.LS., F.Z.S.

+ ‘Handbuch der Fischkrankheiten.’

t+ ‘Les Maladies des Poissons d’eau douce d’ Europe.’

§ ‘ Lecons sur les Sporozoaires,’ Paris, 1884, etc.

|| “ Recherches sur les Myxosporidies,” Bull. Soc. France et Belgique, ser. 4, vol. v.

Paris 1894. ete.

4 ‘Die Protozoen als Parasiten und Krankheit Serrigers,’ Jena, 1901. “Studien

zur Naturgeschichte der Protozoen,” Zoologische Jahrb. xi. 1889, ete.

** “ Myxosporidia, ete.” Rep. U.S. Commissioner for Fish and Fisheries, 1892.

Washington, 1894. ;

BLINDNESS IN MARINE FISHES. 635

The several subjects described hereinafter were received by

the writer from the Superintendent of the Brighton Marine

Aquarium, Mr. EK. W. Cowley, to whom thanks are due.

Acknowledgment of much assistance in the preparation of this

paper is also accorded to Mr. John Hopkinson and to the late

Mr. G. Harold Drew.

Hofer * traces the progress of the more usual form of blindness

in freshwater fishes in the following manner: (a) Corneal

“cataract” due to injury or pathogenic bacteria attacking the

integument; (3) gradual necrosis of the cornea-epithel and

outer surface of the cornea; (vy) perforation of the cornea.

Complications which may secitte ave “ Keratoglobus” in the

earlier stages, and Saprophytic growth attendant upon the

bacterial lesions.

In the case of several freshwater fishes, the same author

describes a not uncommon form of blindness due to parasites

(Diplostomum larve), but of all the marine fishes which herein

receive notice, none has been affected in this manner. More-

over, it may be recorded that no evidence of Saprophytic growth

occurred on even the worst examples.

1. Traumatic Corneal Opacitis in a Conger Eel

Conger vulgaris).

fo) 5

The subject of this note was a Conger Kel which measured

3 feet 7 inches. It had lived in the Aquarium for nearly six

years, retaining complete health until November 1912, when,

according to the Superintendent of the Aquarium, Mr. E. W.

Cowley, it went completely blind within the space of a week.

It was destroyed immediately, when, from an observation of its

behaviour, both eyes appeared to have become wholly useless.

An examination of the head of this specimen showed that the

right cornea-epithel was perforated in two places and the left in

five, each perforation being just large enough to admit a seeker

-5 mm. in diameter. Extensive penetration of sea-water followed

probably by bacterial lesions had occurred throughout the under

surface of the cornea-epithel, those on the left side extending to

the adjacent epidermis of the gill-covers, destroying the colour-

cells, and rendering the part affected opaque white. The eye

itself, when superficially examined, was so completely dimmed

as to render observation of the characteristic yellow iris difficult,

except in strong light.

The term ‘corneal opacitis” would be somewhat loosely

applied in this instance, since upon removal of the integument

the cornea proper was found to be completely free from injury,

and but slightly affeeted with opacitis.

* Op. cit. p. 291 etc.

+ For a determination of this fact the writer’s thanks are due to Miss Lorraine

Smith, of the British Museum.

636 MR. G. E. BULLEN ON

The condition termed by Hofer * ‘ Keratoglobus,” 7. e. out-

ward bulging of the cornea away from the pupil, did not affect

either eye, the lenses retaining their normal position.

Blindness, therefore, in this instance was due apparently to

mortification and bacterial lesions of the integument of the head

passing over the eye attendant upon perforation of the cornea-

epithel.

Hofer? remarks: “In the case of a fish weakened by disease,

or in a dying condition, the organs of sight are deeply sunk

in their orbits, whereas in a healthy fish the eyes somewhat

protrude.” In the present example this condition was wholly

apparent, the eyes presenting the flabby appearance to be seen in

a fish some hours after death.

It is highly probable, therefore, that perforation of the integu-

ment above the cornea, resulting in blindness, was largely due,

in the present instance, to the susceptibility of the fish, alr eady

weakened by graver organic disease, to injury ; and that incipient

corneal opacitis may sometimes be taken as a diagnostic of other

disorders.

Corneal Opacitis in a Greater Weever

(Trachinus draco).

The chief difference between the condition of the eyes in the

present example and that of the Conger Hel already described,

lay in the fact that apparently no lesion of the cornea-epithel

appeared to have taken place. The subject under consideration

had lived for some years in the tanks at Brighton, and had

attained the considerable measurement of 15? inches. In this

case partial blindness only had come about at the time when the

specimen was examined. The left eye was less affected than

the right, this showing a more or less evenly distributed film of

dimness, as if the specimen had been immersed for a short time

in weak formalin. The right eye, however, not only showed

distinct spots of necrosis unevenly distributed throughout the

surface of the cornea, but there was also a pronounced “ kerato-

globe.” With reference to this latter feature, it may here be

remarked that a gathering of aqueous humour in front of the

pupil, causing the cornea to bulge outward to a considerable

extent, may occur, doubtless as the result of concussion, in fish

the eyes of which are otherwise apparently unaffected. At the

Brighton Aquarium occasionally numbers of a shoal of mackerel

may be observed with both eyes protruding in this way, but

their undoubted lack of effectual vision appears in no way to

interfere with their capability for maintaining position in a

moving shoal. But it is a noticeable fact that they appear

much thinner than their fellows consequent upon a more limited

* Op. cit. p. 293.

+ Op. cit. p. 290. The translations adhere as nearly as possible to the original

text.

BLINDNESS IN MARINE FISHES. 637

absorption of food. Not unfrequently, moreover, such examples

present a certain broken-backed appearance, which, so it is stated

by the attendant in charge of the tanks, is due to the fish, when

first installed, persistently hammering themselves against the

glass!

3. Corneal Opacitis and Cataract in Pollack

(Gadus pollachius),

The Pollack forming the subject of the present note were four

young specimens ranging from 74 to 83 inches in length. At

the time when they came under the writer’s observation (in Sept.

1912) they had lived in the Aquarium, in a tank apart from

other fish, for a period of about nine months. When first

installed their sight appeared to be normal; but it seems that

these particular fish had been taken on a position close inshore

off Shorehain, where there is an outflow of chemical waste.

Mr. E. W. Cowley informed the writer that fish which were

known to have come from this locality were as a rule refused for

stock purposes, in view of the fact that, after a short period in

the Aquarium, they invariably showed signs of disease. Hofer *

states : “ Trout, etc., have been fr equently, attacked by perforation

of the cornea and cataract through existing in waters impreg-

nated with iron from drains”; and goes on to say, * It is not yet

established whether these consequences are to be ascribed to

iron salts in general or to any particular one.” A lack of

similar observation, viz.: the effect of industrial wastes,

sewerage, etc., upon those marine fishes often frequenting

estuaries and tracts of water in which such impurities are known

to oceur, prohibits any comparison. Yet it is a noticeable fact

that the examples about to be described exhibit a form of

blindness most frequently met with in various freshwater fishes

(e. g. Pike, Perch, Carp, ete.).

At the time when the present specimens came under the

writer's observation, they were without doubt totally blind, both

eyeballs of each example presenting the opaque white appearance

similar to that seen in the case of fish which have been boiled.

The cornea in every case was but slightly affected, no apparent

lesion being discer nible on any part of the integument covering

aiveameine general transparency, moreover, was consider: ably gr eater

than it was in the case of either of the two preceding examples.

On dissection it was found that the cornea protruded in front of

the lens to a considerable extent (cir. 24 mm.). A transverse

section, taken through the centres of both eyeballs from two of

the specimens, showed that the extent of necrosis attacking the

interlocking fibres was practically the same, viz., equivalent to

two-thirds of the entire diameter of the section. Vertical

sections, ¢. é. from anterior to posterior aspect of the lens, showed

*Opcitaps 201.

638 ON BLINDNESS IN MARINE FISHES.

a slight increase on the anterior margin, the total extent, how-

ever, being slightly less than that observed in the transverse

sections, amounting roughly to slightly less than half the diameter

in affected tissue. The term “cortical cataract,” as applied to

the human subject, would perhaps aptly describe the condition

observed when in its earlier stages *.

The behaviour of these fish in the Aquarium is a matter

deserving of some comment. Habitually they were observed to

swim in mid-water, maintaining one position often for several

minutes together, and seldom exhibiting much activity in move-

ment. It is a somewhat remarkable fact, but one which the

present writer, by an observation extending over several weeks,

can personally vouch for, that on no occasion were any of the

specimens seen to come in contact with each other, the sides

of the tank, or the glass. Pollack with normal vision installed

In an adjacent tank were easily attracted when a moistened

finger-tip was rubbed rapidly up and down the glass causing

vibration thereby, less freely by gentle tapping. The blind fish,

even when situated within a few inches from the glass, made no

effort to respond to this attraction.

When food, in the form of finely-cut fish, molluscs, ete.,

was offered, it was allowed to sink to the bottom and remain

there for several minutes before a search after it was made.

This took the form of a wholly haphazard grubbing about on the

bottom, the fish often making a vigorous “ bite” immediately

beside a fragment, this operation being repeated until the morsel

was secured. Although a sufficiency of food was placed in the

tank daily, and on one occasion one individual fish was observed

within the space of 15 minutes to secure three fragments of

moderate size, all the specimens when examined post-mortem were

found to be very ill-nourished and considerably below normal

weight.

Nore :—The writer is indebted to Mr. J. O. G. Ledingham, of

the Lister Institute of Preventive Medicine, for the following

report respecting the histological condition of the lens in one of

the above specimens. This gentleman writes “I cut Celloidin

sections of the fish’s eye you sent to me in formalin. They were

very difficult to cut owing to the fact that the lens proved to be

completely calcareous. This was the only obvious lesion I could

detect. I saw no micro-organisms in the sections; but, as you

will understand, the sections were not satisfactory for the

demonstration of micro-organisms.”

* Vide L. B. Harman, “ Aids to Ophthalmology,” p. 88.

STR ARTHUR H. CHURCH ON TURACIN. 639

42, Notes on Turacin and the Turacin-Bearers.

By Sir Artnur H. Cuurcu, K.C.V.O., F.R.S.*

[Received May 26, 1913: Read June 3, 1913.]

In bringing, at the suggestion of the Secretary, these notes

before the Zoological Society I have a threefold object.: First,

to give a summary history of our knowledge of the occurrence in

the Plantain-eaters of turacin, the remarkable (1 may say, the

unique) decorative pigment which is distinctive of the three genera

Musophaga, Turacus, and Gallirex ; secondly, to describe the

properties of the pigment; and, thirdly, to correct some prevalent

errors on the subject of turacin—errors due in part to initial

mistakes made by observers, and in part to incorrectness in the

transmission cf facts by copyists.

Although the earliest observation as to the occurrence in

certain Turacoes of a singular red pigment seems to be assignable

to the year 1818, yet I have not found any publication of the fact

by the original observer until more than half a century later.

Then, on January 17th, 1871, a Corresponding Member of this

Society, M. Jules Verreaux, described at a meeting his observa-

tions and experiments on the subject, referring them back to

the year 1818. Verreaux’s remarks are to be found in the

‘Proceedings of the Zoological Society’ for 1871 (pp. 40 eé seq.);

they refer to the Corythaix albicristata of Strickland. Many of

these Turacoes were met with in the Knysna district of the Cape

of Good Hope, and were carefully observed by Verreaux. He

noticed that, in the torrential rains that prevailed at the time,

these birds left the tree-tops and sought refuge in the dense lower

branches. One bird, which he had seized by the wing, escaped,

and he says that great was his surprise to see the inside of his

hand coloured a blood-red. Some days later the experiment was

repeated with three more specimens caught in a soaked condition ;

it was easy to remove the pigment from the feathers by friction

and to reduce them to a pale rose hue. What M. Verreaux here

adds to his observation as to the solubility of the red pigment in

rain-water is, | venture to think, hard to believe. He wrote in

French, and I give his own words :—‘‘ Mais ce qui nous surprit

le plus, fut de voir cette méme coloration rouge vif revenir des

que loiseau était complétement séché. Nous avons renouvelé

cette opération deux fois par jour, et chaque fois nous avons eu

le méme résultat.’” M. Verreaux further extends his observa-

tions to other species of Turacoes, finding in them the same

colouring-matter, possessed of the same mobility and of the same

power of renewal when the feathers became dry. He even

affirms such recovery of colour in the case of birds that have been

killed as well as of those that have been captured. One cannot

* Communicated by the SrecRETARY.

640 SIR ARTHUR H. CHURGH ON TURACIN.

but ask, how is it possible for a mature feather to be again

supplied with this localized red pigment after it has lost its

original charge and after the vessels in the quill have dried up,

their function having been fulfilled? The opinion of VY. Fatio,

as to an oil making its way up the mature feather and dissolving

and distributing pigmentary deposits already present, does not

apply to turacin which is insoluble in oily media. As to the case

now being considered, [ have not been able to obtain any con-

firmation of this renewal of the pigment (apart from a moult)

from any trustworthy naturalist. The late J. J. Monteiro, the

late Dr. Benjamin Hinde (of Bathurst on the Gambia), and many

other competent observers of these birds in their native haunts

and in captivity have expressed themselves strongly on this

point. It may be admitted that in the folded pion which has

had its pigment moved by soaking, some of the red solution may

travel from the feathers beneath to the surface as it dries; thus

we should have a case of transference of pigment, not of its

renewal.

Amongst those who, after Verreaux, have observed this want

of fixity in the red pigment of the Turaco I may name the late

Rowland Ward, the late W. B. Tegetmeier, also J. J. Monteiro

and Dr. B. Hinde.. The last-named, in a letter dated May 1865,

said the moment soap touches the feathers the colour runs,

although it is difficult of extraction by pure water. But he

added ‘“‘ the birds which I sent home washed themselves neazly

white in the water given them to drink.” In my memoir of

1869, I mentioned Mr. Tegetmeier as having introduced the

subject now being discussed to my notice. His own attention

had been drawn to the matter by a correspondent of ‘ The Field,

who sent him a washed-out feather. Mr. Tegetmeier forwarded

this feather to me and asked me to ascertain if there were any

possibility of its having been dyed.

Such, in brief, was the position of the problem when in 1866 I

began my researches. The chief results of these were published

in the Phil. Trans. of 1869 and 1892, while im 1894 a Friday

Evening Discourse in the Royal Institution presented a digest of

the whole enquiry *. Although it would be unbecoming in me,

a chemist, to enter into classificatory questions before a company

of zoologists, I may venture to remind you that, according to

many systematists, the Order Cuculiformes includes two suborders

of equal rank, the Cuculide or Cuckoos and the Musophagide or

Plantain-eaters. The former is an extensive suborder and

cosmopolitan in range, while the latter contains less than 40 species

and is confined to the Ethiopic region of Continental Africa. Of

* <“Turacin, a new Animal Pigment containing Copper,’ ‘Student,’ i. (1868)

pp- 161-168; with a coloured plate.

“Researches on Turacin, an Animal Pigment containing Copper,”

clix. (1869) pp. 627-636 (with 4 figs. of spectra).

“Researches on Turacin, an Animal Pigment containing Copper,’ Part IL.,

Phil. Trans. clxxxii. A. (1892) pp. 511-530 (with 9 figs. of spectra).

Proc. Roy. Inst. xiv. (1894) pp. 44-49.

hil. Trans.

SIR ARTHUR H. CHURCH ON TURACIN. 641

the six genera of Musophagide three contain the crimson pigment

which I named turacin. These are:—WMusophaga, 2; 7 Ur aCUS,

21; Gallirex, 3; or 26 turacin-bearers in all. The ‘oe ee remain-

ing genera from which turacin is absent are :—Corytheola, 1 ;

Chizerhis, 4; and Gymnoschizorhis, 2; or 7 in all. There are

two remarks that may be interposed in this connection. The first

is that the crimson pigment is identical in all the species, and

occurs not only in from 6 to 18 of the primary and secondary

pinion-feathers, but also in all other similarly-coloured feathers

or parts of feathers which are found on some of these birds—for

example, in the head-feathers of Mwsophaga violacea, in the crest-

feathers of Zuracus donaldsowi, 7. fischeri, and 7’. meriani, and

in the head-feathers of Gallivex johnstoni behind the crest. The

second remark refers to the pinion-feathers of three out of four

species of Chizerhis, which have white patches bare of any pig-

ment pretty much in the same position as the crimson patches in

the turacin-bearers, although mostly confined to one side of each

shaft—a curious coincidence, at all events.

I may now turn to the properties of turacin. We havealready

seen that it is soluble in water. Not, we may add, in hard water,

but in clean rain water or, better, in distilled water. It is still

more easily dissolved by weak alkaline liquids, extremely dilute

ammonia being the best solvent. From this crimson solution, the

colouring-matter, the turacin, is precipitated as gelatinous flocks

on neuti: alization by hydrochloric or other acid. The flocks that

separate when collected and dried form a deep red amorphous

mass, crimson by transmitted light when in thin layers, but

exhibiting a surface-lustre of a purplish hue, not unlike that

reflected from erystals of potassium permanganate. Thus the

appearance of solid turacin cannot be said to be accurately de-

scribed when it is spoken of as ‘“‘a metallic red or blue powder,” ” as

in the Enc. Brit. 11th ed. vol. x. p 226a@. Turacin is insoluble

in alcohol, ether, chloroform, petroleum-spirit, benzol, and the

usual solvents of resins and oils. In order to obtain it in a pure

state special procedure is required to prevent its contamination

with the natural oil of the feathers, while it is not possible to free

it completely from all traces of non-essential mineral matter.

The action of heat on dry turacin presents several points of

interest. It suffers no change at a temperature rather above that

of boiling-water, but at or near the boiling-point of mercury it is

pr ofoundly modified without loss of w eight and becomes insoluble

in alkaline liquids. If this altered ‘yu acin be now exposed to a

high temperature in the presence of air its combustible con-

stituents burn away, leaving a black ash, amounting in the purest

samples to about 9°8 per cent. of the original weight. This ash

consists almost entirely of oxide of copper, the amount corre-

sponding to at least 7 per cent. of that metal in the turacin itself.

Here again the statement in the Ene. Brit. (Joc. cit.) needs correc-

tion, for this is a fixed percentage, not one that varies from “5 to

8.” In further describing the action of heat upon dry turacin it

642 SIR ARTHUR H. CHURCH ON TURACIN.

must be mentioned that this colouring-matter, when suddenly and

strongly heated, yields a volatile, red, copper-containing derivative,

which, undissolved by weak ammonia-water, is not only soluble

in, but may be crystallized from, ether. It is the vapour of this

substance catching fire which gives rise to the green flame so con-

spicuous when a particle of turacin is heated in theair. Hf I

were discussing the nature and relationships of turacin from a

chemical view-point, much would have to be brought forward as

to its composition and probable formula. Here let it suffice to

say that analysis gives these percentage-numbers :——

Warsbomireren.- Pre th 53°69

JalyGleoyxeinl jcaanaeesAnaooncoc 4-60

CIS} 0) SETS: 5 AAA RRA (Ou

INTMUROPYEIM yo nqanossenecesG50 6:96

Osxaime men iien a saceeeeeeeee 27°74

These figures, though deduced from many careful determina-

tions, do not lead unmistakably to one definite empirical formula.

With a colloidal compound like turacin, which does not admit of

purification by distillation or crystallization, there is always the

chance of a disturbing factor being present in the shape of a trace

of some impurity. Possibly such an expression as C,,H,,.N CuO,,

is worth suggesting. In this the ratio of nitrogen to metal is as

4:1, the same ratio which occurs in hematin from blood between

nitrogen and iron, and between nitrogen and magnesium in sore

chlorophyll constituents and derivatives. With all these bodies

turacin is also brought into relation when its absorption-spectrum

is considered, especially in regard to that broad band in the violet

and ultra-violet (between the lines 4 and M) which is common to

all these colouring-matters, and which was figured and described

by the late Prof. Arthur Gamgee in a paper read before the Royal

Society in 1896—a paper in which the author confirms my results,

especially those relating to the discovery of turacoporphyrin.

The spectrum of turacin may now engage our attention: in

the feather itself the pigmented web shows two broad bands, one

with its centre at wave-length at or near 585, the other with its

centre near 538. <A solution in water containing a trace of

ammonia shows the same two bands somewhat shifted towards

the violet end of the spectrum; there is also seen a very faint

band about the solar line F, but nearly as broad as band No. 2.

When turacin which has been precipitated from its alkaline

solution by strong hydrochloric acid is again dissolved in weak

ammonia-water, and the solution spectroscopically examined, a

narrow and faint fourth band situated on the less refrangible side

of D makes its appearance. It is almost certain that this band is

due to an alteration-product of the original turacin. The two

well-marked bands, as they are shown when examining a feather

with the spectroscope, closely resemble those of the oxy-hemo-

globin of the blood; while the corresponding bands seen in the

SIR ARTHUR H. CHURCH ON TURACIN. 643

spectrum of an alkaline solution of turacin are not unlike those

of CO-hemoglobin. However, the absorption-spectrum of turacin

is sufficiently characteristic to enable one to use it as a criterion

of the presence of this pigment in the red feathers of birds. So

when Dr. C. F. W. Krukenberg announced the occurrence of

turacin in a species not belonging to the Musophagide, and not

even African, it became necessary to test the assertion. The bird

in question is a Cuckoo from the Philippines, Dasylophus super-

ciliosus. I obtained a skin, then an assurance of identity from

the Bird Department of the British Museum, I removed the

sparse red feathers from the head of this Cuckoo and submitted

them to spectroscopic scrutiny. They did show an absorption-

band, but it was situated midway between the two bands of

turacin. Moreover, the colouring-matter, which is orange-red, not

crimson, cannot be extracted by ammonia and contains no copper.

Hitherto, therefore, turacin has not been found to occur outside

the Musophagide. It is right to mention here that Dr. Kruken-

berg’s statement as to its occurrence in a Dasylophus was made in

consequence of a report addressed to him at his request by the

authorities of the Senckenberg Museum in Frankfurt, and was

not the result of an experiment made by himself*. It has

unfortunately found its way into various articles and books, e. g.

Ene. Brit. x. p. 226 a, where we read ‘‘ But turacin is not, as was

supposed, confined to the feathers of the Plantain-eaters, since it

has been obtained from a Cuckoo, Dasylophus superciliosus.” 1

have been able to secure a threefold disproof of this statement,

for although the occurrence of this pigment in any bird other than

a Plantain-eater seemed unlikely, the closely-allied Cuckoos might

have supplied an instance.

The occurrence of so large a percentage of copper as 7 in

isolated turacin needs a word of comment. It does not imply a

large amount of this metal in the plumage of a single bird; a

high estimate gives less than 14 of a grain of the metal. And it

must be remembered that many recent analyses of vegetable

and animal matters show the wide distribution of copper in both

kingdoms. Two of the latest researches in this direction were

made by Mr. J. W. Dougal7y in 1911 and by M. B. Guérithault

in 1912. Im fact, there is no difficulty in accounting for the

presence of copper in Turacoes and for its amount.

I purposely exclude from the present paper any discussion of

the curious green pigment named turacoverdin by Krukenberg ;

of the relationships between hematoporphyrin and the turaco-

porphyrin obtained from turacin by the action of acids; and of

the supposed synthesis of turacin by treating hematoporphyrin

with an ammoniacal copper solution.

* ©. F. W. Krukenberg, Vergleichend-Physiologische Vortriige (1886), Bd. i.

p. 152. f

+ Pharm. J. 1911 (4) xxxii. pp. 405-7.

+ Bull. Sci. Pharmacol. 1912, xviii. pp, 633-639.

Proc. Zoou. Soc.—1913, No. X LILI. 4

644 DR. P. CHALMERS MITCHELL ON THE

43. Observations on the Anatomy of the Shoe-bill (Baleni-

ceps rex) and allied Birds. By P. CHatmers MircHett,

M.A., D.Sc., LL.D., F.R.S., Secretary of the Society.

[Received and Read June 3, 1913. ]

(Plates LX XX.-LXXXIII.* and Text-figures 119-132.)

INDEX.

Anatomy : Page

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Syistematicucnaieskonn «suis Cie eae aes ele aS

In the end of June, 1912, a young example of the Shoe-bill

(Baleniceps rex) arrived at the Gardens as a gift from the

Sirdar, Sir Reginald Wingate, G.C.V.O., Honorary Member of

the Society. It was not quite adult, but stood between three

and a half and four feet in height. It was placed in a warmed

enclosure communicating with a grassy paddock containing

a small pond, and it fed well, although its appetite was small,

on fish, frogs and pieces of meat. It appeared to thrive through

the winter, but in the end of March, 1913, it showed difficulty

in breathing. It was removed to the Sanatorium and kept for

a few days in a warm, moist atmosphere, but died. The post

mortem examination showed the presence of mycosis in the

lungs, although not in sufficient quantity to cause mechanical

obstruction to respiration, and the interior of the larynx and

bronchi was nearly free, but subsequent microscopic examination

by Mr. Plimmer, F.R.8., the Society’s pathologist, revealed

extensive infiltration of the tissues by the mould. As Dr.

Beddard, the Society’s Prosector, was engaged on other work, I

gladly took the opportunity of making some observations on the

anatomy of this rare bird, and give the results here. My work

was to a certain extent limited by the necessity of not injuring

the skeleton, which was destined for the British Museum

(Natural History). Certain portions of the viscera were sent to

the Museum of the Royal College of Surgeons.

T am indebted to my colleague Mr. D. Seth-Smith, the Society’s

Curator of Birds, for leave to reproduce on Plate LX XX. the

excellent photograph of this bird, which was an immature male.

Pterylosis—A. D. Bartlett (1) discovered and described the

powder-down patches, and Professor Giebel (21) has written a

* For explanation of the Plates see p. 708.

Bale & Danielsson, I*4 imp, D, Seth- Smith, photo:

BALANICEPS REX.

By Zs Se US, Teil, ILOOR

D. Seth- Smith, photo. Bale & Damelsson, Lt? amp,

SKULLS OF: 1, BALA‘NICI

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vik Fetal

P, Zo So Is, Il, ILOoOX IU

D. Seth- Smith, photo, Bale & Damelsson, Lh‘ imp.

SKULLS! OF: 1, DISSURA. 2. TANTALUS. 3, CANCROMA:

een

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, Sa aaie mn

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PP 4. S&S, UNS, Wl ILecoui.

D. Seth- Smith, photo.

Bale & Danielsson, Lt? imp.

SIMUILLG Ors 1, NRDIBA, 2, IPISWECANIOS., Si, IALOUUS-

OO.

ANATOMY OF THE SHOE-BILL. 645

careful account of the external characters and pterylosis, but

fortunately I finished my examination and notes of the pterylosis

before having the opportunity of consulting Professor Giebel’s

paper, and find that my observations and inferences follow his

very closely.

The down feathers are distributed practically uniformly over

the pteryle and the apteria as in Steganopods and Storks, where-

as in Scopus and Herons they are found only on the apteria.

The oil-gland bas a tuft of feathers as in nearly all the members

of the group. The contour feathers have a small aftershaft ;

this structure is present in Herons and Scopus, variable and

sometimes absent in Storks.

The feathered tracts generally are in marked contrast with those

of Herons and resemble those of Storks in being relatively wide

and fading off into the apteria, those in Herons being remarkably

narrow and very sharply defined at their lateral and posterior

edges. The neck is continuously feathered for almost the whole

of its length, and shows no trace of the long lateral apteria

characteristic of Herons and present in Scopus. Far back, near

the base of the neck, there is a lateral space at each side. ‘The

spinal tract forks between the shoulders, leaving two parallel bands

of feathers continuous with tne neck area, and separated by a

wide median apterion. ‘These dorsal bands are wider than those

of Herons and cease about the level of the axilla. Parallel with

them at each side is a very broad and strong humeral tract, wider

even than in Storks and much wider than in Herons. ‘The dorsal

tracts reappear again as a narrow diamond-shaped band, weaker

in the middle line, and stretching between the powder-down

patches to the oil-gland. The powder-down patch at each side is

an enormous, elongated oval area extending from a little way

behind the axilla to the rectrices, and reaching some way down

the side of the ramp. ‘The patches are separated in the middle

line. The feathers are long, very thickly set, and break down

into a yellowish powder. When the bird had been in spirit, they

formed unpleasant, greasy masses. ‘Their discovery led Bartlett

to associate Baleniceps with Herons as opposed to Storks. As,

however, there are only two patches in Baleniceps as compared

with four to six in Herons generally, eight in Cancroma,

as they are absent in Scopus and exist in many other birds,

e.g. Hurypyga, Rhinochetus, Mesites, some Tinamus, Birds of

Prey, Parrots, and Goatsuckers, it is difficult to attach any

systematic weight to their presence. Nitzsch’s observation that

they may disappear with age (e. g. Gypaétus) still further destroys

their systematic importance.

The femoral tracts bordering the powder-down patches are

rather badly defined. The arrangement on the ventral surface is

almost identical with that given by Nitzsch for Ciconia. The

median apterion is rather indefinite except at the base of the

neck and near the vent. On each side of it is a very broad

feathered tract.

43*

646 DR, P. CHALMERS MITCHELL ON THE

The rectrices are 12 in number as compared with from 8 to 12

in Herons, and 12 in Scopus and Storks.

The primary quills of the wing are 11, not 10 as Giebel states.

Of these six are on the metacarpal, three on phalanx 1 of digit

JI and two on phalanx 2 of digit I, the last bemg much smaller

than the others. There are 11 in Herons, 10 in Scopus and 11 or

12 in Storks. There are 19 secondaries as compared with 11 to

18 in Herons and 14 to 25 in Storks.

Text-fig. 119.

Wing-stracture of Baleniceps.

Diagram of the distal secondary quills and coverts, showing the diastataxic condition.

Right wing; external view. The quills are in outline, the major coverts are

shaded ; the transverse rows are represented by dots, showing the insertion.

8. First Secondary. C.R. Carpal remex.

aw. Diastataxic gap. P. First Primary.

C.C. Carpal covert.

In the figure (text-fig. 119) I give a diagram of the distal

secondaries, showing the arrangement known as “ aquintocubital ”

until W. P. Pycraft and I showed simultaneously (28, 36) that it

was not due to the loss of a secondary feather, after which my term

“ diastataxic ” has been used. The major coverts are inserted

proximally to the quills and cross over them, and this arrange-

ment is repeated in the case of the degenerate carpal covert and

carpal remex, whereas the major coverts of the primary quills

are distally placed. ‘The diastataxic gap is very evident and the

major covert, which occupies the gap, is tied down to the

membrane supporting the quills by a special slip. In the same

fashion the carpal remex has a membranous slip supporting it.

ANATOMY OF THE SHOE-BILL. 647

There is nothing peculiar in this arrangement; the diagram

resembles closely the similar diagrams that I have given for

other diastataxie birds (¢.g., 32, fig. 23). There is some difference

of opinion as to whether the secondary quills, major and minor

coverts, and the feathers nearer the outer border of the wing

represent horizontal or transverse series. Most writers, following

the obvious lead given by the quills and major coverts, have

preferred to regard the rows as horizontal. I have taken the

other view, seeing in the quills merely the enlarged members of

the transverse rows which happen to lie along the margin of the

wing and to have become the flight feathers, and the points of

insertion of the feathers in the plucked wing have always

appeared to me to lie in transverse or rather diagonal rows

stretching upwards from the quills and reappearing on the under

side of the wing in the feathers with reversed surfaces. These

diagonal rows were plain in the wing of Aaleniceps, but I was

surprised to find what I have not noticed in any other bird,

although I am by no means prepared to say that it does not

occur, that there was a transverse row too many. The row

corresponding te the carpal covert and remex curved upwards and

backwards ; the next most proximal row had a similar curve and

belonged to the distal secondary quill; then there appeared to be

an extra row in front of, and not behind the second secondary

quill. More proximally the rows were in regular correspondence

with the quills and gradually changed their inclination. I tried

to correlate the arrangement with what W. P. Pycraft has called

the “‘intercalary row” (a transverse row which in his opinion is

associated with the mode of origin of the diastataxic gap by

“faulting” of the horizontal rows in development), but was

unable to make anything of it. No one appears to have pub-

lished any observations on the theory of diastataxy since Pycraft

and myself. In our communications to the Linnean Society (28,

36) we showed that the condition was not due to the loss of a

feather, Pycraft arguing from ontogeny and I from comparative

anatomy. I showed that whereas most pigeons were diastataxic,

a few were eutaxic and had arrived at this condition by a secon-

dary closing of the diastataxic gap. I also showed that the

eutaxic pigeons were in other respects more specialized than

their diastataxic allies. In later papers (29, 31) I showed that

similar conditions existed amongst Kingfishers and amongst

Gruiform birds. The general inference seems to be clear: that

the eutaxic groups are more specialized birds and that in

association with their general specialization they have lost the

primitive diastataxic arrangement. My argument, however,

may be anatomically sound with regard to the groups presenting

both conditions, and yet not applicable to birds generally. All

birds may have been eutaxic originally ; certain families may

have become diastataxic, and amongst these certain members

may have secondarily reverted to the eutaxic condition. Pycraft

assumes that the eutaxic condition was primitive, and if he be

648 DR. P. CHALMERS MITCHELL ON THE

correct, my facts would have to be interpreted according to the

alternative I have just given. But I am not convinced by

Pycraft’s deduction from his observations on the ontogenetic

changes in the wing. The development of the individual is

partly a process of latent differences becoming visible, and

because a diastataxic bird appears to be eutaxic at a very early

stage, it does not follow that eutaxy was primitive. J am much

more impressed by the general view that passerine birds are

plainly the most specialized of all birds, that they are eutaxic,

and that the members of diastataxic groups which have become

eutaxic are in other respects most passerine-like. In the absence

of any convincing theory of the phylogenetic origin of diastataxy,

all peculiarities in the arrangement of feathers are interesting

and may come to have significance, and so I have digressed

with regard to the wing of Baleniceps.

It is plain that the pterylosis of Baleniceps is of the same

general character as that in Storks, Herons and Scopus. To my

eye, the general appearance and coloration suggest affinity with

Herons rather than with Storks. The sedate habit of standing

slently on any little eminence, the absence of the habit of

clattering with the beak, which we noticed in the Gardens, and

the reported heron-like bending of the neck in flight, confirm

this view. But the actual details of the feathering do not

confirm it; without any doubt, so far as pterylosis can be relied

upon as indicating affinity, baleniceps is more Stork-like than

Heron-like.

Petherick, however, (34) has recorded that the young ran about

with extended wings making a “yrattle-like noise produced by

the snapping of their bills.”

Foot and Claw.—The hind toe (hallux) is usually carried

pointing backwards, but is freely movable in every direction. It

is on the same level as the other toes, as in Herons and Scopus,

not slightly elevated as in Storks. There is no trace of a web

uniting any of the toes, whereas in Herons and Scopus there is

usually a distinet web uniting the third and fourth toes, and in

Storks all three front toes are united by web.

A good deal of confusion, which I am able to dispel, has crept

into the literature with regard to the condition of the claw of the

third digit of the foot. Itis well known that the inner edge of

this claw is pectinated in Herons and unbroken in Storks.

Professor Reinhardt (37, p. 378) stated that Baleniceps had not

a comb on the middle claw, adding that this absence afforded ‘a

strong warning not to class it with the Boatbill, as this peculiar

serrature never fails in any member of the Heron tribe.” Gadow

(16, p. 137) divided the Arde into Family 1. Ardeide, dividing

the latter into the sub-families Ardeine and Balenicipitine,

mentioning as a character of the former ‘“‘ Mittelkralle gezihnt,”

and of the latter “ Mittelkralle nicht gezihnt,” and Family 2,

Scopide, including in the definition of the latter, ‘* Mittelkralle

gezihnt.” Beddard (3, p. 289) cites Professor Reinhardt’s opinion

ANATOMY OF THE SHOE-BILL. 649

that ‘as the middle claw is not pectinated, Baleniceps cannot

be referred to the Herons,” adding ‘‘ Professor Reinhardt would

associate Baleniceps particularly with Scopus,” but omitting to

mention that in the same paper Reinhardt stated that the claw

was pectinated also in Scopus. Mr. A. H. Evans (8, p. 87)

divides the Sub-order Ardez into the Families Ardeide (in

which he places Baleniceps) and Scopide, and states that in the

Sub-order the “claw of the middle digit is toothed on the inner

side, save in Baleniceps.” It is surprising that as Gadow,

Beddard, and Evans all seem to have thought the matter worth

comment, they should have omitted to notice Giebel’s (21, p. 351)

very definite description. Guiebel stated that in the two examples

ot Baleniceps he had examined the pectination was clear and

sharp, the actual teeth being not so small, numerous and deeply

incised as in Cancroma and Nycticorax, but larger, separated by

wider intervals, and, beginning at the point, reaching nearly the

middle of the nail. He described them as closely corresponding

with those of Scopus, adding that the latter had not received full

attention from systematists.

I hope that the drawings reproduced in the figure (text-fig.

120), which were made by Mr. Berridge from the actual specimens,

will explain the matter. Ina large number of birds belonging

to different groups, the claw of the third toe of each foot is not

symmetrical about the middle line; the ulnar or abaxial edge is

relatively straight, and the radial edge, that nearest the body, is

curved out into a sharp-edged scoop. The foot of the White Ibis

(fig. 120, 1) shows this condition well. In Baleniceps (fig. 120, 2),

owing to the lateral compression of the claws, the scooped edge is

not quite so conspicuous in a dorsal or lateral view, but it exists.

Moreover, as Giebel described, this sharp edge is marked by a few

serrations between each of which there is rather a wide space,

but which are so conspicuous on the claw that they can be felt

not only along the edge but as slightly ribbing the surface. In

Scopus (fig. 120, 3) the condition of the claw is almost exactly

similar to that in Baleniceps. In the example I examined and

from which the drawing was made the serrations were cut a

little deeper than in Baleniceps, but according to Giebel, in the

specimens of Baleniceps he examined, the serrations were deeper.

The exact amount of serration is no doubt subject to individual

variation. In the corresponding claw of the left foot of the

Umbre I examined, each tooth was much narrower and more

pointed, resembling the Laleniceps condition more closely. In

the Little Bittern (fig. 120, 4) and in the Goatsucker (fig. 120, 5),

the teeth are much more numerous, regular, smaller and more

closely set together, and when the comb is highly developed,

it may stand out conspicuously from the edge of the claw,

sometimes by not being developed along the whole edge, and

sometimes because the thin edges of the teeth give the comb a

yellow semi-transparent coloration, readily visible against the

duller and more opaque unbroken part. Almost every gradation

650 DR. P. CHALMERS MITCHELL ON THE

between the sharp unbroken edge and a fully formed comb can

be found in nearly allied birds, and it must be a matter of

individual taste at what particular stage of elaboration it is

possible to describe the serration as a comb. It is at least certain

that Baleniceps cannot be separated from Scopus and the Herons

and associated with the Storks because of absence of pectination.

Text-fig. 120.

AWariasweeuneneee

ARAN

Modified Claw in some birds.

Claw of the third toe of the right foot, in each case the left-hand figure showing the

dorsal aspect, the right-hand figure the axial side.

Red-billed White Ibis (Hudocimus longirostris).

. Shoe-bill (Baleniceps rex).

. Tufted Umbre (Scopus wnbretta).

. South American Little Bittern (Butorides cyanurus).

. Nacunda Goatsucker (Podargus nacunda).

oo NO

It seems obvious, however, that the formation of a comb falls

in the category of what I have termed “ multiradial apocentri-

cities,’ modifications from the normal or primitive type, which

as they have occurred repeatedly and independently, afford no

information as to the systematic position of the animals in which,

they occur. By the kindness of Mr. Seth-Smith I have looked

through a number of the skins in the collection of the Society,

and with the assistance of Mr. Ogilvie-Grant a still larger number

ANATOMY OF THE SHOE-BILL. 651

in the Bird Department of the British Museum. Most of the

Steganopods have a scoop-like edge forming a primitive stage

like that shewn in the figure (fig. 120, 1), but some of them, for

instance the common Cormorant, havea well-formed comb. The

small claws of Storks seem to have no trace even of the initial

asymmetry, but Ibises are certainly more nearly allied to Storks

than to Spoon-bills, and although the White Ibis (fig. 120, 1) has

only the scoop-like edge, the “Glossy Ibis (Plegadis falcinellus)

has a well-formed comb. Hur, ypyga and Rhinochetus have the

curved edge but no serration; Cursorius, Dromas and Glareola

have weil-marked combs. Among the Strigide, Scops and some

of the smaller owls have the cur ved cutting-edge without serra-

tion ; Aetwpa is in the same condition. Bubo hasa well developed

cutting-edge with a few slight serrations ; Striw flammea has a

well-marked comb occupying the upper part of the edge of the

claw. By a curious accident of nomenclature I was led to

examine some of the Birds-of-Paradise. Fiirbringer mentions

Falcinellus as a genus in which the claw is pectinated, and as I

had forgotten that that name had been used for a genus of Ibis,

and knew that it was used for a Bird-of-Paradise, I examined the

latter, and found that the claw in the Rifle-birds usually showed

a sharp cutting-edge and that occasionally (e. g. Pédlor his and

Epimachus) there were slight nicks in it.

H. R. Davies (7, p. 368) in discussing the function of this organ

remarks that the ‘ pectinated claw should not be regarded as a

structure peculiar to nightjars, owls, herons, cormorants and

gannets, and different from anything found in any other bird,

but merely as a highly modified form of a structure found in a

less modified form in many birds.” There seems to be no doubt

but that the chief use of the modification of the claw is for

scratching, possibly for removing parasites, and its presence

may be compared in a general way with the condition in the

mammalian Dassies, in which all the digits are protected by

flat nails, except the inner digit on each hind foot which is

provided with a sharp claw used in scratching.

Rhamphotheca.—The horny covering of the beak is compound in

Baleniceps, the premaxillary portion being separate, as in Scopus.

In Storks and Herons it is simple. The edge of the horny lower

jaw is delicately serrated.

The Syrinw.—This has been studied and figured by Beddard (3),

and I have to add to his description only that the first two

incomplete bronchial rings are partly calcified, that the bronchi

are relatively rather long, and that the distal bronchial rings

are practically complete. On the most careful examination, I

could find no trace of anything corresponding to what Beddard

took to be fibrous vestiges of the intrinsic muscles present in

Herons ; they were as completely absent as in Storks. I cannot

follow Beddard, moreover, in his view that the structure of the

syrinx is “‘conclusively in favour of regarding Balaniceps as a

Heron and not as a Stork.” So far as the syrinx of Balceniceps

652 DR. P. CHALMERS MITCHELL ON THE

agrees in structure with that organ in Herons and in Scopus, it

conforms not with a structure that is specially Ardeine, but with

one that is found in so many different groups that Beddard

himself has spoken of it as the typical avian syrinx. Storks, as

he himself has shown, display a series of stages most conveniently

regarded as degenerations in different degrees from the typical

avian syrinx. Daleniceps shows degeneration, if not complete

absence of the intrinsic muscles ; it has not degenerated so far as

most of the Storks, but the fact is that as there is no typical

ciconine and no typical ardeine syrinx, the condition in

Baleniceps affords no clue to its relationship with either of these

groups.

Carotid Arteries —The normal condition, present in a large

number of birds of different groups, and what development and

comparative anatomy would seem to indicate as the primitive

avian condition, is the presence of both right and left carotids,

separate and well developed. This is the condition usual in

Steganopods, Herons, Scopus and Storks. I was surprised there-

fore to find that only the right carotid was present in Baleniceps,

and although I sear ched carefully, IT could find no trace of even a

degenerate ‘left artery. I do not attach systematic importance to

the condition of the carotids, as this often varies within a Family.

In Botauwrus, for instance, the two carotids fuse very close to

their origin, and in another member of the Ardeide (Ardetta)

the right carotid only is present.

ALIMENTARY CANAL.

The tongue is only a vestige as in Storks, Scopus and Cancroma,

whereas it is long in Herons.

Stomach.—The stomach is a capacious, elongated, rather thick-

walled sac, extending posteriorly in the Ime of the cesophagus,

from which it is to be distinguished externally only by a gradual

increase of calibre. Distally it ends in a blunted angular point,

and just proximal of this it gives off, on the right side, a globular

chamber from which the duodenum arises (text-fig. 122, P, p. 657).

Externally there is no trace of any specially tendinous area, and

no constriction to mark off a proventriculus from a gizzard. The

interior of the stomach, including the chamber at the pyloric

end, is lined with a thin but well-marked layer consisting of the

hardened secretion of the gastric glands. The cavity of the

stomach is distinctly marked off from that of the csophagus by

the corrugated edge of the membrane-like layer of secretion. The

general cavity is sharply marked off from that of the pyloric

chamber by a well-marked constriction (text-fig. 121). The

greater part of the interior of the larger chamber is marked by

longitudinal folds, but towards the posterior end these pass into

irregular corrugations which are continued into the pyloric

cavity. When the internal surface is scraped and the wall

squeezed, the large glandular apertures may be seen to be

distributed over the whole area of the stomach and pyloric cavity,

ANATOMY OF THE SHOE-BILL. 653

with a slight tendency to be arranged in longitudinal bands, but

there is no indication of separation into a glandular proventri-

culus and a muscular gizzard, and no trace of aggregation into

specialized patches or areas.

Text-fig. 121.

Stomach of Baleniceps.

The distal end of the stomach and the pyloric chamber have been laid open to show

the constriction separating the general cavity from the cavity of the pyloric

chamber and the minute aperture, A, into D, the duodenum.

We have to recognize in the first place that the absence of

distinction between proventriculus and gizzard gives no clue to

the position of Baleniceps in the assemblage of Pelargo-Colym-

biform birds. F. 8. Leuckart (23) discussed this formation in

1841, citing the earlier authors, such as Blasius and Cuvier, who

had called attention to it, described it in a number of birds and

associated it with diet. It is tempting to associate such an

undifferentiated condition with a primitive structure, but I do

not think that such a view is tenable. At one time I myself

thought that it might be possible to derive information useful for

systematic purposes from the condition of the stomach, and I

examined and made drawings of the organ in a large number of

birds. But throughout the group, from Divers to Eagles, the

extent to which gizzard may be separated from proventriculus by

external or internal configuration, by specialization of muscle and

tendon, or by aggregation of glandular areas, varies so irregularly

as to suggest adaptation to habit rather than genetic tendency.

The typical fish-eaters on the whole have a bag-like sac, weakly

muscular and diffusely glandular ; those that live more on flesh

654 DR. P. CHALMERS MITCHELL ON THE

or on mixed diet have the muscular portion more specialized and

a tendency to the concentration of the glands. Hven a formation

so remarkable as the aggregation of the proventricular glands into

two large circular masses has apparently little or no systematic

significance. Within the group generally, the arrangement of

the proventricular glands is diffuse over the whole area, but

there is a tendency for them to lie in longitudinal bands, which

miy be numerous, as for instance in the Sea-eagles and in

Baleniceps, ov in two bands one anterior and one posterior. I

found these bands rather short and rounded off in the American

Grebe (“chmophorus major) and even more definitely rounded

off in Garden’s Night-heron (Vycticorax gardeni). The condition

I described and figured for the African Vantalus (Pseudotantalus

ibis), where the glands are in a couple of rounded bosses, and

which I noted as occurring also in Leptoptilus crumeniferus and

L. argala, and in Carphibis spinicollis (25), is obviously a simple

derivative from the pair of rounded bands. The state of affairs

noted by Garrod in Levaillant’s Darter (20) and by Forbes in the

Indian Darter (11) differs from that in the Storks only by

the circular form of the two patches being a little more advanced

and by a slight tendency for the circular masses to retreat into

eversions of the stomach wall, a condition which is completed in

Plotus anhinga (19) by the two patches having retreated into

a rounded diverticulum. It is clear that the absence of such

circular patches in Laleniceps tells us nothing as to the place of

that bird in the system.

When I found that there was a well-marked pyloric chamber,

a fact which appears not to have been recorded before, I at once

remembered the existence of such a chamber in the Pelican from

my own notes, and in Plots from the observations of Garrod and

Forbes. But as Leuckart (loc. cit.), Gadow (14), Cazin (6) and

many others have shown, a pyloric chamber in varying degrees

of completeness of separation occurs in many birds, notably in

Herons, Storks and Darters; and even if we try to follow Cazin

in limiting the term to cases where the constriction from the

larger chamber is very well-marked, its presence gives no sure

eround for associating Baleniceps more closely with any one of

the groups of allied birds.

Liver.—As Beddard (3) has noted, the right lobe of the liver is

very much larger than the left,a condition which he was inclined

to think showed affinity with the Herons rather than with the

Storks. In a later work (4), however, he states that the ‘“ relative

sizes of the liver lobes appear to be of no importance syste-

matically ”—an opinion with which I concur. There is a large

gall-bladder, and the cystic and hepatic ducts open nearly

together, but the cystic duct distad of the hepatic duct, just

beyond the end of the straight distal limb of the duodenal loop

of the intestines, the disposition being very like that figured by

Beddard in the case of the Indian Darter (4, p. 32) except that

the hepatic duct passes through the substance of the lobulated

ANATOMY OF THE SHOE-BILL. 655

pancreas. On referring to my own notes, I find that a closely

similar arrangement occurs in a very large number of birds

belonging to widely separated groups.

Intestinal Tract.—The most notable general feature of the

alimentary canal is the extremely small bulk it occupied in the

body compared with the size of the bird. The actual specimen

stood nearly four feet high; when the alimentary tract and

mesentery had been removed by cutting the duodenum close to

the stomach and the rectum close to the cloaca, the little handful

of viscera placed on the dissecting board was not so large as the

similar mass from a duck. The calibre of the whole tract was

narrow and fairly uniform, except that the cecum and large

intestine were rather wider. The aperture leading from the

stomach (7. e., from the pyloric chamber of the stomach) to the

duodenum was excessively small (text-fig. 121, A, p. 653);

grain of millet would have had difficulty in passing through. ‘The

minuteness of this aperture is no doubt an adaptation similar in

purpose to the hair-like brush found by Garrod in Plotus anhinga

and the similarly placed plug found by the same anatomist in

Levaillant’s Darter (Garrod, 19 and 20), which he surmised to be

devices for preventing the passage of fish-bones into the gut.

The minuteness of the exit from the stomach and still more

the further guarding of the aperture by a plug of hair-like

structures may have another advantage than prevention of the

passage of fish-bones. The hair-plug occurs also in the Turkey-

buzzard (Cathartes aura), which is certainly not a habitual fish-

eater. In his “Last Journal” (12) under the date Aug. 20,

Forbes mentions dissecting an example of Plotus levaillanti and

finding the stomach full of nematodes, none of which, however,

had penetrated beyond the plug, although several had been caught

init. I found a number of nematodes in the stomach of my

specimen of Baleniceps. We know now that intestinal parasites

may do much damage to the animals they infest, and it is

possible that the plug of hairs in the Darters and the very small

exit from the stomach in Baleniceps, serve the useful purpose of

preventing nematodes, which have been eaten with the food,

from entering the intestines, keeping them in the stomach where

they may eventually be killed.

The aperture by which the distal end of the small intestine

communicates with the cecum and large intestine (text-fig. 123, B,

p. 658) is only just a little larger than the opening into the duo-

denum. The structure of the alimentary tract shows that the

Shoe-bill is adapted to make the most of a limited diet, consist-

ing probably entirely of animal food, fish, frogs, or even small

mammals; that the food must be retained for a considerable time in

the stomach until it is very well macerated, for large lumps could

neither enter the intestines from the stomach, nor even if they

reached the intestines leave them by passing into the cecum and

large intestines. From much unpleasant experience in dissecting

she alimentary tract of birds and mammals, I have learned to

656 DR. P. CHALMERS MITCHELL ON THE

note differences in the odour on the dissecting table. In the case

especially of vegetarian and omnivorous creatures where the

digestive tract is bulky, and large quantities of food are taken,

there is usually a very offensive odour, showing that putrefaction

attends the processes of intestinal digestion. In other cases, and

amongst birds, notably in birds-of-prey and many fish-eaters like

the Divers, the odour of the alimentary tract behind the stomach

is rarely offensive, and, sometimes, even attractive and aromatic.

Although I was unable to examine this specimen of the Shoe-

bill until it had been dead for several days, the contents of the

intestinal tract were not offensive.

In the figure (text-fig. 122) I represent the course of the ali-

mentary canal, from the stomach to the cloaca, dissected out in a

fashion which, as I have described in former memoirs (26, 30),

seems to me to give much information as to its morphology and to

afford a useful basis of comparison with the conditions existing in.

the different groups of birds. The secondary foldings and modes

in which the gut is packed in the body-cavity are naturally not

shown by this method, but the relation of the gut to the primitive

mesentery, the portions of it which have been expanded into

loops, and the configuration of these loops, appear with diagram-

matic clearness. The first specialized loop is the duodenum ; it

is relatively not quite so long asin Herons generally, but it shows

on its distal limb a minor expansion, represented in the drawing

as two short folds, and comparable with the condition which I

have figured in Wycticorax and Ardea. Then follows a second

definite loop with a minor loop on its proximal limb, then a well

marked loop, and then a few irregular twists, after which comes

the portion bearing the remnant of Meckel’s diverticulum (text-

fig. 122, m.), the vestige of the yolk-sac, lying in the line of axis of

the main branch of the portal vein. In Baleniceps this was very

slender, and bound closely to the inferior edge of the gut by a

ventral mesentery. It might quite easily have been overlooked, if

it had been sought for in the usual fashion, merely by running

the gut through the fingers, but when the tract was laid out in

the way I recommend, so that the blood-vessels were visible and

the mesentery undisturbed, it was at once obvious. The part of

Meckel’s tract between this diverticulum and the usual position

of the ceca is thrown first into a series of short ill-defined loops

and then into a long and definite supra-duodenal loop (text-

fig. 122, S.D.F.) closely attached to the duodenum in the un-

disturbed condition and supplied from the duodenal vein by what

I have termed a “bridging” or short-circuiting vessel, which

traverses the mesentery and must be cut through in process of

laying out the gut (text-fig. 122 x, x). Then follows a rather

irregular piece of gut forming two of the folds which I have

described as ‘‘ supra-ceecal” kinks, the presence of at least one

of these being characteristic of the great assemblage of birds

containing the eagles and vultures, herons and storks, penguins

and petrels.

ANATOMY OF THE SHOE-BILL. 657

Just behind the distal extremity of these kinks, the mesentery

narrows so as to bring the distal end of Meckel’s tract very

close to the proximal end of the duodenum. This almost circular

Text-fig. 122.

Intestinal Tract of Baleniceps.

S. Stomach.

P. Pyloric chamber.

D. Duodenal loop.

1, 2, 3. The three loops of Meckel’s tract anterior to the diverticulum,

typical of the Pelargo-Colymbomorphine Brigade.

m. Meckel’s Diverticulum.

S.D.F. Supra-duodenal loop.

x,«. Cut ends of bridging vessels from duodenal loop to supra-

duodenal loop.

K1,K2. Supra-cexcal kinks,

Ca. Caecum.

expanse of mesentery grows from the simple primitive loop of

which the main branch of the portal vein is the axis and which

658 DR. P. CHALMERS MITCHELL ON THE

earries the yolk-sac at its extreme point. If, as happens in many

birds belonging to widely separated groups, Meckel’s tract is

twisted in the course of growth, the twist in the mesentery is

seen here with the result that in the dissection of the gut as

shown in my figures, the mesentery carrying the posterior region

of the gut may be tucked or folded under the mesentery carrying

Meckel’s tract, so that although the two are morphologically

continuous, the continuity may ‘not be apparent except where the

whole gut is short or very simple. In my diagrams, I have

simplified this region, showing the morphological continuity, as

the secondary twisting was not a part of my argument.

The ceca (or caecum) are to be looked for at this point, which

marks the transition from Meckel’s tract to the hind gut at the

Text-fig. 123.

SI.

Cxcum of Baleniceps.

S. I. Cut small intestine.

L. 1. Cut large intestine.

B. The lateral wall of the cecum has been cut away to show at B, the

aperture into the small intestine, lying distally of the point

where the intestine appears to pass into the cecum.

beginning of the area drained by the posterior branch of the

portal vein. In Baleniceps only one is present, as shown in the

general diagram (text-fig. 122, Ca.). The end of Meckel’s tract

meets the large intestine almost at a right angle, and the cecum

is in continuation of the line of the large intestine. The calibre

of the large intestine is rather greater than that of Meckel's tract,

the difference being greater than is represented in the general

diagram and rather less than in the enlarged figure (text-

fig. 123), which was drawn from the specimen after it had been

washed out and slightly stretched in the process of opening. ‘The

ANATOMY OF THE SHOE-BILL. 659

aperture from the small intestine to the large intestine (text-

fig. 123, B) is relatively extremely small and is considerably more

posterior (nearer the cloaca) than the region where the one portion

of the gut joins the other, the actual passage running in the

conjoined walls for a certain distance. There is not more than

the very slightest fold or bulge on the wall of the large intestine,

which may be imagined rather than definitely stated to be a relic

of an originally paired condition of the ceca.

The presence of a single cecum in Baleiiceps has already been

noted by Forbes (10) from a prepared specimen of that region

of the gut mounted in the Museum of the Royal College of

Surgeons, and this specimen corresponds in every particular

with the example which I have dissected and figure in this

communication to the Society. Beddard, however, (38) wrote as

follows :—‘‘In the intestines I could not discover any trace of

exca at all; I believe that the single cecum which characterizes

the Ardeide (there are two in the Ciconiz) may be extremely

minute, and might therefore easily escape recognition in the

spirit-preserved alimentary tract.” I have probably examined

carefully at least as many ceca of birds as any other anatomist,

and Dr. Beddard’s supposition seemed to me extremely improbable

on general grounds. When I found that the specimen in the

College of Surgeons’ Museum was extremely like my own dis-

section, it seemed still more improbable that a structure so

definite and peculiar could be present or absent in different

individuals. Dr. Beddard examined viscera which had been

preserved in spirit and which had been previously handled by

some other investigator. On consulting with him, he was able

to add to the information given in his memoir, that the late

Professor Stewart was rather unwilling that so rare a specimen

should be cut about too much, and he agreed with me that it was

quite possible that the portion of the gut to which the cecum is

attached had: been removed before he examined it. Thanks to

the kindness of Mr. R. H. Burne I have now had the opportunity

of comparing the gut from the example I dissected with the

actual material examined by Dr. Beddard in 1888. Dr. Beddard’s

material was in three pieces and the greater part of the mesentery

had been cut away, but enough of the latter had been left to

enable me to identify with complete certainty the general dis-

position of the gut, to recognize the duodenal loop followed by

the subsidiary loops into which the hepatie ducts open, the large

loop with its proximal minor loop, exactly as in the diagram from

my specimen, the short loop with the remnant of Meckel’s

diverticulum (which Dr. Beddard, apparently, had not noticed) in

precisely the same relative position on the loop and pointing

forwards, the short twists preceding the supra-duodenal loop, and

the latter loop. It was evident, moreover, that the remainder of

the intestinal tract had been cut away, and that Dr. Beddard had

failed to find the cecum because he had not quite the whole of

the small intestine before him, and no part of the large intestine.

Proc. Zoon, Soc.—i913, No. XLIV. 44

660 DR. P. CHALMERS MITCHELL ON THE

Examination of the preparation of the cecum in the College of

Surgeons’ Museum, which Forbes had seen, as cited by Beddard

in a footnote, made it most probable that that was the portion of

the intestinal tract removed from the specimen Beddard examined.

There is therefore no evidence in favour of Dr. Beddard’s

suggestion that the cecum in Saleniceps may be absent, or so

small as to be unnoticed,

The large intestine from the cecum to the cloaca is relatively

long in Baieniceps and is rather wider in calibre than the small

intestine; it is thrown into a series of short irregular loops,

threaded, so to say, on a mesentery which is much shorter than

the course of the gut itself, and which in the usual way is drained

by a large branch of mesenteric vein.

I have shown on a former occasion (30) that the characters

of the intestinal tract are capable of affording a large amount

of information as to the inter-relationships of the groups of

birds. The primitive gut may be regarded as a tube not much

longer than the length of the body-cavity it traverses and

suspended from the dorsal body-wall by an antero-posterior

mesentery. Jt is fixed at its anterior end, where it joins the

stomach, at its posterior end, where it enters the cloaca, and near

the middle of its length, on its ventral surface, where it is con-

tinuous with the yolk-sac. The great embryonic vein forms the

chief radius of this crescentic loop, running up to the dorsal wall

from the yolk-sac, and receiving a large tributary, which runs

parallel with the dorsal wall, from the hind end of the body. In

the course of growth the gut becomes much longer than the

distance separating its anterior and posterior fixed parts, and the

lengthening takes place by the outgrowth of subsidiary loops from

the primitive gut. The position of these loops with regard to the

fixed points of the gut, and their number and character, differ in

different cases, with the result that when the alimentary tracts

are laid out on the dissecting board in the fashion in which I have

described, they form definite patterns. In their main features,

these are constant in individuals of the same species, closely

similar in the species of a genus, and show definite relationships

in the families and greater groups. A type of pattern persists

through the large divisions and shows a gradual increase in

definiteness and specialization in the different members of these

groups, with the result that the patterns can be arranged in

family trees. These correspond so closely with infor mation that

can be derived from other anatomical characters, that I regard

them as being an extremely useful guide to the relationships of

birds. Obviously patterns are more easy to place in the system

when they are highly specialized and complex, but even the

shortened guts of fruit-eaters may retain marked indications of

pattern.

In the great assemblage of birds which Gadow (16) has

called the Pelargomorphine Legion, in which Baleniceps must

certainly be placed, the pattern ‘of the gut is relatively complex.

ANATOMY OF THE SHOE-BILL 66.

The duodenum is a definite loop, tending to be rather wide

and further sub-divided in the Falconiformes, to be excessively

long, twisted and rolled up with the first loop of Meckel’s tract

in the Ciconiiformes, to be very long, narrow and with a minor

loop on its proximal limb in the Ar ‘deiformes. In Baleniceps, i

is not highly specialized, but the differentiation it displays is

more like that in Herons than in Storks, especially in the forma-

tion of the secondary expansions at the base of its distal limb.

In the same assemblage Meckel’s tract 1s long, and is suspended

round the circumference of an almost circular expanse of mesen-

tery which grows out from a very short portion of the primitive

dorsal mesentery It is roughly symmetrical about the main axis

which runs out to the remnant of Meckel’s diverticulum, the

latter being invariably present. On the proximal half of the

tract there is a tendency to the formation of three minor loops

between the duodenum and Meckel’s diverticulum. These tend

to remain comparatively simple and similar in the Steganopods

and in the Faleoniformes. In the Ciconiiformes the tendency is for

the first to become very long and secondarily twisted up with the

duodenal loop, the second and third becoming nearly obliterated.

In the Ardeiformes the first and second are generally rather long

and definite and may form minor loops, the third is often reduced

to a little bunch of small loops closely set together. In Baleeniceps

the three loops are rather more distinct than in either the Storks

or the Herons, remaining in the more generalized Steganopod con-

dition, but such differentiation as exists approaches the Ardeine

pattern and shows no trace of the Ciconine peculiarities.

In the assemblage Meckel’s diverticulum is actually or very

nearly at the extremity of the axial loop of the tract. When it

is not quite terminal, it 1s always on the proximal side of the

loop and is then bent parallel with it, the free tip pointing

proximally. The axial loop may grow out to a great relative

length. ‘These dispositions are distributed very irregularly

throughout the whole assemblage which I am discussing. The

axial loop, for instance, is long and has the diverticuium at the tip

in some Storks and in many of the smaller Hagles and Falcons.

It is short, but usually has the diverticulum at the tip in some

of the Herons and in Scopus, Cathartes and Polyborus. It is

short and bears the diverticulum proximal to its apex in Phaethon,

Pelecanus, Fregata, Pseudotantalus, Serpentarius and Neophron.

The latter condition exists in Baleniceps, and it is plain that its

occurrence has no systematic significance.

(=)

In the Pelargomorphine Legion the distal part of Meckel’s tract

is less highly differentiated, and therefore affords fewer dis-

criminating characters. The ‘portion of it that follows Meckel’s

diverticulum is usually thrown into a bunch of short, irregular

loops, and then follows a well-marked supra-duodenal loop,

supplied from the mesenteric vein, and one or more definite

supra-cecal kinks. In Baleniceps, there is first the irregular

region, then a definite short loop, then a long typical supra-

44%

662 DR. P. CHALMERS MITCHELL ON THE

duodenal loop, and then two supra-cecal kinks. This region

supphes no definite information which might help to place

Baleniceps inside the Legion.

In the Legion the colie ceca are much reduced and apparently

practically functionless except in the Anseriformes, in most of

which they are very large and functional. Although reduced in

the Steganopods, they are rather less so than in the Herons

and Storks and occasionally contain fecal matter. I think the

presence in Baleniceps of a single cecum, by no means so large

as either of those in the Anseriformes, but definitely functional,

communicating with the hind-gut and containing fecal matter,

may be taken to be established. The presence of one cecum,

instead of the normal pair, associates Baleniceps with the Herons.

I should be disposed to guess that the loss of one cecum of the

pair had taken place whilst both were functional, as there seems

no particular reason why one of two vestigial organs should be

suppressed, except as an occasional abnormality, and that the

condition in the Herons, where there is a single functionless

cecum, 1s secondary to that in Baleniceps.

The characters of the large intestine in Birds generally are not

sufficiently differentiated to afford much information of systematic

value. There seems to have been a general tendency to the

reduction of this area to an extremely short and straight course

from the ceca to the cloaca, a tendency which has been inde-

pendently followed by the higher members of a large number of

groups. Baleniceps has a relatively long and capacious large

intestine, and in so far has remained in a rather more primitive

condition than most of the members of the Pelargomorphine

Legion.

To sum up, the characters of the intestinal tract of Baleniceps

are those of the Pelargomorphine Legion, and such specialization

as it displays associates it with Ardeine birds rather than with

Ciconine birds.

In a communication to this Society, Dr. Beddard (5) has

made some additions to or corrections of my observations,

particularly with regard to the presence of a specialized supra-

duodenal loop in birds in which I did not record it, which are the

more valuable as my work stretched over a number of years, as

material was available, and it was only in its course that I began

to recognize the significance of the various points and what had

specially to be looked for. Dr. Beddard also on several grounds

throws doubt on the value of my mode of displaying and com-

paring the intestinal tract patterns. These grounds are due to

misapprehension. He thinks that my method of figuring the

tract gives ‘an appearance of simplicity that is misleading, with

the result that birds which are separated by marked characters

are represented as being almost identical.” Certainly the patterns

(even if correct) do not in every case afford enough information

to place clearly, or to separate clearly cases where the patterns

are very simple. I was rather careful to insist on this point in

ANATOMY OF THE SHOE-BILL. 663

my memoir. Healso thinks that I do not distinguish sufficiently

between what he terms “ fixed loops definitely formed by a narrow

mesentery, and the irregular folds into which any mobile coil of

the intestine may fall when disposed on the dissecting board.”

The whole tract 1s of course supported by mesentery, and

Dr. Beddard’s “fixed loops” and irregular folds are merely the

beginning and final result of differentiation. The loops to which

I have called attention are fixed by their morphological position,

and they may be wide or narrow, long or short. More serious,

however, is Dr. Beddard’s s misapprehension of the morphology of

the gut which leads him into very curious comparisons. He

states “that there are no essential differences between the

intestinal tract in Birds and Crocodiles.” He accepts as “‘ per-

fectly correct” a diagram I gave (26, p. 137) of the alimentary

tract in the Alligator, in which the canal is displayed as showing

a series of almost exactly similar loops from the stomach to

the cloaca, suspended on a crescentic fold of mesentery. As

the pancreas les in the first of these loops, the latter may by

analogy be called the duodenum, but it is simply the first of a set

of regular loops. Meckel’s tract and the delimitation between

that and the large intestine are not shown ; it is quite clear that

Meckel’s tract is not differentiated. Comparison with the next

figure, that of the tract in an embryonic pheasant, shows the

essential difference. Immediately posterior to the duodenum a

mesenteric area, corresponding to a very short length of the whole

distance from the stomach to the cloaca, grows out into an

enormous nearly circular tract, of which the great vein from the

yolk-sac forms nearly a diameter. This region is Meckel’s tract,

and from the point where it returns to the dorsal line again and

where the. ceca, if present, are given off, the large intestine

begins, and corresponds to a much larger part of the primitive

distance from the pyloric extremity of the stomach to the cloaca,

than the combined length of origin of the duodenum and Meckel’s

tract. This mode of development of the gut dominates its adult

morphology. So also Dr. Beddard does not appreciate the mor-

phological importance of the position of Meckel’s diverticulum,

the remnant of the yolk-sac. Fortunately it persists throughout

life in most of the different groups, and its presence rules out

such comparisons as Dr. Beddard makes between particular

loops in Rhea and a Tinamu (with a Passerine intervening in

the argument !). He is trying to identify different morphological

material, belonging to different somites of the embryo, and this

error makes his conclusions invalid. It would be of great interest

to examine young chicks of those birds in which the rudiment of

the yolk-sac does not usually persist, and this would clear up some

of my dubious cases. But so far as they go, the gut-patterus afford

an amazingly conclusive body of evidence as to “the Avian system.

Croaca.—The rectal portion of the large intestine expands

suddenly to enter the large cloaca. The first chamber of the

cloaca, called the coprodeeum by Gadow, is separated by a thin

664 DR. P. CHALMERS MITCHELL ON THE

transverse fold (text-fig. 124, A) from the second, much narrower

urodeum. The paired ureters open into the latter (text-fig. 124, U)

on the dorsal surface but rather laterally placed. The actual

apertures are situated on a small ridge and were very dificult to

see, considering the size of the bird. I found them eventually by

passing a horse-hair backwards through the ureter, and I think

Text-fig. 124.

Cloaca of Baleniceps. Ventral view.

The sphincter has been cut through im the middle ventral line and folded outwards ;

portions of the dorsal wall of the gut have been removed.

LI. Cut end of large intestine.

A. Fold separating Coprodzeum from Urodeum.

B. Fold separating Urodeum from Proctodeum.

U. Aperture of Ureter.

G. Genital aperture.

C. Glandular crypts.

D. Cut sphincter muscle.

F. Aperture of Bursa Fabricii; «, Dorsal wall of Bursa.

it possible, although I am not certain, that there were several

very small apertures at each side instead of one large one.

Rather lateral and slightly posterior to the ureters were the

minute openings of the vasa deferentia (text-fig. 124, C). There

was no genital papilla, but the specimen was a very immature

male, and the testes were minute. The vas deferens accompanied

ANATOMY OF THE SHOE-BILL. 665

the ureter until the latter nearly had reached the wall of the

cloaca, and then twisted outwards. A transverse fold also sepa-

rated the urodeum from the proctodeeum (text-fig. 124,B). In

the proctodeum, at each side and just at the edge of the sphincter,

were four or five little glandular apertures leading into small

cavities lined with ir regular ridges. I find in my notes of dissec-

tions of Ostriches, both male and female, that similar glandular

crypts are present in that bird. In the middle line of the procto-

deum, just behind the fold separating that chamber from the

urodeum, lies the large, elongately oval aperture of the Bursa

Fabricii (text- -fig. 124, “B. The bursa is a very large chamber,

lying above the cloaca, running forwards almost to the rectum.

The inner wall is lined by irregular, heavy ridges, making it

resemble the reticulum of a ruminant stomach. In the figure,

part of the dorsal wall of the coprodeeum and urodeum is repre-

sented as cut away to show the cavity. The bursa was empty.

There was no trace of a penis.

Our knowledge of the Bursa Fabricii is due chiefly to Forbes,

later writers having added very little to his observations and

conclusions (9). In Struthious birds, especially when they are

young, there is practically no constriction separating the procto-

deum and the bursa, the latter being simply a forwardly

directed and dorsally placed continuation of the cavity of the

posterior division of the cloaca. In the different groups of birds

there appears to be a general tendency for a convergent modifica-

tion of this simple arrangement ; the constriction between procto-

deeum and cloaca becomes more and more pronounced, until the

bursa becomes a tubular or pyriform sac opening by a very small

pore into the dorsal wall of the cloaca. This progressive change

is most marked in Passerines and in those birds in other groups

which most nearly mimic the passerine type,and may lead to the

complete disappearance of the aperture and of the bursa. There

is of course no reasonable doubt but that the Passerines present

the most specialized results of avian evolution. ‘To a certain

extent, ontogenetic changes in the bursa show a similar course of

change, the aperture of the bursa narrowing, and the bursa ttself

tending to contract and even to disappear with age. There is

probably, therefore, no special significance in the condition of the

bursa in the example of Baleniceps I dissected, its large size and

wide aperture being perhaps due to youth. Forbes, however,

states that in the Storks and Herons he examined, the bursa was

large and its aperture small. He also mentions the absence in

these birds of the reticulum of ridges in the lining wall of the

bursa, although he found them in Steganopods much as I describe

them in Baleniceps. I cannot draw any systematic conclusions

from these facts.

A small penis is stated to be present in Storks, absent in

Herons, so that in the absence of that organ balewniceps resembles

the latter group, but I attach no systematic value to this.

666 DR. P. CHALMERS MITCHELL ON THE

Muscutar ANATOMY.

Muscurs oF Heap.

Dermo-temporalis.—This thin but extensive sheet of muscle

arose at each side by a fleshy, narrow head, behind and above the

temporal fossa and close to the origin of the biventer maxille,

and spread out on the skin of the ventral surface and sides of

the neck, precisely as in the Pelican.

Biventer maxille was very large and strong, arising from a well-

marked area on the back of the head behind the quadrate and

inserted to the posterior end of the lower jaw. As in the

Pelican, I could not separate this from the underlying digastric

or depressor mandibule.

Temporal.—External portion. A very strong mass, arising

from the dorsal part of the temporal fossa back to the biventer,

runs downwards and forwards, dipping under the ramus of the

maxilla to be inserted to the outer and upper surface of the lower

jaw opposite the orbit.

Pyramidal portion. Strong fan-shaped muscle arising trans-

versely under the post-orbital process and converging toa rounded

tendon which runs forwards and outwards to the inner side of the

lower jaw. Parallel with this, and possibly a separate portion of

it, is a flat band of muscle running from close to the articulation

of the quadrate, alongside the tendon of the pyramidal portion to

be inserted just anterior to it.

Quadrato-mandibular portion. Very strong fleshy muscle

running transversely from the whole of the anterior surface and

forward process of the quadrate to the ramus.

Quadrato-orbital portion. Long muscle from just behind the

optic foramen across to the whole inner edge of the orbital

process of the quadrate.

Pterygoid——An enormous mass of muscle, partly separable

into layers, on the lower surface of the jaw, from the posterior

angle and ventral posterior portion of the mandible running

forwards to the pterygoids.

The temporal and pterygoid muscles are practically identical

with what I have found in the Pelican.

Biventer cervicis.—I examined this muscle to see if there were

any trace of the peculiar formation described by Garrod in the

case of the Darters, but found that the muscle with its anterior:

and posterior bellies was quite normal.

Hyor Muvsctes.

Mylohyoid anterior —As in the Pelican a very slender and thin

sheet of muscular fibres superficial to the other muscles of the

under surface of the jaws, and running transversely across from

the ramus of the jaw to spread out on the interspace between the

two rami, but without meeting its fellow in a median raphe.

Mylohyoid posterior.-—Arises as a broad strap from the outer

surface of the angle of the jaw just below the ear and divides into

ANATOMY OF THE SHOE-BILL. 667

a thin wide sheet of fibres which runs over the ventral surface,

meeting its fellow of the other side and forming a platysma

myoides, and a better defined band which runs across to be in-

serted to the ceratohyal, superficial to both divisions of the

geniohyoid.

Geniohyoid.—Two well-marked divisions. A very strong

division arises from the last joint and cartilaginous end of the

ceratohyal, round which it is wrapped, and runs forwards to the

outer surface of the lower jaw just under the orbit. The second

division arises from the first joint of the ceratohyal and runs

straight forwards as a sheet of fibres which meets the corres-

ponding fibres of the same division of the other side. These

fibres can be traced up to the junction of the rami of the

mandible.

Genioglossus.—Probably in relation with the degeneracy of the

tongue, this muscle was absent.

Ceratoglossus.—Fleshy from the outer side of the first joint of

the ceratohyal to the tip of the tongue. There was no separate

tendon.

The hyoid muscles, like those of the head, of Baleniceps were

excessively like those of the Pelican, but I attach no systematic

importance to the similarity, as I have very little material with

which to compare these muscles in a number of different groups,

and the material J have shows that, apart from obviously adaptive

features, these muscles are much alike in widely separated groups.

CAUDAL MUSCLES.

Pubo-coccygeus externus.—A flat band of muscle from the

posterior dorsal margin of the end of the pubis, narrowing to its

insertion on the under surface of the sheath of the external

rectrix.

Pubo-coccygeus internus.—This is a much wider and thinner

muscle, deep of the externus, and arising from a greater area of

the pubis with a reach on to the ischium. It is inserted to the

heemapophyses of the posterior caudal vertebree.

Levator coccygis.—The two levators form a strong diagonal

mass of musculature on the dorsal surface of the tail, anterior to

the oil-gland. They arise from the ilium and the lateral pro-

cesses of the caudal vertebre and are inserted by a series of

tendinous slips to the spinous processes of the caudals and to the

membrane covering the rectrices.

Depressor coccygis.—Avises from the transverse process of the

last sacral vertebra by a strong tendon just at the articulation

with the ilium, and from the transverse processes of the first

three free caudals; insertion to the transverse processes and

hemapophyses of the posterior caudals.

Ilio-coccygews.—Only the outer of the two slips which usually

represent this muscle is present. It arises from the illum just

dorsal to the origin of the depressor coceygis, and is inserted to

the outer surface of the capsule of the external rectrix.

668 DR. P. CHALMERS MITCHELL ON THE

I have no standard for comparison in the case of the caudal

muscles. ‘The chief difference from the condition in Leptoptilus

is the absence of the inner slip of the ilio-coccygeus.

MUSCLES OF THE SHOULDER AND WING.

Cucullaris.—Vhe cervical portion is well developed, forming a

definite sheet of circular fibres which stop abruptly in line with

the proximal edge of the rhomboideus externus where they are

inserted along the clavicle.

Lthomboideus externus.—Orvigin tendinous from the neural

crests of five and a half vertebre beginning at just opposite the

junction of the scapula and clavicle. The fibres run outwards

nearly transversely to all the scapula except the down-turned

posterior end and forwards to part of the clavicle (text-fig. 125,

Rh. 2).

LRthomboideus profundus or internus.—Origin tendinous, a little

short of the externus proximally and reaching just beyond it

distally. The fibres run outwards and backwards to no part of

the clavicle but to the whole length of the scapula including the

down-turned end (text-fig. 125, Rh. 1).

The two rhomboid muscles are nearly equal in thickness.

These two muscles, according to Fiirbringer, and my own obser-

vations confirm his view, are in process of creeping forwards.

Their condition in Baleniceps shows a considerable degree of

specialization, but I have not material to compare the condition

in allied birds.

Latissimus dorsi anterior.—A broad strap of muscle arising

from the anterior dorsal vertebre only and running downwards

and forwards, dipping under the anconeus, to a fleshy insertion to

the shaft of the humerus distal of the insertion of the posterior

division of the muscle and unconnected with it (text-fig. 125, L.A.).

Latissimus dorsi posterior,—Fleshy origin, the anterior edge of

which touches but is not fused with the posterior edge of the

anterior division. Origin wider than that of the anterior

division, but not reaching quite as far back as the proximal edge

of the ilium. Its fibres converge to form a band about the same

width as the lat. dorsi anterior (text-fig. 125, L.P. 1), pass under

that muscle with a more proximal slope, to be inserted along a

strong tendon (text-fig. 125, L.P. 2) which is inserted to the

scapula under the scapular anchor of the anconzus, proximal to

the insertion of the lat. dorsi anterior, and which joins the

anconeus belly distally.

Latissimus dorsi metapatagialis.—Absent.

The anterior division is like that in Leptoptilus. The connec-

tion of the tendon of insertion of the posterior division with the

anconeus occurs also in Leptoptilus, but I have noted a somewhat

similar arrangement in Bubo maximus. The loss of the meta-

patagial division has been noted by Fiirbringer in Plotus, but it

is usually present in the Herons, Storks, and Steganopods.

ANATOMY OF THE SHOE-BILL. 669

Serratus superficialis anterior.—¥rom the ventral end of the

last cervical rib and the first dorsal rib converging to a flat

Text-fig. 125.

Rh...

Ve —— Dela)

TH Ui inane Del. 2

WO SH YK WN \\\ oly hing DY Rrra €

fis iat mi i yk Ri So Seah

yf ft! = TARR

fi) ANT

Shoulder-muscles of Baleniceps.

Right wing, external aspect. Muscle striped: tendon dotted.

Rh. 1. Rhomboideus profundus, cut across.

Rh. 2. Rhomboideus externus, cut across.

Del. 1. Tendinous anchor of Deltoides major.

Del. 2. Cut surface of Deltoides major reflected.

Del. 3. Distal portion and insertion to humerus of Deltoides major.

S.P. Scapulo-humeralis posterior.

Pee.m. Tendon of insertion of Supra-coracoideus (Pectoralis minor or secundus).

$.C. | External scapular head of Sub-coraco-scapularis.

A.S,. Anconzeus scapularis, showing scapular origin, and anchor to humerus. It

has been divided to show the Latissimus dorsi.

Anconeus scapularis, part of the belly.

Anconeus humeralis.

Latissimus dorsi anterior, insertion.

Cut edge of Latissimus dorsi posterior. :

Tendon of insertion of Lat. dorsi posterior from humerus to junction with

Anconeeus scapularis.

Vestige of Expansor secundariorum. The distinctness of this is exaggerated

in the drawing.

670 DR. P. CHALMERS MITCHELL ON THE

tendon which is inserted to the postglenoid scapula between the

two parts of the sub-coraco-scapularis, its insertion being quite

covered by the outer part of that. \

Serratus superficialis posterior.—From the first two uncinate

processes and area of their ribs in line with them to about three-

quarters of an inch of the posterior inferior border of the scapula ;

covered .by the third portion of the serratus superficialis.

Serratus superficialis metapatagialis —Origin from the same

two ribs as the ser. super. posterior, but entirely from below the

uncinate processes, and reaching down almost to the sternum.

Insertion to the metapatagium, Cwith a strong tendinous slip to

the tip of the scapula.

The first of the three serrati, according to Fiirbringer, is very

variable even within families. ‘The second is constant in many

families, variable in others. The limitation of the origin to

dorsal of the uncinates is somewhat rare but has been noted in

Phenicopterus. The insertion of the serratus metapat. partly to

the scapula appears to be extremely rare. Fiibringer has noted

it in Crex, where the condition 1s much as I find it to be in

Baleniceps, and in Fulmarus and Bucorvus, where the scapular

insertion alone occurs. The relation to the scapula is probably a

vestige of the origin of this muscle as a separated portion of the

serratus superfic. posterior.

Serratus profundus.—From the last two cervical and first two

dorsal ribs to the scapula in four digitations. According to

Fiirbringer, this arrangement is normal in Herodii.

Biceps brachii.—This arises by a flat narrow tendon from the

acrocoracoid (text-fig. 128, B. 1, p. 675), alongside but not covered

by the origin of the coracobrachialis externus and separated by

that muscle from the tendon of insertion of the supracoracoideus

(pectoralis secundus). It passes under the insertions of the

pectoralis major without being connected with them, and passing

into a rounded belly (text-fig. “128, B. 2) runs down parallel with

the humerus to be inserted toa knob on the ulnar face of the

radius (text-fig. 127, Bi., 1, p. 672). It is then continued across

to the opposite face of the ulna (text-fig. 127, 4, 5) by a deep

broad tendon and a narrow more superficial tendon, first sending

a strong slip (text-fig. 127, 2) to the radial end of a radio-ulnar

ligament.

The biceps obviously presents a highly specialized condition in

Baleniceps, the specialization consisting of the complete loss of

the usual humeral head. The two divisions are well separated

at their origin, and the radial and ulnar tendons of insertion

separate rather high up in most Steganopods, Storks and Herons.

As Furbringer has pointed out, the humeral head in such cases

ean be traced to the radial insertion. As both radial and ulnar

insertions are well marked, indeed rather unusually complex in

Baleniceps, 1 infer that the loss of the humeral head is com-

paratively recent.

Biceps patagialis —This slip to the patagial tendons is absent

ANATOMY OF THE SHOE-BILL. 671

in Baleniceps, as in Storks, Herons, Scopus, and mest Steganopods.

The absence, however, is not of much value; the slip is present

in some Steganopods, in Spoonbills, in Phenicopterus, and is

present or absent within the same family in a number of cases.

Deltoides propatagialis (text-fig. 126, Del. pat.).

I have already mentioned that there is no biceps propatagialis.

Text-fig. 126.

cry A

Patagial muscles and tendons of Baleniceps.

Del.m. Deltoides major. Ane. Anconzus scapularis. Hum. Humerus. Bie.

Biceps. Del.pat. Deltoides propatagialis. Pec. Pectoralis major cut across.

P.l. Pectoralis slip to longus tendon. P.b. Pectoralis slip to brevis tendon.

Lon. Longus tendon. Brev. Brevis tendon with the slips named «, 3, and

y by Fiirbringer. Ex. Extensor metacarpi radialis.

672 DR. P. CHALMERS MITCHELL ON THE

There is no cucullaris propatagialis. The patagial tendons arise

solely from the deltoides propatagialis and from the pectoralis

propatagialis, and there are no anchors to the humerus.

The origin of the deltoides patagialis is from the clavicle,

acrocoracoid ligament and part of the scapula, the latter origin

not being found in Herons. Distally it sphts into two peaks, a

smaller from which the longus tendon arises and a larger for the

brevis tendon. This is a more specialized condition than in

Steganopods, Storks and Herons generally, where even the longus

and brevis tendons have a short common course, but in Scopus

and Leptoptilus there ave separate peaks for the tendons.

Text-fig. 127.

Bo.

Insertion of Biceps hgament.

R. Radius. U. Ulna. Bi. Biceps tendon: 1, msertion to radius; 2, insertion to

radio-ulnar ligament; 4, insertion to ulna; 5, second superficial insertion to

ulna ; 3, radio-wnar ligament. L.h. Humero-ulnar hgament.

The longus tendon (text-fig. 126, Lon.), after being reinforced

by a slip from the pectoral, enlarges in width and becomes elastic,

this portion being doubled, and being anchored by a very faint

(much fainter and more diffuse than would appear from the

drawing in fig. 126) set of fibres from the distal portion of the

brevis.

The brevis tendon (text-fig. 126, Brev.) is highly specialized.

The main mass of the muscle passes into a strong rounded tendon

which is reinforced by the pectoralis slip and represents the con-

joined a and /3 slips of Furbringer, the beta slip being the direct

continuation of the muscle, but giving off half-way down the

patagium a broader and weaker alpha slip. The latter itself

becomes doubled distally, gives off a weak anchor to the elastic

portion of the longus tendon and is inserted to the extensor

metacarpi radialis tendon. ‘The beta slip broadens out as it

reaches the fascia over the extensor, sends forward a stout anchor

which covers and is fused with the tendinous head of the extensor

metacarpi, and sends downwards a branch which forms a forked

fan reaching the distal edge of the forearm. The gamma slip of

ANATOMY OF THE SHOE-BILL. 673

Fiirbringer is the most proximally placed, and is distinct although

very flat and weak throughout its whole length from its origin at

the proximal side of the brevis muscular peak to its insertion to

the recurrent beta slip.

The course of evolution of the brevis tendon appears to have

been from a wide rather diffused band to first a specialization of

portions of that band into the slips distinguished by Firbringer,

then to a separation of these slips, and finally to the loss of one or

more of them. In the Storks, Herons and Scopus the slips are

at least separate distally ; in Storks and Herons the separation of

alpha and beta is only distal and does not begin so high up as in

Baleniceps, in which, although the actual separation occurs only

about half-way down the patagium, the identity of beta can be

traced right up to its origin. Beddard figures an almost

similar condition for Scopus (2, fig. 2). So also the very com-

plete separation of gamma and beta occurs in Scopus and

Baleniceps, and is much less distinct except distally in Storks and

Herons. The resemblance between Baleniceps and Scopus appears

to be rather close; the most important differences being the

‘greater distinctness of the anchor to the longus in Scopus, and

the presence of an anchor to the humerus in the same bird.

Delioides major (text-fig. 125, Del. 1, Del. 2, Del. 3, p. 669;

text-fig. 126, Del.m.; text-fig. 128, De.).—This large muscle arises

fleshy ‘from the scapula but with a distal tendinous anchor just

external to that of the anconzeus, and is inserted fleshy to nearly

half-way down the humerus. It is very nearly divided into

the two portions visible in Leptoptilus and other storks. The

tendinous anchor occurs in the Herons and Storks that I have

dissected, and Beddard has recorded it in Scopus.

Deltoides minor.—This muscle, possibly owing to the large size

of the deltoides major, is not to be distinguished as a separate

muscle: probably it is absent. In Storks it is small and quite

separate.

Scapulo-humeralis anterior.—This small muscle is absent. In

Steganopods, Storks and Herons it les very close to the teres

major, so that it is possible that it may have fused with this in

Baleniceps. Beddard does not mention it in his description of

the shoulder muscles of Scopus, so that possibly it may also be

absent in that bird.

Scapulo-humeralis posterior (Teres major) (text-fig. 125, SP).—

A strong but relatively rather small muscle arising from about

the distal half of the scapula and inserted to the humerus between

the two heads of the anconeus. A relatively narrow insertion,

according to Furbringer, also occurs in Steganopods, Storks

and Herons. In Saleniceps it has no accessory anchors or

attachments.

Swb-coraco-scapularis.—The coracoid head (Coracobrachialis

brevis of Garrod) is single and much smaller than the scapular

heads. It arises only from the proximal half to third of the

inner face of the coracoid, as in Storks and Herons, and converges

674 DR. P, CHALMERS MITCHELL ON THE

to join the scapular heads near their tendon of insertion. The

external (text-fig. 125, 8.C., p. 669) and internal scapular heads

arise from about the second fifth of the under surface of the

scapula, where they are separated by the insertion of the serratus

superficialis anterior. The three heads unite to form a strong

rounded tendon inserted to the median tubercle of the humerus.

‘

AnCOnCUS.

Anconeus scapularis.— Origin by a strong forked tendon

from scapula (text-fig, 125, A.S.1, A.S. 2), passes into a rounded

muscular belly which sends an anchor to the humerus near the

insertion of the latissimus dorsi and receives a strong tendon

from the latissimus dorsi posterior. Passes into a strong flat

tendon just before reaching the elbow.

Anconeus humeralis (text-fig. 125, A.H.).—Origin from the

whole length of the humerus, the origin being cleft proximally.

Passes into a tendon at the distal end of the humerus, and this

runs parallel with but united only by membrane to the tendon of

insertion of the anconeus scapularis. Insertion to the olecranon

of the ulna.

The forked head of the scapular portion, the anchor to the

humerus, and the general relations of the two divisions of the

muscle are very much like what I have observed or find recorded

in Storks and Herons. The absence of any extension of the

scapular head to the clavicle or coracoid is rather a primitive

feature.

Anconeus caput coracoideum (Hupansor secundariorunr) (text-

fig. 125, 8.)—At the elbow there was a slip of muscular fibres

connected with the feathers and giving rise to a very delicate

tendon which I traced up the under surface of the skin close to

the anconzeus, but which then appeared to become diffuse and be

lost in the subdermal fascie. There was no trace of it passing

through the edge of the teres major, as usually happens when it

is well developed, or in the axilla.

This was one of the muscles to which Garrod paid great atten-

tion, hoping to find it useful in classification, but further obser-

vations have not justified his anticipations, as it 1s present or

absent in very closely allied birds. It is usually absent in

Steganopods, but present in a few cases. It is present in Storks,

and in Herons except Ardetta and Cancroma. According to

Beddard it is absent in Scopus. Its vestigial presence in Balent-

ceps is therefore interesting but of no systematic value.

Pectoralis thoracicus.—The great pectoral (text-fig. 126, Pec.,

p. 671; text-fig. 128, Pec.) in Baleniceps is an enormous mass of

muscle arising from the clavicle, the membrane between the clavicle

and coracoid, from the whole of the keel and from all the posterior

part of the sternum with a considerable overlap to the ribs. I

could not trace any definite horizontal division of the muscular

mass. The insertion is by two very distinct tendons, which

cross each other in a remarkable fashion. The greater and more

ANATOMY OF THE SHOE-BILL. 675

proximal mass of the muscle converges to a strong flattened

tendon (text-fig. 128, 1), which is inserted to the humerus rather

distally and not far from the posterior end of the deltoid inser

tion. The more distal portion of the muscle converges to a mucl.

broader tendon, which forms the posterior border of the whole

muscle and then dipping under the first tendon of insertion runs

in to the humerus proximally of it (text-fig. 128, 2). There is

also a strong anchor to the humerus, shown as cut and reflected

in the figure (text-fig. 128, 3).

Text-fig. 128.

Pectoral muscle of Baleniceps. Tendon dotted ; muscle striped.

Acro. Acrocoracoid process. Hu. Humerus.

Pec. Pectoralis majer, cut across.

1, 2. Insertion tendons of pectoralis to humerus. 8. Anchor to humerus,

divided and reflected.

P.pt. Pectorales propatagiales.

Su. Tendon of insertion of supracoracoideus (Pect. minor).

De. Deltoides major.

Cor. ex. Coracobrachialis externus.

B.1. Tendon of origin of biceps. B.2. Cut belly of biceps.

The large area of origin of the great’ pectoral is of course

associated with a powerful wing, and is probably purely adaptive

Proc. Zoou. Soc.—1913, No. XLV. 45

676 DR. P. CHALMERS MITCHELL ON THE

The strong proximal anchor (text-fig. 128, 3) ito the humerus also

occurs In many Steganopods, mm Storks, Herons, and is probably

vepresented in Scopus, 12 which bird Beddard mentions a strong

insertion to a fibrous aponeurosis attached to the crista of the

humerus and covering the biceps. It is characteristic of Storks.

as opposed to Herons, that in the former birds the great pectoral

is completely divided into two msucles, as it 1s in the Pelican

and some other Steganopods. ‘The insertions of these portions as

described by Weldon correspond almost exactly with the double

insertion in Baleniceps, and I was able to separate the mass of

muscle quite easily into portions corresponding with these inser-

tions, although, in the absence of the separate insertions, I should

not have described the muscle as doubled. But, whatever the

distinction be worth, the great pectoral muscle of Daleniceps is

more Ciconine than Ardeme. Beddard’s description of the condi-

tion in Scopus is not sufficiently detailed to follow in this matter,

but he speaks of it as ‘‘ partly doubled,” and the humerus shows

marks of a double insertion.

Pectoralis propatagialis (text-figs. 126, 128, pp. 671, 675).—-As IT

have already stated, there are separate slips from the pectoral for

the longus and brevis tendons. Both slips are entirely tendinous,

and that for the longus is smaller and more superficial (text-fig.

126, P.1., P.b.; text-fig. 128, P.pt.). In my dissections of Herons,

I find similarly cdistinet slips for the brevis and longus from the

pectoral ; Beddard mentions them for Scopus, but-in the case of

Storks the usual arrangement appears to be the more primitive

condition of a single slip, which joins the patagial tendon before

that has divided into longus and brevis.

Pectoralis abdominalis.—This is absent in Baleniceps as in

Storks. It is present in Herons, but Beddard does not refer to:

its presence or absence in the case of Scopus.

Supracoracoideus (Pectoralis minor).—This is a small and

rather narrow muscle elongately oval, with a centrally placed

tendon like the mid-rib of a leaf. Its fleshy origin is limited to:

a very small part of the sternum, including no part of the keel,

and part of the coracoid and the membrane between the coracoid

and clavicle. It is widely separated on the coracoid from the

origin of the coracobrachialis posterior (pectoralis tertius) and its

tendon of insertion (text-fig. 128, Su.) is free from any fibres.

that could represent a deltoides minor. Its general relations and

small size are closely paralled in Herons and Storks.

Coracobrachialis externus or anterior.—A strong muscle arising

fleshy from the acrocoracoid only (text-fig. 128, Cor. ex.) and

covered only at the extreme edge by the tendon of origin of the

biceps. Insertion on a bro adly oval area to the planum: bicipitale-

of the humerus.

Coracobrachialis internus (Pestoralis tertius).—A very stout

almost doubled mass of muscle from the distal dorsal two-thirds

of the edge of the coracoid opposite the origin of the supracora-

ANATOMY OF THE SHOE-BILL. 677

coideus, but quite separate from that. Its fibres converge to a

strong tendon inserted to a peak of the median tubercle of the

humerus.

Muscurs oF ForREARM AND NECK.

Brachialis inferior.—A very strong flat sheet of muscle with «

fleshy origin and insertion, occupying the angle between the

humerus and ulna, the insertion to the ulna being twice the

width of the origin from the flexor aspect of the humerus,

Pronator sublimis or brevis.—Short muscle from the inner

condyle of the humerus to the first quarter of the radius. This

insertion 1s rather shorter than in Leptoptilus.

Pronator profundus or longus.—As in Leptoptilus a larger

muscle, from the inner condyle of the humerus to a little beyond

the surface of the radius covered by the brevis and more on the

ulnar aspect than the brevis.

Entepicondylo-ulnaris.—Absent, as in Leptoptilus, but accord-

ing to Gadow present only in Rasores and Tinamus.

Hetepicondylo-ulnaris.—From outer condyle of the humerus to

first third of ulna on its radial face. A very thick and strong

muscle, closely united with the flexor digitorum profundus, As

in Leptoptilus.

Eetepicondylo-radialis.—A thin muscle arising by a flat tendon

from the outer condyle of the humerus along with the extensor

digitorum communis, inserted to a quarter of the radius; as in

Leptoptilus, except that the insertion is shorter in the latter

bird.

Flexor carpi ulnaris.—From the inner condyle of the humerus

with a sesamoid ; runs down the inner surface of the ulna to the

great tuberosity of the ulnar carpal. Arising as a fleshy belly a

thinner tendon connected with the quills runs down to end on

the carpal alongside the great tendon. As in Leptoptilus.

Ulni-metacarpalis ventralis.—Fleshy from the last third of the

ulna on the radial face; tendon crosses over a slide on the radial

carpal and is inserted on a hump of the second metacarpal near

the attachment of the pollex.

Ulni-metacarpalis dorsalis.—Short muscle arising by a tendon

from the distal end of the ulna on its lateral face ; it divides into

a shorter portion running straight across to the upper part of

metacarpal ITT and a broader portion inserted to about two-

thirds of the upper surface of metacarpal III, where that is free.

Similar in Leptoptilus, except that the first portion is tendinous,

the second fleshy, while both are fleshy in Baleniceps.

Hetensor metacarpi radialis—Two heads, outer tendinous,

inner fleshy, from the outer condyle of the humerus. The outer

belly is quite separate from the inner belly and is connected with

the brevis tendons of the patagium (text-fig. 126, Ex., p. 671).

Insertion to the base of metacarpal I, the tendons from the two

45*

678 DR. P. CHALMERS MITCHELL ON THE

bellies remaining separate until their insertion, so that the muscle

is completely double. In Leptoptilus the tendons fuse distally.

Extensor metacarpi ulnaris.—Avises from the external condyle

of the humerus by a tendon superficial to that of the ectepi-

condylo-ulnaris ; then a long fleshy belly, then a thin tendon

passing over a groove in the distal end of the ulna from which it

receives a strong anchoring slip, absent in Leptoptilws, to its in-

sertion on metacarpal IT just where metacarpal IIT is given off.

Flexor digitorum sublimis.—A strong band of tendon runs

from the inner condyle of the humerus to the ulnar carpal, from

the upper side of which the flexor digitorum sublimis arises as a

delicate fleshy muscle giving rise to a slender tendon which passes

over a groove in the ulnar carpal and is inserted to the base of

phalanx 2 of digit II, but with first an insertion to the base of

the first phalanx of that digit, which I do not find recorded in

my notes on Leptoptilus.

Flevor digitorum profundus.—Arises fleshy from the second

and third fifths of the ulna, very closely connected with the in-

sertion of the ectepicondylo-ulnaris. The tendon begins where

the origin from the ulna ceases, and runs down the radial face of

the ulnar carpal under the ligament from the radius to meta-

carpal IT, and then follows the tendon of the superficial flexor to

be inserted just beyond it to phalanx 2 of digit IT. It receives

a strong slip from the short extensor of the thumb, which I did

not record in the case of Leptoptilus. Gadow mentions somewhat

similar relations with the thumb in the case of Owls and

Feliornis. ae

Extensor digitorum commuinis.—Arises tendinous from the

external condyle of the humerus and passes into a slender belly a

quarter way down the forearm, but receives no fibres from the

ulna. Its tendon of insertion passes througha groove in the end

of the ulna and then sends a branch to the base of phalanx 1 of

digit Iand a stronger tendon to phalanx 2 of digit IT. As in

Leptoptilus.

Extensor pollicis longus.—Two slender fleshy heads from the

adjacent surfaces of the radius and ulna at their proximal ends,

with accessory fibres froma large part of the length of the radius

on its ulnar face. Tendon unites with that of the extensor

metacarpi radialis at its insertion. As in Leptoptilus.

Extensor indicis longus.—One head fleshy from the distal half

of the radius and a second much smaller, tendinous from the

distal end of the radius and from radial carpal. Insertion to the

second phalanx of digit IT, but attached by fascia to the first

phalanx. As in Leptoptilus.

Tnterosseus dorsalis.—Arises fleshy from the opposite faces of

metacarpals II and Il]; fibres run to a centrally placed tendon,

like the midrib of a leaf, and this is inserted to the base of the

second phalanx of digit II. As in Leptoptilus.

Interosseus palmaris.—More ventral and stronger than the

foregoing muscle but with similar origin and arrangement.

ANATOMY OF THE SHOE-BILL. 679

Tendon inserted to phalanx 2 of digit IT. In Leptoptilus I have

noted it as reaching only the first phalanx.

Abductor indicis.—Strong muscle arising fleshy from the whole

of the radial side of metacarpal II. Inserted to the base of the

phalanx | of digit IT. As in Leptoptilus.

Flexor digitt [1T.—Arises fleshy from the ulnar side of meta-

carpal ITI and is inserted to the base of the first phalanx of the

corresponding digit. As in Leptoptilus.

Adductor pollicis—A strong muscle from the metacarpal to

the tip of the pollex.

Hetensor pollicis—This muscle, which is usually described as

single, is represented by two distinct and well-developed muscles,

a condition which has been described in the ease of Struthio, but

not in other birds. Most probably, if it were carefully looked

for, it would be found elsewhere. The first of the two isa strong

slip from metacarpal I and the tendon of the extensor metacarpi

radialis to the radial side of the thumb. It is this muscle that

gives off a slip to the flexor digitorum profundus. The second

muscle is from the head of metacarpal II to the base of the

thumb.

MuSscLEs oF THE THIGH AND Lec.

Iio-tibialis internus (Sartorius).—A large and strong strap

arising from the anterior and lower border of the ilium and from

the fasciee over the gluteus medius, and closely united along its

distal border with the gluteus maximus. Normal insertion to

the tibia. I find no notable difference as compared with Storks

and Herons.

Ilio-tibialis (Gluteus maximus).— The origin is entirely ten-

dinous and a median tendinous area separates the fleshy anterior

and posterior borders. Posteriorly the origin extends backwards

half-way over the origin of the biceps, that is to say what Garrod

called the post-acetabular part of the muscle, the ilio-tibialis pos-

terior, is present. Garrod tried to use the presence or absence of

this in his systematic arrangements, but without much success.

It is absent in. the Steganopods generally, usually absent in Storks

but present in Ciconia, absent or very slightly developed in

Herons. So far as I can judge, the presence of the post-acetabular

portion of this muscle is a primitive condition, and it has been

lost or reduced independently in many groups of birds.

Ilio-trochanterici.— The externus (text-fig. 129, Gl.a) and the

posterior (text-fig. 129, Gl.2) are both present in the normal

condition. The minimus and quartus are represented by a single

tendon of origin and muscular belly (text-fig. 129, Gl.3). In

Leptoptilus I found these quite distinct in their origin and inser-

tion; in Vycticorax they had a common tendon to the femur but

separate insertions to the ilium. These muscles, however, vary so

much from bird to bird that I cannot attach any significance to

their distinctness or fusion.

680

DR. P. CHALMERS MITCHELL ON THE

Text-fig. 129.

Va:

"EL. Pro.

Diagram of Muscles of the leg in Baleniceps.

Left leg, external aspect. Tendon is dotted.

FEMUR. Femur. FIB. Fibula.

Gl.a. llio-trochantericus externus (Gluteus anterior). Gl. 2. Il.troch.

posterior (Gluteus minor). GI. 3. Il.troch. anterior et medius

(Gluteus minimus and quartus).

Ob.ex. Obturator externus.

Ob.in. Obturator internus, surrounded by Gem., Gemellus.

FE.C. Origin of Femoro-caudal.

A.D. Adductor longus (the upper muscle) and Adductor magnus.

BIC, Insertion of Ilio-fibularis or biceps, passing through a sling.

Gas. External head of Gastrocnemius, cut and reflected to show relation

to short arm of the Biceps sling.

F111, Fl1.10, Fl.1.1V. Tendons to respective toes of the Perforated

Flexor muscles.

Amb. Ambiens head of Perforated Flexors.

Fl.1.E. External head of Perforated Flexors. :

FI.2.11, F1.2.11I. Perforated and Perforating Flexors of digits II & ITT.

Fl. Hal. Flexor longus hallucis.

Fl. Pro. Flexor profundus.

Ex.Com. Extensor communis.

ANATOMY OF THE SHOE-BILL. 681

Ilio-femoralis internus (Pectineus).—Strong, little fleshy slip

from the ventral edge of the preacetabular portion of the ilium to

the back of the femur below the neck. As in most birds.

Femori-tibiales (Crureus and Vastus).—The external muscles

are fused to form a common mass; the vastus internus is large,

arising from nearly the whole length of the femur.

Caud-ilio-femoralis (lKemoro-caudal).—This is a broad thin

strap of muscle arising fleshy from the femur (text-fig. 129, F-H.C.)

and running upwards and backwards under the biceps and be-

tween the semitendinosus and the semimembranosus and passing

to the underside of the tail, where it becomes a thin tendon

spreading out into a sheet which meets its fellow of the other

side, the combined insertion being to the tendon of the depressor

ecoccygis where that is inserted to the hzemapophyses of the

posterior caudal vertebree. There is no accessory femoro-caudal,

The accessory fem.-caud. is, I believe, invariably absent in Stega-

nopods, Herons and Storks, although it is present in Spoonbills

and the Flamingo. The fem.-caud. itself tends to be degenerate.

It is usually present in Storks, but is very slender in /%ssuia and

absent in Leptoptilus; it is weak in the Herons and absent in

several genera.

Caud-ilio-flecorius (Semitendinosus and Accessory semitendi-

nosus).—Origin fleshy from the ischium behind the biceps and

extending on to the fascia posterior to the ischium; meets the

rather small but distinct accessory or femoral head in a tendinous

raphe, and the combined muscles are inserted to the middle belly

of the gastrocnemius (text-fig. 130, C.J]. 1 & 2). The muscle is

much weaker than the semimembranosus.

Ischio-flecorius (Semimembranosus).—This ig the usual broad

strap underlying the semitendinosus, and in this case much

thicker and wider than the latter. It has a wide origin from the

lower edge of the ischium and the fascia over the obdurator

externus, is unconnected with the semitendinosus, but receives a

strong tendinous slip (text-fig. 130, Sl.) from the inner adductor

and then is inserted to the tibia by a flat tendon.

Gastrocnemius.—There are the usual three heads of which the

tibial head is the strongest. The outer head arises from the

external condyle of the femur in common with the short arm of

the biceps sling (text-fig. 129, Gas.). The middle head is the

smallest and arises from between the condyles of the femur by a.

flat tendon. The tibial head is enormous and arises from the

tendon of the ilio-tibialis and from the cnemial crest of the tibia.

The three heads unite in the usual way rather less than half-way

down the leg to form the tendo achillis.

Relations of the Caud-ilio-flexorius, [schio-flexorius, and Gastro-

cnemius.— Weldon (38) called attention to the varying relations

of these muscles and the differences they presented in Storks,

Ducks, and Phenicopterus. I have noted them in a number

of Storks and Herons and paid a good deal of attention to

them in Gruiform and Limicoline Birds (31 & 32). With

minor variations as to the precise interconnections of the

682 DR. P. CHALMERS MITCHELL ON THE

semitendinosus and semimembranosus, the condition which I

found in Laleniceps seems to be typical in Herons and Storks

(text-fig. 130). ‘The middle head of the gastrocnemius is joined

by the semitendinosus just after the latter has been met by

its accessory. ‘The accessory origin from the femur is parallel

to but separate from that of the middle head. The accessory is

present in Herons, Storks, Seopus, and in Daleniceps ; it is

frequently absent among the Steganopods. In my opinion the

middle head of the gastrocnemius is a separated portion of the

accessory semitendinosus, and separation of the two, with sub-

sequent disappearance of one or of both, is a secondary or

specialized condition.

Tlio-fibularis (Biceps)— Very strong fleshy origin from the

whole of the post-acetabular ridge of the ilium to the beginning

of the origin of the semitendinosus. The strong belly converges

to a rounded tendon which is inserted to the fibula after passing

through a sling in the usual way (text-fig. 129, BIC.). The short

arm of the sling has a strong anchor to the fibula, which I happen

to have noted in Herons, but which is present also in many birds.

belonging to widely separated groups.

Ischio-femoralis (Obdurator eaternus).—Arises by strong tendon

from external condyle of the femur (text-fig. 129, Ob.ex.) and

inserted fleshy to surface of the ischium.

Obdurator (Obdurator internus).—Origin by a strong tendon

surrounded by a gemellus muscle (text- fig. 129, OB.in.Gem.) from

the external condyle of the femur proximal to the obdurator

externus. Garrod (18) believed that in most cases the insertion

of this muscle to the inner aspect of the pubis and ischium

could be distinguished as oval or triangular, and attached some

systematic value to the condition. He described it as oval in

Steganopods and Storks and triangular in Herons; in Baleniceps

it is plainly oval.

Pub-ischio-femorales (Adductor longus and Add. nagnus).—The

external or longus is only, about half the width of the inner

or magnus, but. their origins and insertions are practically co-

extensive. In my notes i find that they were nearly equal in

Herons and Storks, but I have not paid special attention i the

point. The slips ta ‘om the magnus to the tibia (text-fig. 130, Sl.)

and the slp to the semimembranosus (text-fig. 130) I nave noe

noted in Storks or Herons.

Peroneus superficialis (longus).—Strong muscle from crest of

tibia and fascia over the tibialis anticus ; usual insertion by broad

tendon to the fascia of the ankle and a long tendon running down

to join with the tendon of the perforated flexor of the third toe.

Precisely the same relations exist in Storks and Herons, but also

in so many other birds that no systematic importance can be

attached to them.

Peroneus profundus.—A. short but stout muscle from the tibia

below the fibula; its tendon passes over the ankle-joint to be

inserted to a knob on the outer side of the tarsus-metatarsus.

- ANATOMY OF THE SHOE-BILL. 683

According to Weldon and my own notes, this muscle is absent in

Leptoptilus, but: I found it present and with an extension of its

origin to the fibula in Herons.

Tibialis anticus.—The outer head arises by a strong tendon

from the external condyle of the femur and runs in a deep groove

to join the fleshy head from the tibial erest. Insertion by a

forked tendon to the tarsus-metatarsus, in a pit about an Inch

below the joint. The conditions are practically the same in

Herons and Storks.

Text-fig. 130.

[sf K—= ary rT

Gaatrocnemius and its relations in Baleéniceps.

Ad. Adduetor magnus SI. Tendinous slips from adductor to tibia.

Isf. Ischio-flexorius (Semimembranosus).

C.11.1. Femoral head of Caud-ilio-flexorius (Accessory Semitendinosus).

C. 11.2. Belly of Caud-ilio-flexorius (Semitendimosus).

Ge. 1, external, Ge.2, middle, Ge. 3, tibial portion of Gastrocnemius.

Soleus.—This little muscle has the usual relations, but is

relatively rather stronger than in Storks and Herons; it arises

fleshy from the inner side of the tibia and is inserted to the

annular cartilage of the ancle-joint.

Extensor communis digitorum: arises fleshy from the crest and

external surface of the shaft of the tibia (text-fig. 129, Ex.com.).

‘The strong tendon passes through a bony and a fibrous bridge

and runs down to the digits where it divides symmetrically into

684 DR. P. CHALMERS MITCHELL ON THE

two, each branch again dividing into two, the four tendons

running respectively to the second digit, to each side of the third

digit and to the fourth digit. The arrangement is practically

identical in Storks and Herons.

Flexor perforans et perforatus (Flexor secundus) of digit I1.—

Origin is from the external condyle of the femur distal to the

short arm of the biceps sling (text-fig. 129, Fl. 211, p. 680) and

from the fascia over the knee-joint, immediately superficial to the

corresponding flexor of the third digit with which it is closely

connected. The tendon passes in the normal fashion to the

second digit, perforating the tendon of the flexor primus and

being perforated by the branch of the tendon of the flexor

communis.

Flexor perforans et perforatus (Flexor secundus) of digit I1I.—

This has two heads, one just deep of the corresponding flexor of

the second digit and practically common with it, and a second

from the edge of the fibula. Its tendon receives a strong slip

from the tendon of the perforated flexor (flexor primus) of its own

digit and then is inserted to digit IIT in the same fashion as the

corresponding flexor of digit II. I did not record the existence

of the second head of this flexor in my notes on Storks and

Herons, but otherwise the flexores secundi have identical relations

in Baleniceps, Storks and Herons. It must be noticed, however,

that these relations are found in a very large number of birds

belonging to different groups.

Flexores perforati (Flexores primi) of digits II, III, IV;

Rudiment of Aimbiens-—The perforated flexor muscles (text-

fig. 129, Fl. 1. 11, Fl. 1. ILI, Fl. 1. LV) are very closely united.

Distally the tendons for the respective digits separate out ;

proximally the common muscular belly arises from three distinct

heads and the arrangement is such that fibres to each tendon can

be traced to each head. The largest head is fleshy from the

intercondylar notch of the femur; there is an outer rather broad

tendinous head, superficial to the biceps tendon and arising from

the head of the fibula (text-fig. 129, Fl. 1. E). The third head is

a round and very distinct tendon, passing under the biceps tendon

and running partly te the head of the fibula and partly to the

fascia of origin of the flexores secundi (text-fig. 129, Amb.). The

inner fleshy head is normal and oceurs in practically identical

form in all birds that I have dissected. The outer tendinous

head also is usually present, and exists in Storks and Herons, the

chief differences it presents being in the extent to which it is

muscular. In Baleniceps, the tendinous portion is longer and

the muscular portion relatively shorter than in Storks and Herons,

thus showing a degenerate condition. The rounded tendon under-

lying the biceps is more interesting. In birds where the curious

muscle known as the ambiens is present, the tendon of that

muscle passes through the knee-joint, passes under the biceps

tendon, sometimes with an anchor to the edge of the fibula, and

then forms a third head of origin of the perforated flexors,

ANATOMY OF THE SHOE-BILL. 685

precisely similar in position and relations to the rounded tendon

under the biceps in Balwniceps. But for the fact that the

tendon stops short at the head of the fibula and does not pass

through the knee capsule to a normal ambiens muscle, it can-

not be distinguished from the ambiens head of the perforated

flexors. Garrod (17) first called attention to the interest of the

ambiens muscle and regarded it as a major key to the classi-

fication of birds. He divided the Class into two Subclasses, the

Anomalogonate, containing the Piciformes, Passeriformes, and

Cypseliformes in which the ambiens is never present, and the

Homalogonate, containing all the other groups of birds and

showing that in them the ambiens was normally present. Among

the Anomalogonatz there is no species in which the ambiens has

been found; among the Homalogonate there are families and

genera in which it is absent, and Garrod believed that in

such cases it had been secondarily lost. In a much later con-

tribution to the subject (24), I showed that in the Night Heron

and in Helectus, birds without an ambiens but belonging to

Garrod’s Homalogonatee, there existed an ambiens head to the

perforated flexors, absent in the Anomalogonate, and plainly

suggesting that it was a remnant of the ambiens muscle. In

a memoir on the anatomy of the Hoatzin (27) I was able to

describe from dissections of different examples of that bird, a case

of this possible degeneration in actual progress. Garrod had

dissected both legs in three examples of the bird and in all cases

found the ambiens small but normal above the knees, but in five

out of the six legs it was lost at the knee-joint. He does not

appear to have had his attention called to the importance of the

ambiens head of the perforated flexors. I examined each leg in

two examples, and found in every case an ambiens head to the

perforated flexors, but the ambiens muscle in some instances

absent above the knee, in others small and lost at the knee-joint.

It may therefore be taken as established that the ambiens head

of the perforated flexors represents a vestige of a complete

ambiens muscle, and its existence in Baleniceps is of morpho-

logical rather than systematic importance. The ambiens is

usually present in the Steganopods, present in the Spoonbills,

present in some genera of Storks absent in others, absent in

Scopus, absent in Herons and Saleniceps, but in the last two

cases its recent loss is shewn by the existence of the vestige to

which I have now called attention.

Flexor profundus or perforans and Flexor longus hallucis.—The

deep flexor as in Storks and Herons arises by fleshy digitations

(text-fig. 129, Fl. Pro.) from the side of the fibula and from down

the shaft of the tibia to form a strong round tendon, The flexor

longus hallucis comes from the inner surface of the outer condyle

of the femur (text-fig. 129, Fl. Hal.) and similarly forms a round

tendon. The two tendons pass down to the flexor surface of the

foot in the usual way. The deep flexor (text-fig. 131, B) breaks

up into a branch for digits 2, 3, 4, and the hallucis tendon,

686 DR. P. CHALMERS MITCHELL ON THE

crossing over the profundus, runs to the hallux, but sends a long

slender branch which joins the profundus tendon just before

that divides for the digits. The condition corresponds with what

Gadow (16, p. 195) calls type I. The hallucis tendon crosses

over the profundus to reach the hallux, but sends a vineulum to

it. In Storks and Herons the condition is essentially simular, but

in the former group the vinculum is stronger and may be in

separate slips; in Scopws and the Herons the vinculum is much

more slender and may be absent. For comparison | figure the

condition in a Stork (text-fig. 131, 8), a Heron (text-fig. 131, A),

and in Baleeniceps (text-fig. 131, B).

Text-fig. 131.

£5

f¢

—

Roe

Diagram of Deep Flexor Tendons in A, Nyecticorax; B, Baleniceps :

8, Leptoptilus.

The longus hallucis tendon is in outline, the flexor profundus is shaded.

1. Hallux. 2,3 &4, 2nd, 3rd & 4th digits.

Popliteus.—There was only one of these little muscles stretch-

ing across between the head of the fibula and the tibia. In

Leptoptilus I noted two.

SUMMARY OF MuscutAar ANATOMY.

Garrod’s hope, excited by his extraordinarily interesting pioneer

work, that muscular anatomy would furnish a sure clue to the

classification of birds has not been fulfilled. Garrod relied chiefly

on the presence or absence of certain muscles which he found to

vary from group to group. Gadow, who has attempted on a

complete scale to apply to the system Garrod’s group of muscles,

using the additional facts made known by Beddard and other

writers, appreciated that as these muscles were a common heritage

of all birds, the presence of any of them in any group of birds

could not be taken as a guide to the systematic position of that

ANATOMY OF THE SHOE-BILL. 687

group. He was disposed, however, to attach value to the loss of

any of these muscles, and accordingly regarded the loss of this or

that muscle as one of the characters to be employed in judging of

the relationships of groups. Even this cautious use seems to me

to be going too far. At present I do not know of any reason

why we é should suppose that a particular muscle may not have

been lost independently many times; that is to say of any reason

why a bird that has lost its femoro-caudal muscle should be more

nearly related to another bird with a similar loss than to a bird

which has retained the possession once common to all three. The

loss is what I have described as a multiradial apocentricity.

Possibly when we know as much of the development and mor-

phology of the muscles used by Garrod, as Fiirbringer has taught

us in the case of the shoulder and wing muscles, we shall be able

to make more definite use of muscular anatomy in systematic

ornithology. As, however, muscular anatomy has been used

freely, I may give a summary, of the chief facts from which more

confident anatomists would draw inferences

Comparison of Herons, Storks, Scopus and Baleniceps.

Peculiar to Baleniceps.

Absence of latissimus dorsi metapatagialis. (? Scopus.)

Absence of humeral head of biceps brachii. (Unique.)

Absence of deltoides minor. (¢ Scopus.)

Absence of teres minor. (¢ Scopus.)

Origin of serratus superficialis posterior confined to dorsad of

uncinate processes. (Same in Phenicopterus.)

Accessory origin from tip of scapula of serratus metapatagialis.

Common to Laleniceps and Scopus.

Condition of deltoides patagialis and patagial tendons.

Expansor secundariorum vestigial or absent (so also in most

Steganopods).

Common to Baleniceps and Herons.

Presence of peroneus profundus.

Ambiens reduced to a distal vestige (said to be absent in

Scopus, present in Storks).

Deep flexor tendons.

Common to Laleniceps and Storks.

Pecuhar arrangement of tendon of insertion of latissimus

dors. post.

Practical doubling of pectoralis major (also in some Steganopods).

Presence of post- acetabular portion of gluteus maximus (at

least in some Storks; ? Scopus).

Oval origin of obdurator internus (also in most Steganopods).

Common to Baleniceps, Scopus, Herons, Storks and most Stega-

nopods (but also in many other groups).

Absence of biceps slip to patagium.

Absence of accessory femoro-caudal.

688 DR. P. CHALMERS MITCHELL ON THE

Clearly, so far as the evidence from muscular anatomy goes,

Baleniceps is an ally of Scopus, Herons and Storks, and shares

many characters with these birds and Steganopods. The two

facts that impress me most are the close similarity of the rather

highly specialized patagial muscle and tendons in the case of

Baleniceps and Scopus, and the very remarkable condition of the

pectoralis thoracicus found in Baleniceps and Storks, but which

probably also exists in Scopus, and certainly in Pelecanus and

some other Steganopods.

OstTEOLOGICAL NOTES.

As the osteology of Baleniceps has been described at length

in W. K. Parker’s well-known monograph (83), and as I have

not the time at my disposal to make the elaborate study of

the different types of Storks and of Herons which is necessary

before final conclusions can be drawn from the skeleton of

Baleniceps, | must content myself with a few notes on some of

the salient points which struck me as requiring special study.

Occipital condyle.—In Baleniceps this, seen from in front and

below, has much the appearance of a moderately dolicocephalic

human cranium. It is sessile, elongated antero-posteriorly, and

its posterior margin, where it projects slightly into the foramen

magnum, is convex. In the Herons the condyle is transversely

elongated, and the posterior margin, where it projects into the

foramen magnum, 1s the broadest part, is concave and slightly

erooved, as if to form the beginning of two condyles. In Scopus

the condyle is also transversely elongated but not so much as in

Herons, and its foraminal margin is abruptly truncated. In

Anastomus, Dissura, Xenorhynchus, and Ciconia the long axis is

transverse, and the foraminal margin is concave or notched. In

Tantalus, on the other hand, the condyle is nearly spherical, and

although a notch may just be indicated, the foraminal margin is

convex. The configuration of the condyle in Lalwniceps is, there-

fore, unlike Scopus, Storks or Herons, but it is most nearly

approached by the Z'antalws Storks.

Paroceipital processes.—The broad, thin and shell-like pro-

cesses which bend down over the articulation of the quadrate

are repeated on a smaller scale in Z’antalus, where however the

lamine are relatively thicker and less extensive. In other Storks

they are replaced by similarly situated, thick and curving ridges.

In Scopus and Herons all appearance of the shell-like arrange-

ment is absent.

Basitemporal plate.—In Baleniceps the anterior margin of this

has a erescentic free edge which nearly meets at each side a curved

lamina projecting from the basisphenoid, so that the Eustachian

tubes are nearly floored in below. Parker wrote that ‘in the

Heron these parts are essentially a miniature” of those in Bale-

niceps, adding that this ‘is certainly not a faint and superficial

mark of affinity.” But the similarity extends to Scopus and

ANATOMY OF THE SHOE-BILL 689

Storks. In Herons the anterior margin is slightly pointed, and

in Scopus and the Storks, including Zantalus, it is rather more

sharply pointed, but the general relations and particularly the

relations to the laminze from the basisphenoid, which I shall now

describe, are more like those of Laleniceps in the case of Storks

than of Herons.

Basisphenoid.—Seen from below this has the usual “T-shaped”

appearance, the cross bar of the “'T” being contiguous with the

anterior edge of the basitemporal, the main limb forming the

rostrum. In Baleniceps delicate crescentic lamelle project back-

wards nearly meeting the front edge of the basitemporal plate and

with it forming a floor for the Eustachian tubes. The condition

of these lamine in Zantalus most closely resembles that of

Baleniceps; in the other Storks the laminz are less complete,

and they are least complete in the Herons*and Scopus. The

rostrum from the “T” cross bar to the attachment of the

pterygoids is a stout, broad beam of nearly equal width through-

out its length in Laleniceps. The other birds in the set 1 am

considering present a series ranging from aleniceps through

Tantalus, the typical Storks, and Scopus, to the Herons which

present the end of the series most remote from Laleniceps. The

rostrum gradually in the series changes from an even beam to

a sharply contracting, almost triangular outline, and its smooth

ventral curved surface becomes first slightly ridged, and then

strongly carmate as in Herons.

Orbital septum.—This is completely ossified in Baleniceps,

Scopus, all the Storks, including Zantalus; very incomplete in

the Herons, including Cancroma.

Lacrymal.—As Parker has described, the lacrymal of Baleniceps

is highly peculiar, although no doubt'the peculiarity is partly

adaptive in relation to the enormous beak. It is a stout vertical

strut forming the anterior wall of the orbit, firmly anchylosed

below with the jugal and maxilla, and above with the nasal. On

the roof of the skull it forms the external portion of the fronto-

maxillary hinge which runs as a transverse suture across the

forehead, being thus entirely anterior to the hinge. A thin

vertical lamina projects from it into the cavity of the orbit, which

is pierced by a large lacrymal foramen, external to the nasal

cavity. In Scopus the lacrymal is a vertical beam scooped out

on its orbital face for the lacrymal canal, but hanging down

freely along the front of the orbital cavity, until it almost meets

but does not actually touch the jugal. It has no contact with

the maxilla and depends from the orbital edge of the frontal,

behind the fronto-maxillary hinge, and with no more than the

minutest overlap to the nasal on the distal aspect of the hinge.

Jn Storks of the genus Vantalus the lacrymal is suspended from

the orbital edge of the frontal behind the hinge, with just a

trace of overlap across it to the nasal. From this point of

suspension the flat external face hangs vertically downwards,

gradually narrowing, and free from the maxilla and not reaching

690 DR. P. CHALMERS MITCHELL ON THE

the jugal below; it has a stout lamina projecting into the orbital

cavity transversely to the long axis of the skull and pierced for

the lacrymal canal. In the typical Storks and in Anastonus,

the external face of the lacrymal is roughly triangular, the convex

basal line being attached to the orbital edge of the frontal, but

definitely extending forwards across the hinge to form a very

loose connection with the nasal, not more than a fifth of the

whole dorsal edge of the bone. From this, the rapidly narrowing

triangle hangs down in front of the orbit and is far from reaching

the jugal and has no connection with the maxilla, The inwardly

projecting flange is a very thick beam pierced for the lacrymal

canal. In the Herons, the relations are a still further exaggera-

tion of the difference between Storks and baleniceps. The outer

surface of the lacrymal is triangular with a very broad base of

attachment to the frontal behind the hinge, a small loose overlap

to the nasal in front of the hinge, and with no connection with

jugal or maxilla. The inwardly directed flange is absent and

there is no lacrymal canal.

In the case of the lacrymal bone, therefore, Baleniceps and

Ardea stand at opposite ends of a series, Zantalus being nearest

to Baleeniceps.

Nasal region.—The nostrils in aleniceps are impervious, a

stout nasal septum being developed. Gadow states that they are

pervious in Scopus, but this is a mistake; a very thin lamina

of bone very slightly fenestrated separates them. In all the

Herons and Storks they are pervious, the cavity from one nostril

to the other being large and quite open. The skull is holorhinal

as in Scopus, Ardea, Cancroma and all the Storks including

Tantalus, but in many, especially the larger Storks, the proximal

end of the nasal bone shows a line of weakness running up

towards the naso-frontal hinge and leading to the schizorhiny

seen in /bis. From the anterior border of the nostril a groove

runs along the surface of the beak to the extreme anterior

end, only the hook of the beak projecting beyond it. Beddard

appears to lay some stress (4, p. 484) on this point because he

says that the groove is “precisely like that of Scopus and Can-

croma.” It is like that of Scopus, but in Cancroma the groove

is much wider and more shallow and does not reach the extreme

anterior end. It is much more exactly repeated in the Pelican,

the Ibis and the Flamingo. In Ardea it is represented by a groove

which runs about half-way from the nostril to the anterior end

of the beak; and in most of the Storks it is represented by a

line of weakness in the bone reaching about half-way to the tip

of the beak. This is specially well marked in Z'antalus.

The nasal processes of the premaxille are so firmly fused with

the adjacent nasals that their exact outline cannot be seen. It

is clear, however, that they do not invade the frontal region but

terminate distad of the fronto-maxillary hinge. This also is the

ease in Cancroma and Storks, including Z'antalus, but in Ardea

ANATOMY OF THE SHOE-BILL. 691

and Scopus the nasals break the transverse line of the hinge,

fitting into the frontals.

The anterior tip of the premaxilla is produced in Daleniceps

to form the strong down-turned hook of the bill. Scopus repeats

this on a smaller scale ; in Cancroma the sharp point is not bent

into a hook. In the other Herons and in Storks it is straight.

Palate—The palate is desmognathous in Salwniceps, the

maxillo-palatines being fused in the middle line, and the vomer

represented by a triangular, very thin, ossification, the knife-like

base of which divides the internal nares when seen from below.

The condition in Scopus is almost identical, but at its proximal

end, where it touches the central lamine of the palatine, the

rather larger vomer shows a broader edge with the faintest

suspicion of doubling. In the Storks, the vomer is relatively

smaller even than in Laleniceps and there is no trace of forking.

In Cancroma and Ardea the vomer is relatively very much larger

and its edge is quite distinctly cleft between the palatines, each

blade being attached to the palatine lamina contiguous with it.

Tn this respect Baleniceps and the Herons are at the opposite

ends of the series. With regard to the palatines, the most

striking feature in Laleniceps, fully described by Parker, is the

coalescence of the internal laminz to form a strong keel stretch-

ing back from the posterior nares to the pterygoid articulations.

Allowing for differences in shape and proportion, the similarity

with Scopus is close. The median keel is still more strongly

marked in the Pelican and in Plotus; it is represented in Storks

by a delicate median ridge, but in Cancroma and Ardea the

internal lamine of the palatines remain completely separate.

Pterygoids.—I notice no significant. differences between the

pterygoids of Daleniceps and those of Scopus, Storks and Herons,

Basipterygoid articular processes are absent in all, and I have

not found even any rudimentary trace such as is common in the

Pelican. The ventral distal end of each pterygoid is smoothly

rounded in Laleniceps and Ardea; in Cancroma, Scopus and

most of the Storks it shows a sharp keel running out as if to

meet the outer lamina of the palatal.

Quadrate.—This is substantially alike in Balewniceps. Scopus,

Herons and Storks, but the orbital process in Balwniceps is almost

triangular, the blunted apex projecting into the orbital cavity.

In Scopus the orbital process is rather blunter ; in Zantalus move

acute, but in Storks generally it tends to expand to a spatulate

end, and in the Herons, including Cancroma, the apex is much

expanded.

Quadratojugal bar.—This is enormously stout in Baleniceps,

and the separate portions of which it is composed cannot be dis-

tinguished. In all the other birds I am considering, it forms a

slender, much elongated rod.

Temporal cavity.—The boundaries of the temporal cavity

present interesting modifications in baleniceps and its aliles.

Proc. Zoot. Soc.—1913, No. XLVI. 46

692 DR. P. CHALMERS MITCHELL ON THE

Unfortunately, owing to the very complete union of the bones

concerned, these cannot be interpreted completely without the

examination of very young skulls, but comparison of adult skulls

shows a good deal worth noting. In Baleniceps the size of the

brain is small in proportion to the size of the bird, with the

result that the skull is very short between the orbit and the

posterior end, Attachment for the powerful temporal muscle is

increased by the strength and size of the postfrontal process

(Pl. TOOL, fig. 1), which depends as a triangle of bone behind

the orbit, with the external face strongly ridged. The blunted

apex of the triangle reaches nearly half-way down to the quadrato-

jugal bar and is continued to a strong tubercle on the latter by a

fibrous band. The central portion ‘of the band bas a separate

ossification, and it would not be surprising to find, in an old bird,

that calcification of the fibrous band had joined this central ossi-

fication with the postfrontal above and the jugal below, to form

a complete beam of bone separating the orbital and temporal

cavities externally. Owing to the complete fusion of the bones,

it is impossible to be certain as to the exact composition of the

postfrontal. A more primitive skull like that of Dromews (in

which the proportion of the temporal region to the rest of the

skull closely resembles that in Laleniceps) shows that the ali-

sphenoid contributes the main portion of the postfrontal, and

that the frontal grows down over it only about half-way. In

Baleniceps it appears as if the frontal covered the alisphenoid

right down to the lower end of the postfrontal, and on the

poster ior face the alisphenoid may itself be covered by a process

of the squamosal. On the other side of the temporal cavity in

many birds the squamosal sends forwards and downwards from

just over the quadrate articulation a stout beam of bone p1o-

jecting tow ards the point of the postfrontal. In LBaleniceps this

is represented only by a narrow edge pr ‘ojecting over the quadrate

articulation. Scopus (Pl. TOOL, fe, 2) has like Baleniceps also

a small brain and narrow temporal space. The postfrontal has

almost exactly the relations of that of Laleniceps, but it does not

reach nearly so far towards the jugal. The spur of the squamosal

is minute, and allowing for differences in the strength of the

muscular attachments, this region is almost the same in Scopus

and Baleniceps.

In a large Stork like Yenorhynchus (Pl. LXXXI. fig. 3) there

is an arrangement strikingly different in appearance but which,

none the lass, can be interpreted easily. The brain is still small

and the temporal cavity narrow. The postfrontal triangle narrows

very rapidly and is continued downwards as a slender bar which

stops short long before the jugal is reached. Clcse serutiny seems

to show that “the frontal contributes a superficial splint-lke

factor, running down almost to the tip on the anterior face, and

that the squamosal forms the greater portion of the lower and

posterior part, but how much Saye alisphenoid contributes it is

impossible to say. The spur of the squamcsal from over the

ANATOMY OF THE SHOE-BILL, 693

quadrate articulation is enormous and forms a powerful process

which runs downwards and forwards to meet and fuse with the

tip of the postfrontal. Ina smaller skull, like that of the Dissura

(Pl. LXXXIT. fig. 1), the arrangement is practically identical,

but the postfrontal, although it meets the squamosal spur, is

much more slender, and the frontal factor does not appear to

reach more than half-way down,

In Tantalus (Pl. LXXXII. fig. 2) the brain is relatively

shghtly longer and larger. ‘The squamosal spur is exactly as in

NXenorhynchus and Dissura, but two things have happened to

the postfrontal. In the first place it is much shorter, and does

not reach the squamosal spur, In the second place the triangular

base is very much wider and is deeply notched in front, with the

result that it has an anterior and smaller portion corresponding

exactly to the anterior margin in all the other birds I have been

describing here, but certainly with no squamosal factor, and a

longer portion running down towards the point of the squamosal

spur, corresponding with the posterior part of the postfrontal

in other birds and certainly consisting chiefly of alisphenoid and

squamosal factors.

The condition in Cancroma (Pl. LX XXII. fig. 3) can now be

followed easily. The brain is still larger relatively ; the squa-

mosal spur is reduced, and the separation between the two parts

of the postfrontal, only just apparent in Zantalus, is well marked.

The purely frontal, anterior portion is the stouter of the two.

In Ardea (Pl. LX XXIII. fig. 1), where again the brain is still

larger, the squamosal spur is relatively rather small, but the

separation between the two parts of the postfrontal is very

wide indeed.

As this matter appears to be of some interest, and as I

have not found it discussed, I shall continue the description

outside the immediate relatives of Baleniceps. The Pelican

(Pl. LX XXIII. fig. 2) shows a further extension of the series.

The squamosal spur is as in Ardea, but the two portions of the

post-frontal are even further separated, and the posterior of the

two is reduced to a mere tubercle, intermediate in position be-

tween the squamosal spur and what would normally be taken to

be the postfrontal. Im Plotus (Pl. LX XXIII. fig. 3), which has

a very large brain indeed, the squamosal spur is small, there is a

mere stump to represent the posterior portion of the postorbital

process and this is actually nearer the squamosal spur than

it is to the anterior representative of the postfrontal.

I do not suggest that the series, as I have arranged it, is

phylogenetic, but it is a striking example of the differences that

identical morphological material may exhibit in allied birds, and

a warning against the hasty drawing of conclusions as to sys-

tematic position from the comparison of one or two presumably

allied birds. So far as this point goes, Baleniceps and Scopus

stand together as birds with small brains, with the squamosal

spur slight and the postfrontal process simple. Storks form a

46*

694 DR. P. CHALMERS MITCHELL ON THE

second group also with small brains, with the squamosal spur

very highly developed, frequently reaching the postfrontal, and

with the latter simple, but in Zantalus showing the beginning

of cleavage. Cancroma and the other Herons form a third

group, characterized by larger brains, with the squamosal spur

reduced as in the first group but with the postfrontal split into

two separate processes of which the anterior tends to become the

more important.

Mandible.—Vhe fusion of the component parts is so complete

that I could not see any trace of sutures. The most notable

feature is the absence of the projecting spur of the angle, an

absence which Baleniceps shares with Scopus, all the Storks and

Cancroma, in all of which the end of the mandible behind the

articular cavities for the quadrate is as if abruptly sawn off,

while in Ardea and typical Herons it is produced backwards as

a long rounded spur.

Vertebral column.—As Parker (33) and Gadow (16, p. 76) have

pointed out, there are 17 cervical vertebre in Baleniceps, 16 in

Scopus, 17 or 18 in Storks, and 18 to 20 in Herons. The carotid

canal is complete in all this group of birds. The individual

vertebree are very much shorter antero-posteriorly in Laleniceps

than in the Herons and Scopus; the Storks are intermediate

between Laleniceps and Herons in this respect, which no doubt

is purely adaptive, but it suggests at least that Baleniceps is

not very closely related with Herons. Parker states that there

are no foramina for the vertebral arteries on the sides of

the atlas in Baleniceps and in its allies. They are certainly

absent in Baleniceps, but present, although small and limited to

the anterior lateral part of the atlas, in Ardea, Cancroma and

Scopus, and present and large in Storks.

The thoracic vertebre have no hemapophyses in Baleniceps,

Cancroma, Ardea and Xenorhynchus, but there is a very small

unpaired process in Tantalus.

Sternum.—The posterior lateral processes are very long in

Baleniceps, projecting well behind the metasternum ; in Scopus,

Herons and Storks, they are short, not projecting behind the

metasternum. ‘The notch separating the posterior lateral process

from the metasternum is rounded in Saleniceps, Scopus and

Storks, angular in Ardea and Cancroma. Parker figured a small

posterior intermediate process, and compared it with that of the

Ibis, but it was completely absent in the skeleton I examined, as

well as in Herons, Storks and Scopus.

The internal spine of the sternum 1s absent in Baleniceps as in

Scopus, Storks and Herons, but the external spine is also absent

in Baleniceps, small in Scopus, small or absent in Storks (best

developed in Vantalus), large and prominent in Herons including

Cancroma.

Furcula.—In Baleniceps the clavicles are joined to form a

very short-stemmed ‘“‘ Y,” the stem of the Y as well as the distal

ends of the diverging arms being strongly anchylosed to the

ANATOMY OF THE SHOE-BILL. 695

projecting anterior end of the keel of the sternum. There is no

trace of a median process opposite the hypocleideum between the

arms of the furcula. Anchylosis with the keel may be regarded

as an adaptive character. It is as complete in the Pelican as in

Baleniceps: it exists 1 some of the large Storks, but in Storks

and Herons generally the connection is by ligament. In Scopus

the furcula is far short of reaching the keel. In Scopus, Storks

and Herons including Cancroma, the furcula is more ‘‘U ”-shaped;

the median forward process between the arms of the “U” is

absent in Scopus and Storks as in Baleniceps, but is well-marked

in Herons.

The proximal end of each clavicle where it reaches the coracoid

has a very strong flange (text-fig. 132, A, Cl.a.) which articulates

Text-fig. 132.

Shoulder-girdle Articulation in Baleniceps and Scopus.

A. Baleniceps. B. Scopus.

Ac. Acrocoracoid process of Coracoid.

Cor. Coracoid.

Pe. Procoracoid process of Coracoid.

Cl. Clavicle:

Cl.a. Acrocoracoid process of Clavicle.

Sc. Scapula.

directly with the anterior border of the coracoid. A similar

flange is present in Scopus (text-fig. 132, B, Cl.a.), and in its place

there is a minute flat articular facet in Zantalus, but it is absent:

in Storks generally and in Herons including Cancroma, the

clavicle being attached to the acrocoracoid only by strong liga-

ments. The development of this flange in Baleniceps is a very

strongly marked character, but too much weight cannot be laid

on this similarity with Scopus, for the acrocoracoid flange of the

clavicle is equally well marked in the Pelican, in Plotus, in

Cormorants and Gannets, and in Birds-of-Prey.

696 DR. P, CHALMERS MITCHELL ON THE

The epicleideal plane of the clavicle passes across the median

face of the acrocoracoid and nearly reaches the scapula, with

which, however, it is united only by ligament (text-fig. 132, A, CZ.).

Its tip, however, does not extend so far as to cross the acro-

coracoid,fand it is therefore wholly concealed when it is viewed

from the outer side, the opposite aspect from that shown in text-

figure 132. In Scopus the tip of the clavicle projects slightly

further. In all the Storks and Herons that I have examined the

tip of the clavicle projects completely across the acrocoracoid so

that its hgamentous connection with the end of the scapula is

freely visible when the shoulder-girdle is viewed from the outside.

Coracoid.—The basal ends of the coracoid, where they articu-

late with the sternum, do not quite meet in Baleniceps ; they

either do not meet, or just meet in Storks ; in Scopus and Herons

they overlap.

The procoracoid process (text-fig. 132, A, Pe.) is very large in

Baleniceps, with a curved border nearly reaching an emar-

gination in the clavicle, with which, however, it is united only

by ligament. This process varies in Storks, being almost as

large but not expanded in some, small in others (e.g. Tantalus).

It is quite small in Scopus (text-fig. 132, B, Pc.) and Herons

including Cancroma.

Pelvis.—Parker lays some stress on the narrowness of the

pelvis in Baleniceps, comparing it in this respect with Herons

and contrasting it with Storks. Certainly such a difference does

exist, but as it was not so striking to my eye as Parker found it,:

JT measured a number of pelves of Storks and Herons, reduced

the measurements of the narrowest and widest portions of each

to a common standard and averaged the results,

Width of Pelvis in percentage of length.

Narrowest part. Widest part.

Baleniceps...... wis 20 50

PAROLE) Viera: = eyes Ne 25 50

Cancroma ......... 28 57

NXenorhynchus ... 35 60

Tantalus ......... 3D 56

ISCOPUS relents sees oF 62

It will be seen that the width at the widest part differs much

less than the width at the narrowest part. In other respects I

find that the pelvis of Baleniceps is rather more like that of

Scopus and Storks than of Herons. There is a well-marked

notch separating the distal ends of the ilium and ischium in

Baleniceps, Scopus, and Storks which is absent in Herons, in-

cluding Cancroma. In Herons, including Cancroma, the dorso-

lateral edge of the post-acetabular ilium projects outwards as a

sharp horizontal ridge, which is very slightly marked in Scopus

and is absent in Baleniceps and Storks.

ANATOMY OF THE SHOE-BILL. 697

Humerus.—A good deal of attention has been paid to the

anterior surface of the proximal end of the humerus in birds, and

I have compared the conditions presented by Baleniceps, Scoprus,

Storks and Herons. Saleniceps is markedly different in this

respect from the others and resembles much more closely the

structure shown by the same bone of the Pelican, which has

been adequately figured by Pyeraft (35, fig. 6, p. 90). The

expanded end of the humerus is very flat and rather symmetrical.

The coraco-humeral groove is extremely faint, rather better

marked than in the Pelican but entirely different from the deep

transverse groove shown by Scopus, Storks and Herons. The

evista inferior is hardly raised above the surface; only the

faintest distal groove separates it from the shaft of the bone.

In the Pelican it is much more salient; Pycraft’s figure rather

under-represents its prominence. In Herons the crista is rather

like that of Laleniceps, but more prominent; in Scopus it is still

more prominent, and in Storks, including Z'amwtalus, it is even

more prominent. ‘The pectoral crest is still longer and better

marked in Laleniceps than in the Pelican, but is generally

similar, and in both the oval impression for the insertion of the

second division of the pectoral is very clear. There is no trace

of thisin the Herons. In the Storks, including Zantalus, it is

well-marked. This region of the humerus in Scopus is charac-

terized by the enormous development of the pectoral crest, which

is very much larger than in any other of the birds with which I

am dealing here, and there is a faint oval impression which

certainly seems to imply the existence of a divided pectoral in

that bird, as in Storks and baleniceps (see supra, p. 676).

Hand.—The hand of birds is usually described as containing

the representatives of three digits, of which that corresponding

with the index finger (on the usually accepted supposition that

the pollex is present) is the largest. The proximal phalanx of

this digit has usually a broad flange on the ulnar side on which

one or more of the primary quills rest. In dissecting this region

in Baleniceps | was struck by the way in which the arrangement

of muscles suggested that this phalanx represented the phalanges

of two adjacent digits united by a narrow sheet of bone. On

examining the skeleton in Laleniceps itself, Scopus, Ardea and

a number of Storks, the suggestion is still more striking. The

broadened phalanx 1s obviously thickened along the ulnar and

radial borders, and if these borders were the phalanges of two

digits united by an ossified sheet of fibre the structure would be

intelligible. The point requires investigation, both by com-

parative anatomy and embryology, and I mention it here, only

to direct attention to it, but I shall be surprised if it does not

turn out that the hands of these and many other birds show four

not three digits.

Tibial bridge.—The bony canal for the tendon of the extensor

muscle of the digits is complete in Laleniceps as in Scopus, Storks

and Herons.

698 DR. P. CHALMERS MITCHELL ON THE

Tarsal bridges.—The hypotarsus is complex in Baleniceps,

Scopus and Herons; that is to say the proximal end of the shaft

is provided with two bony bridges through which the flexor

tendons pass; in Storks these bridges are absent, and the tendons

lie in a groove.

SUMMARY or OsTEoLoGICAL Nores.

There is a strong general resemblance between Baleniceps, on

the one hand, and Scopus, Herons and Storks on the other.

Baleniceps stands alone in its general proportions, in the struc-

ture and relationship of the lacrymal, quadratojugal bar, atlas,

posterior lateral process of the sternum, and the humerus.

Baleniceps and Scopus agree, opposed to the others, in the

impervious nostrils, the union of the inner plates of the palatines,

she processes bounding the temporal cavity, the acrocoracoid

articulation of the clavicle, and the shortness of the tip of the

clavicle.

Baleniceps, Scopus and Storks agree, opposed to the others, in

the basi-temporal plate, the complete interorbital septum, the

very small vomer, the shape of the posterior notch of the ster-

num, the absence of a spina interna, and of a median process

between the diverging arms of the furcula, and the presence of

a notch separating the posterior extremities of the ischium and

lium.

Baleniceps and Storks agree, opposed to the others, in the

large procoracoid, and the smooth edge of the post-acetabular

ium. Of the Storks, Z’antalus shows the closest agreement with

Baleniceps in the occipital condyle, the paroccipital processes

and the basisphenoid.

Laleniceps and Herons and Scopus agree, to the exclusion of

the others, in the proportions of the pelvis, and the complex

hypotarsus.

T have been unable to find any osteological points in which

Baleniceps agrees with Herons to the exclusion of the others.

Sysvematic Posrrion or BAL-ENICEPS.

John Gould (22) named and diagnosed this bird in 1851. He

relied on external characters and compared it carefully with

Pelecanus, Grus, Ardea and Cancroma. He came to the con-

clusion that it was the “Grallatorial type of the Pelecanide.”

He rejected alliance with Ardea and Cancroma apparently because

in the latter the nail of the central toe is pectinated, a character

which he thought to be wholly absent in Baleniceps. I have

shown (supra, p. 648) that much reliance cannot be placed on

this character. Undoubtedly there are many marked anatomical

characters common to baleniceps and the Pelican. I may

mention the presence of a pyloric chamber of the stomach ;

the division of the great pectoral muscle; the condition of the

ANATOMY OF THE SHOE-BILL, 699

tendons of the wing; the absence of intrinsic muscles in the

syrinx ; the long lacrymals; the fusion of the internal lamine

of the palatines; the shell-like paroccipital processes ; the anchy-

losis of the clavicle to the keel of the sternum; the shape

of the head of the humerus. But these can probably be best

explained either as convergent modifications in birds which,

after all, are not very far apart im the system, or the common

inheritance of Steganopods and their immediate alhes. Sub-

sequent writers on Baleniceps appear to have been dominated

by the wish to prove Gould incorrect. J. Reinhardt (37) in

1860 came to the conclusion that the nearest ally of balew-

niceps was Scopus, and that Scopus and Baleniceps together

were nearer the Storks than the Herons. He appreciated that

the pterylosis, especially of the neck, was similar in Laleniceps,

Scopus and Storks and markedly differed from that of Cancroma

and the other Herons. He attached importance to the pectina-

tion on the claw of Cancroma and the Herons: noted that

there was an approach to it in Scopus, but did not recognize

that it also occurred in Laleniceps. He noted that the inter-

orbital septum was complete in Scopus and Storks, incomplete in

Cancroma and Herons, but had not information on that point

in the case of Baleniceps. He compared the bills in considerable

detail, and pointed out essential points in which baleniceps and

Scopus agreed, and differed from Cancroma. Parker (33) read his

great paper before Bartlett’s contribution, but as the latter was

actually published first, it is more correct chronologically to take

Bartlett (1) first, although, curiously enough, the authors quote

one another, and each relies on the other’s opinion. Bartlett in

1861 made the interesting discovery that Laleniceps had a pair

of large powder-down patches on the back, and this for him

settled the Ardeine aftinities of the bird. He included, however,

Hurypyga amongst the Herons, and I have shewn (supra p. 645)

that in the present state of our knowledge, the presence of

powder-down patches is no conclusive evidence as to affinity.

T cannot understand why it has been assumed and stated by

writers who had an opportunity of reading the memoir, that

W. K. Parker’s osteological examination proved LDaleniceps to be

a Heron and net a Stork. In the Introduction to his memoir, he

made the following general statement :—‘‘ It is to the stilted,

wading group of scavengers that Baleniceps belongs, being one

of the Ardew affines, and therefore intimately related to the

White Stork, the Marabout, and the Adjutant. Its nearest

relations, however, are the South American Boat-bill (Cancroma

cochlearia) and the Little South African Umbre (Scopus wmbretta).”

In a footnote to his paper he refers to Bartlett’s discovery as

“having proved beyond al! dispute, that the Balaniceps, like the

Boat-bill, is essentially a Heron.” But these are ex cathedra

statements. He also gives a list of what he regards as ‘“‘ Ardeine

genera ”;—including Ciconia, Leptoptilus, Mycteria, Anastomus,

Aramus, Ardea, Botaurus, Herodias, Nycticorax, Scopus,

700 DR. P. CHALMERS MITCHELL ON THE

Cancroma and Baleniceps. He selected the Heron as the type-

form as ‘it is best known, and has the characters of the family

moderately, but markedly developed.” In his detailed examin-

ation, he is quite as concerned to prove that Laleniceps is like

Scopus or like a Stork as like a Heron. He came to the very

definite conclusion that Baleniceps was not a Pelican, but was

Axdeine, in his broad sense of the word. Quite certainly he did

not prove it to be a Heron as opposed to a Stork; and in this

I entirely agree, after having gone through his own observations

with the skeletons before me, and having made some further

comparisons myself.

C. G. Giebel (21) in 1873 re-opened the matter, chiefly after a

full examination of the pterylosis and external characters. He

corrected Gould and Reinhardt (not quoting the latter, however)

as to the pectination, describing it carefully and showing that it

was present in aleniceps 11 much the same form as in Scopus.

He showed also that these birds agreed in pterylosis, where

they differed from Cancroma, and came to the conclusion that

Baleniceps was much more closely allied with Scopus than with

Cancrona.

¥. KE. Beddard in a special memoir and in his treatise on

the anatomy of birds (3 and 4) assumed that Parker’s osteo-

logical researches had proved Baleniceps to be a Heron, not

a Stork, overlooked the evidence supplied by Reinhardt and

Giebel as to the Stork-like pterylosis of Baleniceps and Scopus,

and although he referred to Giebel’s paper, did not note that

Giebel showed Baleniceps and Scopus to be alike in the peetin-

ation of the claw. He placed Baleniceps definitely among the

Herons as opposed to the Storks, relying apparently chiefly

on the syrinx. I have already stated (supra p. 651) that his

argument cannot support his conclusion. Possibly it may show

that Baleniceps is not a Stork; it does not show that it is

a Heron.

H. Gadow (16) placed both Baleniceps and Scopus under the

Ardez as opposed to the Ciconie, but placed Baleniceps merely

as a sub-family of the Herons, distinguished from the true

Herons by the absence of pectination and various minor characters.

He was misled, I think, by the literature, and does not claim to

have made independent observations.

Those who have followed the old and new facts regarding

Baleniceps that I have been able to bring together must be

impressed by the number of characters in which Scopus and

Baleniceps agree, and by the much greater number of points in

which these two birds agree with Storks than with Herons. If

we adopt the method made familiar by many of those who have

written most copiously on the anatomy of birds, take characters

on their face value, and regard those birds as most nearly allied

which have the greatest number of characters in common, then

the sub-order Arde will contain the single family Ardeide, and

the adjacent sub-erder Ciconie will comprise the families Scopide,

ANATOMY OF THE SHOE-BILL. 701

Balenicepidide, Ciconiide and Ibidide. But if we desire that

classification should represent phylogeny, we must await further

knowledge as to the value of the various characters which

anatomists have tried to use. It is no use bringing pectination

or powder-down patches, or a completely feathered neck into an

argument between Storks and Herons, if these characters have

been independently acquired or independently lost in the case of

many different groups. It is no use for systematic purposes to call

a particular form of syrinx Ardeine if that be a common type in

many different kinds of birds, or to call a doubled pectoral muscle

Ciconine if that be shared by many Steganopods. The fact is

that Steganopods, Scopus, Baleniceps, Storks and Herons have a

large common heritage, consisting partly of actual common

structures (some of which they share with a very much larger

assemblage of birds) and partly of the capacity to be modified

in certain definite directions. ‘The characters latent and patent

composing this common heritage are distributed irregularly

amongst them, partly for reasons that we do not know and partly

in response to similar habits. Until the meaning and history

of each set of characters have been worked out very fully we have

less than no idea as to their real value in indicating affinity.

The only set of characters on which I have sufficient knowledge to

have any confidence is shown in the disposition of the alimentary

tract. In that respect Baleniceps seems to me to have passed

through the condition common to Storks and Herons, and to

display specialization in the same direction as the Herons. But

until we have further knowledge, not so much of Baleniceps in

particular, but of the value of anatomical characters, the safe

course is to regard Laleniceps as the representative of a group of

equal value with Storks and Herons.

In my opinion, however, the relation of Storks and Herons

to the Steganopods requires revision. Pending this, I must add

that John Gould’s description of Laleniceps as the “ Grallatorial

type of the Pelecanide ” is at least as happy as the more confident

statements of later writers.

List of References.

1. Barruert, A. D., On the Affinities of Baleniceps rex. P.Z.S.

ISOM JO. Ika sk

. BepparD, F. E., Anatomy of Scopus umbretta. P.Z.S. 1884,

p- 943.

: a Visceral Anatomy of Baleniceps rex. P.Z.S.

1888, p. 284.

The Structure and Classification of Birds.

London, 1898.

3 99 On the Alimentary Tract of Certain Birds.

JP fais HOTT Fos 24

‘ CAZIN, Maurice, L’Appareil eastrique des Oiseaux. Ann.

Sci. Nat. iv. p. 177 (1887).

Oa oa - wo WD

CO COS

10.

nts

12.

13.

14.

15.

16.

17.

18.

atts),

20.

21.

22.

23.

24.

25.

26.

27.

28.

29.

DR. P. CHALMERS MITCHELL ON THE

. Davies, H. R., Pectination. Nature, 1891, p. 367.

. Evans, A. H., Birds, in the Cambridge Natural History.

London, 1889.

. Forses, W. A., On the Bursa Fabricii in Birds. P.Z.S.

1877, p. 304.

5 On the Pterylosis of Mesites. P. Z.S. 1882,

footnote p. 269.

53 On some Points in the Anatomy of the

Indian Darter. P.Z.S. 1882, p. 208.

a The Last Journal of W. A. Forbes. Ibis,

1883, p. 494.

Fursprincer, M., Water ‘suchungen zur Morphologie und Sys-

tematik der Vogel. Amsterdam, 1888.

Gapvow, H., Versuch einer vergleich. Anatomie des Ver-

dauungssystemes der Vogel. Jen. Zeitsch.

Xl. po. 91S 79):

55 Remarks on the Cloaca and on the Copulatory

Organs of the Amniota. Phil. Trans.

elxxvul. p. 5 (1888).

Vogel in Bronn’s Thier-reich (1891).

Garrop, A, H., On certain Muscles of the Thigh of Birds,

and on their Value in Classification. P. Z. S.

1873, p. 626.

4 On the Anatomy of Chaunaderbiana. P.Z.S.

1876, p.189:

a On the Anatomy of Plotus anhinga. P.Z.S.

IRSA, JO 31330).

4, On the Anatomy of Levaillant’s Darter.

123 Agiss UiSiish, Os O08);

GieBEL, C.G., Baleniceps rew. Zeits. fur Gesam. Naturw.

vol. xli. p. 350 (1873).

GouLp, Joun, On a new and most remarkable Form in

Ornithology. P.Z.S. 1851, p. 1.

Levcxart, F. S., Ueber eine zusammengesetztere Magenbild-

ung bei verschiedenen Vogeln. Zoolog.

Bruchstiicke, ii. p. 64 (1841).

Mircuet, P. Cuatmers, On the perforated Flexor Muscles

in some Birds. P.Z.S. 1894, p. 495.

a On the Proventricular Crypts of Pseudo-

tantalus ibis. P.Z.S. 1895, p. 271.

5 On the Intestinal Tract of Birds. P.Z.S.

1896, p. 136.

a Anatomy of the MHoatzin (Opisthocomus

cristatus). P.Z.S. 1896, p. 618.

up Quintocubitalism in the Wing of Birds.

Journ. Linn. Soc., Zool. vol. xxvii. p. 210

(1899).

os Anatomy of Kingfishers. The Ibis, 1901,

Da oie

30.

31.

32.

33.

34.

35.

36.

37.

38.

ANATOMY OF THE SHOE-BILL. 703

MircuHett, P. CHatmers, On the Intestinal Tract of Birds.

Trans. Linn. Soe., Zool. viii. p. 173 (1901).

5 Anatomy of Gruiform Birds. P.Z.S. 1901,

p. 629.

f Anatomy of Limicoline Birds. P.Z.8. 1905,

p. 155.

Parker, W. K., The Osteology of Baleniceps rex. Trans.

Z.8. iv. p. 269 (1860).

PETHERICK, JOHN, Memoranda on the Hippopotamus and

Baleniceps recently imported to England.

P.Z.8. 1860, p. 195.

Pycrart, W. P., Contributions to the Osteology of Birds.

Part I. Steganopodes. P.Z.S. 1898, p. 82.

a Aquintocubitalism in the Bird’s Wing.

Journ. Linn. Soc., Zool. vol. xxvu. p. 236

(1899).

REINHARDT, J., On the Affinities of Baleniceps. P.Z.8. 1860,

p. 377.

Wetpon, W. F. R., The Anatomy of Phwiicopterus. P.Z.S.

1883, p. 638.

EXPLANATION OF THE PLATES,

Prats LXXX.

Baleniceps rex. From a living young male.

Photo by D. Seth-Smith, F.Z.S.

Prare LXXXI.

Fig. 1. Side of Skull in Baleniceps.

2. = 5 Scopus.

3. ss if NXenorhynchus.

Pratt LXXXII.

Fig. 1. Side of Skull in Dissura.

a 35 Tantalus.

3. 5 3 Cancroma.

Pirate LXXXIII.

Fig. 1. Side of Skull in Ardea.

5 pH Pelecanus.

a 5 i Plotus.

704 PROF. A. DENDY AND MR. R. W. H. ROW ON

44, The Classification and Phylogeny of the Calcareous

Sponges, with a Reference List of all the described

Species, systematically arranged. By ArrHur Denpy,

D.Sc., F.R.S., F.Z.8., Professor of Zoology in the

University of London, and R. W. Haroip Row, B.8c.,

F.L.S., Assistant Lecturer and Demonstrator in Zoology

at King’s College.

[Received May 30, 1913: Read June 8, 1913. |

(Text-figure 133.)

“ Le groupe ne se définira plus par la possession de certains charactéres,

mais par sa tendance a les accentuer.”— Bergson.

INDEX.

Page

JES faeh OND} ORGHTNICOhc He oHeeReatemnerctsonciosucdacodana sea seboanouseapas echO!

PRINCIPLES OF CLASSIFICATION ........2..-c00-020.008--. 09

SysTEMATIC ARRANGEMENT OF THE CALCAREA ...... 715

Family 1. Homoceelide, fam. nov. ..........6....00-.. 716

Baas ICUCASCLO MCR ee eee re ee Ce RIL.

oy Ob LECTORINGIEIES NENING WOW noo sonsanccassea0a60s 736

» 4. Minchinellide, fam. nov. .................. 739

» 9. Murrayonide, fam. NOV. ......-0...ccees 741

st Ie ORRASITRECLAAOLIZ AN ob Gane Semede dablouneataudebeaoosad socauain (Le

Bn. Us Sbeterapidd enact Neben cocoa Reeser 750

pe USI CU AiO HOU eRe speacn ade obsbae ana dab ban soo reebe dor: 1 ZOE.

Ge ZU UNOPISCIUE soonscaeasn ba xcesno spaa0acoo0en “hail

eel OMMelapeidcen Tam Mover seec reer ao

Genera and Species “‘ Incertze Sedis” ............00.605 786

List of rejected Generic Names ..................0..... 786

PEVOGENY OF DHE CATCAR HAS ay.t cen seecceseecheaceennn: Lod

Bir mnssry (OYE UND) 2 O'd ones Alaa aaah arp Ronan eu bod aeacadcaa oso ndodstiae, i 1s01)

INTRODUCTION.

The first Caleareous sponges, ‘‘ Spongia ciliata” and “ Spongia

compressa,’ were described in 1780 by Fabricius, but it was

not until much later that the essential differences between the

Calcareous and Non-calcareous Sponges were recognised. Fleming,

in 1828, however, proposed the genus Grantia for the former

group, including in it all the forms whose skeleton consisted of

ealeium carbonate. (Risso’s earlier genus, Sycon, and Gray’s

Seypha were diagnosed differently.)

CALCAREOUS SPONGES. 705

The next important advance in the history of the group was

the erection of the genus Leucosolenia by Bowerbank in 1866,

for certain sponges which we now include in the Homoccelidee, in

addition to other genera no longer employed. From that time

forward numerous investigators studied the group, and many

new species and several new genera were described, but no really

serious attempt to deal with the question of the classification of

these sponges was made until the time of Haeckel, who in 1870

published his ‘ Prodromus,’ and in 1872 his famous Monograph

of the group, with extremely detailed, though somewhat un-

successful descriptions of all the then known species, including

many which he described for the first time. Haeckel’s so-called

“natural ? system, with its three families of Ascones, Leucones,

and Sycones, based upon the type of canal system, and its twenty-

one genera based upon the types of spicules present, is so well

known, and has been so often criticised, that it needs no further

description by us, especially as it proved extremely artificial, and

expressed only to a very limited extent the true phylogeny of

the group.

The scheme proposed by Poléjaeff in 1883 was considerably

more successful, and his primary division of the group into

Homoca@na and Hererocata has been made the basis of almost

every Classification since proposed. Weare now beginning to

realise, however, that this division also is of a very arbitrary

character.

The next scheme of classification we need notice is that of

Vosmaer, in Bronn’s ‘ Klassen und Ordnungen des Thierreichs ”

[1887], which is almost identical with that of Poléjaeff, with

the addition of the Pharetronide as a fourth family of the

HETEROCGLA.

In 1891 von Lendenfeld proposed a modification of Haeckel’s

system, erecting a fourth family, the Sylleibide, intermediate in

canal-system between the Leucones and Sycones, and reducing

the number of genera in each family to two, according to the

presence or absence of oxea. This was undoubtedly a con-

siderable improvement upon Haeckel’s system, but again it failed

to interpret the interrelationships of the members of the group

correctly, and it has since been almost entirely abandoned, though

certain spongologists, notably Breitfuss, retained it with but

little modification for a considerable time.

During the years 1891-1893 there was published by Dendy

[1891 A, 1892 B, 1893 A| a scheme of classification based on

almost wholly different lines. Retaining Poléjaefi’s Orders

Homocana and Hererocana, and, like that author, including in

a single genus, Leucosolenia, all the species of Homocaa, he

divided the Hrrrrocaia into five families, whose differentiating

characters were based far more on the structure and arrangement.

of the skeleton than on the canal system; and although this

system has not been accepted by all writers, yet we ourselves feel

that it embodies a more natural arrangement of the group than

706 PROF. A. DENDY AND MR. R. W. H. ROW ON

any of its predecessors, and we have made it the basis of the classi-

fication here proposed.

In 1896 Minchin published a paper entitled “Suggestions for

a Natural Classification of the Asconide,” which may fairly be

said to mark a new departure in the taxonomic study of the

Calearea, in that it introduces for the first time the idea of the

position of the nucleus in the collared cell as a character of

taxonomic importance, a character which has since proved,

our opinion, to be of great value. We have not been able, how-

ever, to follow Minchin completely in those modifications of

Dendy’s classification of the group which he proposed, partly in

the paper referred to, and partly in his well-known article in

Lankester’s ‘ Text-book of Zoology ’ [1900].

In 1898 Bidder, in a paper on “The Skeleton and Classification

of Caleareous Sponges,” proposed to carry out Minchin’s ideas

with regard to the nucleus of the collared cell to their logical

conelusion, and to divide the Calearea into two great groups

acecordingly—CaLcaRoNEA and Caucingea. Although not actually

adopting this division, which we consider to be somewhat pre-

mature in the present state of our knowledge, we have ourselves

followed much the same line of cleavage. Although he accepts to

a large extent, with regard to his families, the system proposed

by Dendy [1892 B], Bidder makes certain rearrangements which

do not appear to be altogether satisfactory. He does good

service, however, in indicating for the first time the relation-

ship of Carter’s Clathrina tripodifera, for which he proposes

the genus Dendya, to Haeckel’s Leucaltis clathria (= Hetero-

pegma nodus-gordii Poléjaeft). In the same paper he discusses

the position of the crystalline optic axis of the radiate spicule

systems, and endeavours to assign taxonomic value to this

character also, but whatever may be the theoretical value of his

conclusions, which have since been accepted by Minchin [1909],

we cannot consider that such a character is of any practical use

to the systematist.

In 1908 Jenkin erected two new families, the Chiphoride and

Staurorrhaphide, supposed to be differentiated by the presence of

what he considered to be a new type of spicule, the “ chiactine,”

from all previously recognised families. The peculiarity of these

spicules was believed to consist in the orientation of the various

rays both in relation to one another and to the other parts of the

skeleton, and a special method of development was suggested for

them. Finally, in 1909, one of us (Row) still further elaborated

the ‘* Chiact Theor y, as it was called, and proposed yet another

family, the Grantillide, in which more primitive, but similar,

‘‘nrochiacts ” were supposed to be present, and which was made

by him the starting-point from which the Heteropiide were

supposed to have been derived. As we shall show later, however,

we do not now think that the spicules in question are more than

very slight modifications of ordinary ty pes, ¢ and we have abandoned

all three families.

CALCAREOUS SPONGES. 707

It is now more than forty years since any attempt was made

at a complete revision of the Calcarea, and in the interval

the number of known species has increased from 111, described

in Haeckel’s ‘Kalkschwimme,’ to 436 recognised by us at

the present time. Having recently been engaged in examining

collections of Calearea of considerable importance and extent,

we have had impressed upon us the necessity for a complete

systematic catalogue of all the known species, the descriptions

of which are scattered through an immense number of separate

memoirs. In order to supply this want we have had to go

through practically the whole of the literature of the group,

and there is not a paper, to the best of our belief, which contains

a reference to a new species which we have not seen. It is with

considerable satisfaction that we find, as a result of this work,

that the scheme of classification proposed by one of us twenty

years ago [Dendy, 1892 B] is still applicable in its essential

respects, and requires comparatively little revision in order to

bring it up to date.

In the present memoir we propose to give diagnoses of all the

families and genera employed, and to enumerate all the known

species under the genera to which we assign them. We have

decided to confine ourselves to the consideration of living forms,

since our knowledge of fossil sponges is at present so unsatis-

factory, and the number of described species so great.

We have had access, during the course of our work, to a large

amount of material, comprising a large proportion of the known

species. This has consisted chiefly of the collections in the

Natural History Department of the British Museum, a large

collection brought by one of us from Australia (see Dendy

[1891 A and 1892 B]); the collections made by Mr. Cyril Cross-

land in the Red Sea (see Row [1909]), Zanzibar (see Jenkin

[1908 A]), and Cape Verde (see Thacker [1908]); a collection

made by the ‘Sealark’ Expedition in the Indian Ocean (see

Dendy [1913]); and the magnificent collection made by the

Hamburg South-Western Australian Expedition in 1905 (see

Row P1913 MS.]). In addition, one of us (Row) visited Berlin

and Jena in 1912, and Sotned valuable information from the

study of type specimens at those places.

With regard to the difficult question of synonymy, and the

limits which should be assigned to each species, we have, never-

theless, been obliged to rely in most cases upon the published

descriptions and determinations of the authors responsible, and

we have unfortunately found that these descriptions are fre-

quently very inadequate; while even where they are more com-

plete, the fact that they have been prepared from the point of

view of a different scheme of classification has sometimes made

them difficult to use. We have, however, assumed that all

descriptions are correct, except in cases where they have been

shown by subsequent authors, from an investigation of type

Proc. Zoou. Soc.—1913, No. XLVII. AT

708 PROF. A. DENDY AND MR. R. W. H. ROW ON

specimens, to be erroneous. Further, we have felt that we could

not, merely on the basis of these published descriptions, under-

take the responsibility of determining whether or not a species

described as new by its author should more properly have been

allocated to a previously known species. Thus we have proceeded

on the principle that all species described as new. must be con-

sidered to be distinct, unless they have been proved to belong to

a previously known species by the subsequent reinvestigation of

type specimens or otherwise. We have only given such synonyms

and references as seem absolutely necessary, but where one species

has been merged in another by any authority, and this result

accepted by us, we give the synonym and the appropriate

authority under the species in which it is now placed.

It is thus highly probable that, of the 486 species which we

enumerate, a considerable proportion will ultimately prove to be

identical with one another. This question, however, can only be

decided by a very thorough study of the specific characters and

range of variation in each case, probably necessitating in many

cases a reinvestigation and comparison of the original types.

Professor Minchin, in his paper on ‘“'The Characters and

Synonymy of the British Species of Sponges of the genus Lewco-

solenia” [1905], has set a good example of the manner in which

this critical revision of the group ought to be carried out.

We have, as a generai rule, taken no notice of varieties as

distinct from species, but certain of Haeckel’s so-called “ Specific

Varieties,’ to which he has already given distinctive names,

appear to us, after careful consideration of his descriptions, to

deserve to rank as separate species.

In the preparation of this paper we have throughout borne in

mind the requirements of the systematist, and it is hoped that

its publication will greatly assist the determination and arrange-

ment of species in the future. We have had to set aside a great

number of published generic names as synonyms, but it is hoped

that the appended list will enable the student to trace them in

the present system.

We have indicated in the list of species under each genus that

species which we recognise as the type, our method of procedure

being to take, in the case of old genera, that species, of those

which we assign to the genus, to which the name of the genus

was first applied, while in the case of new genera we have

chosen as our type the species which seems to exemplify best

the special characters on which we have founded the genus.

As the publication committee of the Zoological Society has

decided against the use of brackets around the names of authors

of species in all cases, it must be understood that the authors

cited are responsible for the specific but not necessarily for the

generic names employed by us.

_ CALCAREOUS SPONGES. 709

PRINCIPLES OF CLASSIFICATION.

It appears to us that the chief point to be borne in mind im

attempting to arrive at any natural system of classification is the

importance of utilising as many characters as possible. A classifi-

cation based upon a small number of characters must necessarily

be arbitrary and artificial, and characters which are of great

importance in some cases may be of comparatively little use in

others. here can be no doubt that there are certain significant

characters which do indicate genetic relationships, but these

characters are by no means the same in all cases, and they have

to be carefully sought for and distinguished amidst a host of less

important features. Sometimes it is the canal system that affords

the best clue, sometimes the arrangement of the skeleton, some-

times the form of the spicules, and sometimes even the position of

the nucleus of the collared cells. There must also be remembered

the undoubted fact that the phenomenon of convergence has

played a large part in the evolution of the Calcarea, and has led

in many cases to totally deceptive resemblances, as, for example,

between the genera Leucetta and Leucandra.

It seems likely, however, that the collared cell, or choanocyte,

which itself is by far the most characteristic histological con-

stituent of the sponge organisation, may ultimately prove, as

suggested by Bidder [1898], to afford a means of dividing the

whole of the Calcarea into two main branches, one having the

nuclei of these cells placed basally, and the other having them

apical. Indeed, the acceptance of this principle, if only in a

tentative manner, constitutes the chief difference between our

present views on the subject and those which we previously held ;

but in the present state of our knowledge it is a principle which

must not be pushed too far, and we have only been able to make

use of it as subsidiary to more easily determined characters. The

acceptance of this principle, however, necessitates the wide sepa-

ration of the Lelapiide from the other Pharetronid sponges with

which they have hitherto been associated.

We shall discuss this question in some detail later on, but it

may serve a useful purpose if we give at once a list of all the

species of Calcarea in which we have been able to determine the

position of the nucleus, and the results of our determination.

This list includes no less than 75 species, as follows :—

Nuclei apical. Nuclei basal.

Family Homoca@Lipa.

Leucosolenia bella Row. Leucosolenia falcata Haeckel.

Leucosolenia complicata Montagu, Leucosolenia stolonifer Dendy.

de Minchin. Leucosolenia ventricosa Carter.

Leucosolenia lucasi Dendy. Leucosolenia gardinert Dendy.

47°

710 PROF, A. DENDY AND MR. R. W. H. ROW ON

Nuclei apical.

Nuclei basal.

Family Homoca@1i1p&% (continued).

Leucosolenia variabilis Haeckel, Leucosolenia coriacea Montagu,

fide Minchin.

Leucosolenia depressa Dendy.

Leucosolenia cavata Carter.

Leucosolenia pelliculata Dendy.

Leucosolenia proxima Dendy.

Leucosolenia pulcherrimaDendy.

Leucosolenia vitrea Row.

Dendya tripodifera Carter.

Ascute uteoides Dendy.

Family Levucascip#&.

Leucascus simplex Dendy.

Leucascus insignis Row.

Leucascus clavatus Dendy.

Leucetta chagosensis Dendy.

Leucetta expansa Row.

Leucetta microraphis Haeckel.

Leucetta prolifera Carter.

Leucetta pyriformis Dendy.

Pericharax heteroraphis Polé-

jaeff.

Pericharax pexiza Dendy.

Family Levcautip&.

Leucaltis clathria Haeckel.

Leucettusa dictyogaster Row.

Family MINcHINELLID2.

Minchinella lamellosa Kirk-

patrick.

Family MurRAYoNIDs.

Sycon boomerang Dendy.

Sycon cartert Dendy.

Sycon gelatinosum de Blainville.

Sycon gigantewm Dendy.

Sycon lendenfeldi Row.

Sycon ramsayt von Lendenfeld.

Sycon raphanus O. Schmidt.

Sycon setosum O. Schmidt.

Sycon verum Row.

Murrayona phanolepis Kirk-

patrick.

Family Sycerrips.

CALCAREOUS SPONGES. 711

Nuclei apical. Nuclei basal.

Family HErerRopiips,

Grantessa erinaceus Carter.

Grantessa hastifera Row.

Grantessa hispida Carter.

Grantessa poculum Poléjaeff.

Grantessa polyperistomia Carter.

Grantessa sacca von Lendenfeld.

Grantessa intusarticulata Carter.

Heteropia glomerosa Bowerbank.

Heteropia simplex Row.

Vosmaeropsis dendyt Row.

Vosmaeropsis depressa Dendy.

Vosmaeropsis macera Dendy.

Vosmaeropsis primitiva Row.

Vosmaeropsis wilsont Dendy.

Family GRANTUD.

Grantia compressa Fabricius.

Grantia genuina Row.

Grantia vosmaeri Dendy.

Grantia indica Dendy.

Teichonopsis labyrinthica Carter.

Grantiopsis infrequens Carter.

Ute syconoides Dendy.

Ute spiculosa Dendy.

Synute pulchella Dendy.

Leucandra hispida Carter.

Leucandra australiensis Carter.

Leucandra echinata Schuffner.

Leucandra meandrina von Lendenfeld,

Leucandra minima Row.

Leucandra phillipensis Dendy.

Leucandra thulakomorpha Row.

Aphroceras cataphracta Haeckel.

Family AMPHORISCIDA.

Amphoriscus oblatus Row.

Leucilla australiensis Carter.

Leucilla princeps Row.

Family LELAPIID#.

Lelapia australis Gray.

With the exception of this important feature, which in the

main harmonises very well with our previous conclusions, the

principles that we have followed in arriving at the classification

set forth in the present paper are almost exactly those which

712 PROF. A. DENDY AND MR. R. W. H. ROW ON

were expounded by one of us some twenty years ago | Dendy

1891 A, 1893 A], as the following review of our present position

will show.

The canal system, including the form of the flagellate chambers,

is, we are convinced, of comparatively little taxonomic value in the

higher Calcarea. In the lower forms it necessarily determines the

arrangement of the skeleton, which must he in the walls of the

ascon tubes, however these may be arranged, and in the Sycet-

tide the arrangement of the radial tubes has undoubtedly been

the determining factor in the development of the articulate tubar

skeleton. With the appearance of a definite dermal cortex, how-

ever, the arrangement of the skeleton begins to vary more or less

independently of the canal system, so that with an identical

eanal system we find such different types of skeleton as that

of the Grantiidee, the Heteropiude and the Amphoriscide. In

each of these families, while the type of skeleton remains fairly

constant, the canal system ranges from syconoid to leuconoid, or

at least sylleibid. The syconoid type is again met with on a

totally different line of descent in the homocel genus Dendya,

and also in Lewcaltis, and again gives rise to a leuconoid type

both in the Leucascide and the Leucaltide.

Our view that it is the canal system rather than the skeleton

that has repeatedly undergone convergent evolution is strongly

supported by the distribution of the different types of spicules

and of the two types of collared cells. The form of the spicule,

however, must be used with great caution as a guide to genetic

relationships, for it is largely a question of adaptation. The tri-

radiate is undoubtedly the fundamental spicule form in the group,

but one might almost say that it tends to become quadriradiate

on the slightest provocation. Thus we almost invariably find

quadriradiates in the gastral cortex, whose inwardly directed

apical rays are undoubtedly of great value as a protection against

enemies, such as small crustaceans, approaching through the

osculum. Then, again, the ordinary triradiates of the dermal

cortex not infrequently develop a more or less conspicuous,

centripetally directed, apical ray ; and this latter tendency appears

to have led, in the case of the Amphoriscide, to a constant

skeletal character which forms the most characteristic feature of

the group. Similarly with regard to the distinction between

equiangular and sagittal triradiates, we find that the latter can

always be developed, when the situation in the sponge demands

this form, by the bending back of the oral rays during growth.

This nearly always takes place, for example, in the oscular collar,

where there is no room for the oral rays to extend forward at the

asual angle.

Whether or not there is a fundamental difference between an

equiangular triradiate, however its rays may be bent, and an

alate one in which the primitive oral angle is really different

from the paired angles, and in which there is a correspond-

ing difference in the position of the crystalline optic axis, as

CALCAREOUS SPONGES. 713

maintained by Bidder and Minchin, is another and much more

difficult question to decide. The presence of regular equiangular

triradiates seems most certainly to be very characteristic of the

Leucascid-Leuealtid line of descent, and we have made use of it

as one of the distinguishing features of the members of those

groups. It is, however, extremely difficult in practice to distin-

guish between a sagittal spicule which owes its sagittal character

merely to the backward bending of the oral rays, and one which

is sagittal owing to a real inequality between the primitive

angles.

There can be no question that a superficially sagittal condition

may be attained in different ways, and one of the most interesting

results at which we have arrived in the preparation of the present

paper is that the so-called subdermal sagittal (pseudosagittal)

spicules of the Heteropiide have a quite different origin from

the ordinary sagittal form, the basal ray not being homologous

in the two cases.

With regard to Jenkin’s [1908 B] supposed families Chiphorid

and Staurorrhaphide, we have come to the conclusion that these

are based upon purely imaginary distinctions. It will be re-

membered that Jenkin maintained that in these families a special

type of spicule, the ‘“chiactine,” constitutes the first (or only)

joint of the tubar skeleton. It seems highly improbable, from

purely @ priori reasons, that this joint should be differently con-

stituted in different syconoid sponges. As a matter of fact, no

one, so far as we are aware, has demonstrated how it arises in

ordinary cases, such as Sycon or Grantia, but everybody has been

content to speak of it as being composed of subgastral sagittal

triradiates. It is, moreover, well known that these triradiates

may develop an apical ray, as they do in many species (e. g.

Sycon ensiferum, Sycon verum and Grantia indica), and thus

become quadriradiates. Jenkin thought that he had demon-

strated that in certain cases the subgastral quadriradiates arise

by rotation and re-orientation of the basal rays of gastral quadri-

radiates, owing to the pressure of the developing chambers, and

regarded this mode of origin as distinguishing his so-called

chiactines from other subgastral tri- and quadriradiates. As,

however, he made no attempt to show how the subgastral tri-

radiates amd quadriradiates arise in other cases, this distinction

cannot be regarded as having any value; and we ourselves are of

opinion, from our own observations, that the spicules which con-

stitute the first joint of the tubar skeleton probably arise in the

same way in all cases, although we are not disposed to accept

without further evidence the exact mode of origin described by

Jenkin.

It is certain that, in some cases at any rate, the spicules at the

growing margin of the osculum have not yet assumed their defi-

nitive or jentacian but exhibit a confused arrangement. Some of

them gradually become oriented as dermal cortical spicules, with

their three rays lying parallel to the surface, others as subgastral

714 PROF. A. DENDY AND MR. R. W. H. ROW ON

spicules, with their basal rays centrifugally directed and their oral

rays lying in the deeper part of the gastral cortex. The spicules

of the other joints of the tubar skeleton, on the other hand,

appear to arise in the walls of the chambers themselves, and this

difference in mode of origin probably accounts for the difference

which undoubtedly exists between them and the subgastral

spicules.

We consider that the abolition of the families Chiphoride and

Staurorrhaphide will effect a much needed simplification in the

classification of the group, and also that it is highly desirable to

do away with them from the point of view of practical con-

venience, for to draw a real distinction in practice between a

so-called chiactine and an ordinary subgastral sagittal quadyri-

radiate is quite impossible. The bending of the apical ray,

whereby it is brought to he nearly or quite in the same straight

line as the basal ray, is merely a question of degree, as may be

seen from the examination of the apical rays of ordinary gastral

quadriradiates in various species.

The family Grantillide, proposed by one of us [ Row, 1909]

bas, of course, also been abandoned by us. The rejection of

Jenkin’s chiact theory and the fact that we attach little import-

ance to the mere presence of subdermal quadriradiates, have

removed both the characters on which the family was founded,

and it has consequently been merged in the Heteropiide.

The presence, however, of subgastral sagittal spicules (tri-

radiates or quadriradiates) appears to be very characteristic of

the Sycettid as contrasted with the Leueascid-Leucaltid line of

descent.

The distribution of oxea in the Calcareous sponges presents an

extremely difficult problem, as species possessing them occur side

by side with species that lack them in almost all the large genera

throughout the group. Asa result we have found it impossible

to assign to this character any such important place in our scheme

of classification as previous authors have suggested, though as a

matter of practical convenience we have used it asa basis for

arranging the species of a genus in sections. Certainly the ability

of some sponges to produce oxea may be looked upon as differ-

entiating them, at any rate to some extent, from others which

either have lost this power, although descended from oxea-bearing

ancestors, or else have never possessed it.

Further, we have found that two types of oxeote spicules

can be distinguished—the comparatively large, usually radially

arranged form, and a much smaller for which we employ the term

‘microxea.’ In typical cases the latter are less than 0-1 mm. in

length, and they are usually of a very definite hastate shape,

with an enlargement at a short distance from the distal extremity.

They thus form very characteristic and well-defined skeletal

elements, and it is remarkable to find them recurring in so many

perfectly distinct genera, belonging to most of the families within

CALCAREOUS SPONGES. 715

the group. It would obviously be impossible, in any natural

systematic arrangement, to associate together all the species

which possess microxea, for these species differ amongst them-

selves in nearly every other respect. It is noteworthy that in

some cases these spicules occur in very small numbers, so that

they might easily be overlooked, while in others they are ex-

tremely numerous; and it seems by no means impossible that

some individuals of a species may possess them while others do

not. Nevertheless, as a matter of convenience, we have decided

to make use of the presence or absence of these spicules for the

purpose of distinguishing sections of genera.

In this connection, however, it must be observed that we do not

include, in our conception of the term microxea, those long, hair-

like spicules frequently found surrounding the osculum, or some-

times echinating the surface. These we believe to be merely

slightly modified or imperfectly developed large oxea, and we

include them under that head, under the term ‘ trichoxea.’

One is tempted to explain the sporadic distribution of oxea by

speculations which, in the present state of our knowledge, are

perhaps unjustifiable; but we may perhaps venture to suggest

that the presence of oxea constituted a characteristic feature of

some remote ancestor, and that the faculty of producing them has

never been entirely lost, but requires special genetic conditions

of which we know nothing before it can become active in any

particular species. It is quite possible that our sections are

somewhat artificial, but a grouping of the species by easily

recognisable characters, especially in the larger genera, can hardly

fail to be of use to the systematist.

Further discussion of the principles of classification may con-

veniently be left until we come to deal with the various sub-

divisions of the group.

SYSTEMATIC ARRANGEMENT OF THE CALCAREA.

Class and Order CALCAREA.

Diagnosis. Sponges in which the spicules are composed of car-

bonate of lime (calcite), and consist of either triradiate or

quadriradiate systems, or are oxea (monaxons).

For many years past it has been the almost universal practice

amongst spongologists to divide the class CALCAREA into two

sharply contrasted orders, viz. Homoca@ia, in which the whole

of the gastral cavity is lined by collared cells, and Hererocana,

in which the collared cells are confined to special flagellate

chambers, a practice which was first initiated by Poléjaeff in

716 PROF. A. DENDY AND MR. R. W. H. ROW ON

1883. It appears to us that the time has come, owing to our

greatly increased knowledge of the group, to abandon this primary

division, and we now propose to consider the class as consisting

of a single order only.

One reason for taking this step lies in the discovery of various

intermediate forms. Such are the species of Dendya, which in

the radial arrangement of the ascon tubes approach closely to the

Leueascid type of Hrrrrocana, and the species of Lewcascus itself,

which are but slightly modified from homoccel ancestors and

form the starting-point of a distinct evolutionary series within

the group Herrrroca:ta; while von Lendenfeld’s Homoderma

sycandra is obviously merely a Sycon in which the collared cells

persist in the central gastral cavity throughout the life of the

sponge.

Another even more important reason lies in the fact that, as

noted above, the group Hrrmroca:ta is at least diphyletic in

origin, the genus Dendya and the family Sycettide forming two

distinct starting points from which the evolution of the higher

Leuconoid forms has proceeded.

We therefore propose to divide the class CALCAREA straightway

into families, of which the first will be the Homoceelide, practi-

cally co-extensive with the Homocaa of Poléjaeff. His group

Hererocana, on the other hand, is here definitely abandoned,

being represented by the families Leucascidee, Leucaltide, Min-

chinellide, Murrayonide, Sycettide, Heteropiide, Grantiide,

Amphoriscidee and Lelapiide.

Family 1. HOMOCCELID AE nov.

Diagnosis. The whole of the gastral cavity and its various out-

growths lined by collared cells throughout the life of the

sponge. Sponge colony rarely radiate, and, if so, the central

individual retains the primitive ascon structure, with a

lining of collared cells and without a special gastral cortex.

No true dermal membrane or true dermal cortex is ever

developed.

In 1872 Haeckel proposed seven genera of ‘‘ Ascon” Calcarea,

based upon the permutations and combinations of triradiate,

quadriradiate, and oxeote spicules. ‘This constituted his so-called

“natural” system, but he also had an “ artificial” system based

upon the type of colony formation. Both systems have shown

themselves far from satisfactory in practice and have long since

been abandoned.

In 1883 Poléjaeff, recognising the extreme difficulty of sub-

dividing the group, placed the whole of the species in the genus

Leucosolenia of Bowerbank, which takes priority over all Haeckel’s

genera and is the only genus recognised by Poléjaeff in his order

Homocaa.

CALCAREOUS SPONGES. lua

In 1891 von Lendenfeld proposed a modification of Haeckel’s

“natural” system, retaining only two genera, Ascetta and

Ascandra, the former genus Jacking oxea, ane lnuien possessing

them.

In the same year Dendy, in his “‘ Monograph of the Victorian

Calcarea Homoceela,” while accepting Poléjaeff’s conclusion that

only a single genus could be recognised, proposed to divide that

genus into sections and subsections, according to the type of

colony formation and canal system. Three sections were re-

cognised, Simplicia, Reticulata and Radiata, and the Leticulata

were further subdivided into /ndivisa and Subdivisa, according to

the absence or presence of an endogastric network. Of these

sections the Radiata now constitute the genus Dendya of Bidder,

while the other two are of little value to the systematist.

In 1896 Minchin proposed to distinguish three genera of

Homocasa, Clathrina, Leucosolenia and Ascandra, and in 1900,

in Lankester’s ‘Text- Book of Zoology,’ he recognised two distinct

families, Clathrinidze and Leucosoleniide, and gave (p. 110) the

following classification and diagnosis :—

“Grave A. HOMOC(@LA, Pol., s. Ascones, H.

‘“‘Gastral layer continuous.

“ Waminy |. Cruararinip&®, Minchin. Form reticulate. Tyri-

radiate systems always present, equiangular; monaxons present

or absent. Collar-cells with nucleus at base. Larva a parenchy-

mula, Genera—Clathrina, Gray (= Ascetta, H., pars, Ascaltis, H.,

pars, etc., and Leucascus, D.); Figs. 2, 6, 7, 8; Ascandra, H.,

emend. (= Homandra, Liaf., for Ascandra falcata, H.); Dendya,

Bidder, for Clathrina tripodifera, Crtr. Faminry 2. Lruco-

SOLENUDM®, Minchin. Form erect; monaxons always present ;

triradiates, 1f present, alate; collar-cells with nucleus apical ;

larva an amphiblastula. Genera—Ascyssa, H.; Leucosolenia,

Bwk. (= Ascandra, H., pars, etc.) ; Figs. 3, 4, 5.”

In 1909, Zool. Anzeiger, xxxv. p. 230, in response to criticisms

by Hammer [1908] and Dendy as to ane position of the nucleus

in the collared cells, he emended his diagnoses as follows :—

“Class CALCAREA. Sponges with the skeleton composed of

calcite, in the form of spicules either monaxon, triradiate or

quadriradiate in form.

“Grade 1. Homocana. Calcarea with the gastral layer of collar-

cells continuous, not forming separate flagellated chambers.

“Family 1. Crarurinip&. Oscular tubes generally short,

arising as shallow vents from the network of tubes, form of the

body typically reticulate. Triradiate spicules always present,

equiangular, and with the crystalline optic axis vertical to the

facial plane of the rays; monaxon spicules present or absent.

Collar-cells with the flagellum arising quite independently of the

718 PROF. A. DENDY AND MR. R. W. H. ROW ON

nucleus, which is spherical in form, and situated at the base of

the cell. Larva a parenchymula.

“ Family 2. LeucosoLentip®. Oscular tubes long, arising as

distinct individuals from the stolon-like system of basal tubes ;

form of the body erect. Monaxon spicules always present ; tri-

radiates, if present, typically bilateral in form, with two paired,

and one unpaired angles, and with the crystalline optic axis never

vertical, but always inclined, to the facial plane of the rays.

Collar-cells with the flagellum arising directly from the pear-

shaped nucleus, which is situated at, or near, the apex of the cell.

Larva an amphiblastula.”

Without entering into a long discussion as to the theoretical

value of these diagnoses, we may point out that in some respects

they are in actual practice very difficult of application. Only

very few of the numerous described species of homoccel sponges

have been examined with reference to the mode of origin of the

flagellum in the collared cells, the nature of the larva, or the

direction of the crystalline optic axis in relation to the facial

plane of the spicule. If it were necessary to investigate these

very obscure characters in every case, the classification of the

group would indeed make slow progress.

The more obvious characters which Professor Minchin first

made use of for the subdivision of the group, viz., the equiangular

or alate character of the triradiates and the position of the nucleus

of the collared cell, together with the erect or reticulate form of

the colony, lose their value when we extend our investigations

beyond the familiar British species. The Australian species,

Leucosolenia lucasi, L. stolonifer and Ascute uteoides all have

the characteristic non-reticulate, ‘‘ Leucosolenia” form, and all

possess oxea (monaxons) ; L. stolonifer and A. uteoides, however,

have collared cells with basally placed nuclei, while in LZ. lucast

the nuclei are apical, though unfortunately the position of the

basal granule is—as is always the case in specimens preserved

without very special precautions—-indeterminable. In L. stolonifer

and ZL. lucast, again, some at any rate of the triradiates are

apparently equiangular and indistinguishable from clathrinid

spicules. The test concerning the direction of the optic axis is

far too difficult to apply accurately to be of any general value.

As to the larvee, again, not only are these rarely met with in

the Homoccelids, but Professor Minchin himself has shown that

there is a transition from the one type of larva (parenchymula)

to the other type (amphiblastula). He says’ (Lankester’s ‘ Text-

Book of Zoology,’ part i. p. 75), “The type of parenchymula

larva exemplified by Clathrina reticulum (Fig. 59, 7), affords an

easy transition to the so-called amphiblastula found in Leuco-

solentide, and in the great majority of the Hrrrrocana.”

The genus Clathrina of Gray [1867] was originally based on

the reticulate form of the sponge colony, and this is still almost

the only character which could be made use of in practice as a

CALCAREOUS SPONGES. 719

distinction from Leucosolenia, but here again we know that no

sharp separation can be drawn between the two types of external

form, for one and the same colony may be reticulate in its lower,

and non-reticulate in its upper portion.

In short, we do not think that any spongologist who has

examined a large and representative collection of Homoccelidze

would be prepared to maintain Professor Minchin’s subdivision

of the group into Clathrinidee and Leucosoleniidee. It appears to

us that the most that can be done at present is to pick out and

diagnose in a more or less satisfactory way certain more or less

isolated and well-characterised generic forms, and to leave the

vast bulk of the species in the genus Leucosolenia.

At the same time we are of opinion, as already pointed out in

the introduction, that Professor Minchin has indicated some

characters at any rate which will in the future prove to be of

very great value for taxonomic purposes, and we ourselves have

made extensive use of the position of the nucleus in the collared

cells, as roughly determined from the spirit material, in support

of our views as to the evolution of the heteroccel Calcarea. The

reason why this character appears to be of less value amongst the

Homoceelidee will be discussed in the section dealing with the

phylogeny of the group.

There can be no doubt that the Homoccelide have all been

derived from a common Olynthus-like ancestor, from which a

number of lines of descent have branched out in various directions.

Colony formation seems to have played the chief part in the

process of evolution and many different types of colony have thus

arisen.

At present we are only able to distinguish four genera in this

family, Leucosolenia, Ascute, Ascyssa and Dendya. ‘Three of

these, Ascute, Ascyssa and Dendya, are easily and clearly definable,

possessing well-marked characters, but the remaining genus,

Leucosolenia, is distinguished almost entirely by negative cha-

racters, and contains a very large number of species, presenting

a very great diversity amongst themselves, both in colony-form

and spiculation.

Genus 1. LeucosotentA Bowerbank [1864-1882].

Diagnosis. Diverticula of the gastral cavity, if any, never radially

arranged around a central tube. Skeleton composed of tri-

radiate or quadriradiate spicules, to which oxea may be

added. No uteoid dermal skeleton. Nucleus of collared

cells basal or apical.

For illustrations of this genus see Dendy [1891 A].

The external form in species of this genus ranges from simple

Olynthus-like individuals, which may be connected together by a

720 PROF. A. DENDY AND MR. R. W. H. ROW ON

basal stolon as in ZL. ducasi and L, stolonifer, to complex reticulate

colonies which have acquired by integration a new individuality

of a higher type, sometimes with pseudogaster, pseudosculum,

pseudoderm and pseudopores, as in L. ventricosa, and sometimes

with reversal of the canal system as in LZ. cavata. It may happen

that one and the same colony exhibits a reticulate structure in

one portion, and a non-reticulate in another, as in several forms

figured by Haeckel [1872]. For further information as to the

variations in the mode of colony formation the reader is referred

to Dendy [1891 A].

Haeckel’s Ascandra falcata, which we include in the genus

Leucosolenia, has been made by both Minchin [1896] and von

Lendenfeld [1891] the type of a special genus (with the generic

names of Ascandra and Homandra respectively) on account of

the peculiar appearance of the gastral layer, which is thrown into

folds in such a way that in transverse section there appear to be

present a series of shallow radial tubes. These endodermal ridges

are always supported by the large apical rays of the gastral

quadriradiates, which have either pushed the layer of collared

cells out before them as they grow, or formed a foundation upon

which the collaved cells have spread. It may be pointed out that

other Homoccelid: also show this type of structure, e.g. LZ. canar-

vensis and L. gegenbaurt. We ourselves do not consider that this

character is of sufficient importance, per se, to take generic rank ;

nor do we consider that there is any relationship between the

“pseudoradial” character thus given to the gastral layer, and a

true syconoid canal system, since, in the radial tubes of Sycon, for

example, both dermal and gastral layers are folded, while in

Leucosolenia falcata the gastral layer only is affected. Further,

embryology shows that the radial tubes of Sycon are outgrowths

from the central gastral cavity, and not formed by ingrowths

into it. Had the syconoid type of canal system originated from

some such condition as that of Lewcosolenia (Ascandra) falcata,

the sycon person would have been provided with a dermal cortex

ab initio, whereas the more primitive Sycettide have the ends

of the chambers freely projecting on the surface of the sponge.

The position of the nucleus of the collared cells in this genus

is not, so far as we are aware, correlated with any particular

type of spiculation or canal system, and we cannot, if only for

practical reasons, make use of this character by itself for sub-

dividing the genus. Asa means of grouping the various species

of the genus into sections, however, we may, as in the higher

forms, adopt the criterion afforded by the presence or absence of

oxea.

We recognise the following species as belonging to this

genus :—

Section A. Oxea present.

1. L. ama@porweEs Haeckel.

Ascandra complicata, var. ameboides Haeckel [1872].

CALCAREOUS SPONGES. 721

2. L. aneuLata von Lendenfeld.

Ascandra angulata von Lendenfeld [1891].

3. L. arAcHNotpES Haeckel.

10.

12.

13.

14.

15,

16.

Gel be

Ascandra variabilis var. arachnoides Haeckel [1872].

ARMATA Haeckel.

Olynthus pocillum Haeckel [1870], fide Haeckel [1872].

Asculmis armata Haeckel [1872].

The earlier name is a nomen nudum.

5 by

5 IU

ATLANTICA Thacker.

Leucosolenia atlantica Thacker [1908].

BELLA Row,

Leucosolenia bella Row [1913 MS8.].

BOTRYOIDES (Hllis and Solander). Type species of the genus.

Spongia botryoides Hllis and Solander [1786 }.

Leucosolenia botryoides Minchin [1905].

. Borrys Haeckel.

Ascandra botrys Haeckel [1872].

. CERVICORNIS Haeckel.

Ascandra variabilis var. cervicornis Haeckel [1872].

. CLARKIL Verrill.

Ascortis clarkit Vervill [187 iI(6

. COMPLICATA Montagu.

Spongia complicata Montagu [1812].

Grantia botryoides Lieber kithn [1859], fide Haeckel [1872].

Olynthus hispidus Haeckel [1859], fide Haeckel [1872].

Leucosolenia complicata Minchin [1905].

. CONFERVICOLA [aeckel.

Ascandra variabilis var. confervicola Haeckel | 1872].

. contorTA Bowerbank.

Leucosolenia contorta Bowerbank [1864-1882].

Ascandra contorta Haeckel [1872].

. CORALLORHIZA Haeckel.

Sycorhiza corallorhiza Haeckel [1870].

Auloplegma haeckeli O. Schmidt MS. a Haeckel [1872].

Ascortis corallorhiza Haeckel [1872].

. CoRDATA Haeckel.

Ascandra cordata Haeckel [1872].

. DENSA Haeckel.

Tarrus densus Haeckel [1870].

Nardopsis gracilis Haeckel [1870], fide Haeckel 11879}.

Ascandra densa Haeckel [1872].

722 PROF. A. DENDY AND MR. R. W. H. ROW ON

17. L. piscoveryt Jenkin.

Leucosolenia discoveryi Jenkin | 1908].

18. L. pusra Dendy.

Leucosolenia dubia Dendy |1891 A}.

19. L. ecuiata Ark.

Leucosolenia echinata Kirk | 1893).

20. L. scurnorEs Haeckel.

Leucosolenia echinoides Haeckel [1870].

Olynthus cyathus Haeckel [1870], fide Haeckel [1872].

Ascandra echinoides Haeckel [1872].

21. L. eLeanor Urban.

Leucosolenia eleanor Urban [1905].

22. L. FaBRict O. Schmidt.

Lewcosolenia fabricti O. Schmidt [1870].

23. L. ratcata Haeckel.

Ascandra falcata Haeckel [1872].

24, L, rraciiis Haeckel.

Ascortis fragilis Haeckel [1872].

Leucosolenia botryoides James-Clark [1869], fide Haeckel

[1872].

Leucosolenia thamnoides Haeckel [1870], fide Haeckel

[1872].

Haeckel’s earlier name is a nomen nudum.

25. L. nermest breitfuss.

Ascandra hexmesi Breitfuss [1896 B].

26. L. nispipisstmA Haeckel.

Ascandra variabilis var. hispidissima Haeckel [1872].

27. L. norrwa Haeckel.

Nardopsis horrida O. Schinidt MS., fide Haeckel [1872].

Ascortis horrida Haeckel {1872}.

28. L. incerta Urban.

Leucosolenia incerta Urban [ 1908).

29. L. IRREGULARIS Jenkin.

Leucosolenia irregularis Jenkin [1908 A }.

30. L. nacunosa Johnston.

Grantia lacunosa Bean MS., fide Johnston | 1842).

Grantia lacunosa Johnston | 1842).

Ascortis lacunosa Haeckel [1872].

31. L. naxa Kirk.

Leucosolenia lawa Kirk [18995].

32. L.

33. L.

34. L.

30. L.

36. L.

37. L.

38. L.

43. L.

44. L.

Proc.

CALCAREOUS SPONGES. (2433

LIEBERKUHNI O. Schmidt.

Grantia botryoides Lieberktihn [1859], fide O. Schmidt

[1862], and Haeckel [1872].

Grantia leberkiihniit O. Schmidt [1862].

Leucosolenia robusta Haeckel [1870], fide Haeckel [1872].

Ascandra lieberkiihnit Haeckel [1872].

Lucas! Dendy.

Leucosolenia lucasi Dendy [1891 A}.

MINCHINI Jenkin.

Leucosolenia minchini Jenkin [1908 B].

nitipa Haeckel.

Olynthium nitidum Haeckel [1870].

Olynthium splendidum Haeckel [1870], jide Haeckel

[1872].

Ascandra nitida Haeckel [1872].

Panis Haeckel.

Ascandra panis Haeckel [1872].

pinus Haeckel.

Leucosolenia botryoides Lacaze-Duthiers MS., fide Haeckel

[1872].

Ascandra pinus Haeckel [1872].

RETICULATA Haeckel.

Tarrus reticulatus Haeckel [1870].

Ascandra reticulum, var. reticulata Haeckel [1872].

. RETICULUM O. Schmidt.

Nardoa reticulum O. Schmidt [1862].

Ascandra reticulum Haeckel [1872].

4. SERTULARIA [Haeckel.

Ascandra sertularia Haeckel | 1872).

. STOLONIFER Dendy.

Leucosolenia stolonifer Dendy [1891 A}.

. TENUIPILOSA Dendy.

Leucosolenia tenuipilosa Dendy [1905].

Leucosolenia canariensis ‘Thacker [1908], pars, fide

Row [1909].

TENUIS Schuffner.

Ascandra tenuis Schuffner [1877 ].

VARIABILIS Haeckel.

Leucosolenia variabilis Haeckel [1870].

Ascandra variabilis Haeckel [1872].

Leuconia somesi Bowerbank [1864-1882], fide Minchin

[1896].

Leucosolenia variabilis Minchin [1905].

Zoou. Soc.—1913, No. XLVITI. 48

724

45. LL.

46. L.

48. L.

49. L.

50. L.

+ ie OP

52. L.

53. L.

54, L.

od. L.

PROF. A. DENDY AND MR. R. W. H. ROW ON

VENTRICOSA Carter.

Clathrina ventricosa Carter [1885-1886 ].

Leucosolenia ventricosa Dendy [1891 A].

Srcrrion B. Without oxea.

Acassizit Haeckel.

Ascaltis lamarchii var. agassizii Haeckel [1872].

. BLANCA Michlucho- Maclay.

Gaancha blanca Michlucho-Maclay [1868]. —

Olynthus (dc.) guancha Haeckel [1870], fide Haeckel

[1872].

Ascetia blanca Haeckel [1872].

CANARIENSIS Michlucho-Maclay.

Nardouw canariensis Michlucho- Maclay | 1868].

Nardoa sulphurea Michlucho-Maclay [1868], fide Haeckel

[1872].

Nardoa rubra Michlucho-Maclay [1868], jide Haeckel

[1872].

Ascaltis canariensis Haeckel [ 1872).

Leucosolenia nanseni Breitfuss [1896], fide Thacker

[1908].

Ascaltis compacta Schuftner [1877], fide Thacker [1908].

Leucosolenia canariensis ‘Thacker [1908].

CANCELLATA Verrild.

Leucosolenia cancellata Verrill [1873].

cAROLI Haeckel.

Ascaltis darwinii, var. caroli Haeckel [1872].

CAVATA Carter.

Clathrina cavata Carter | 1885-1886].

Leucosolenia cavata Dendy [1891 A].

CEREBRUM Laeckel.

Ascaltis cerebrum Haeckel | 1872].

CHALLENGERI Poléjaeff.

Leucosolenia challengeri Poléjaeff [1883 ].

CHARYBDMA Haeckel.

Ascaltis gegenbauri var. charybdea Haeckel [1872].

CLATHRATA Carter.

Leucetta clathrata Carter [1883].

Clathrina tripodifera var. gravida Carter [1885-1886],

jide Row [1913 MS. ].

Grantia cliftont Bowerbank MS., fide Row [1913 MS.].

Leucosolenia intermedia Kirk [1895], fide Row [1913 MS.].

Leucosolenia clathrata Row (1913 MS. ].

56.

57.

58.

59.

67.

68.

69.

. GARDINERI Dendy.

i) |

CALCAREOUS SPONGES. 72

. CLATHRUS O. Schmidt.

Grantia clathrus O. Schmidt [1862].

2 Clathrina sulphurea J. Ki. Gray [1867], fide Haeckel

[1872].

Tarrus labyrinthus Haeckel [1870], fide Haeckel [1872].

Nardoa labyrinthus O. Schmidt MS., fide Haeckel [1872].

Ascetta clathrus Haeckel [1872].

. CONVALLARIA Haeckel.

Ascilla gracilis var. convallaria Haeckel [1872].

. CORTACEA Montagu.

Spongia coriacea Montagu [1812].

Grantia multicavata Bean MS., fide Johnston [1842].

Olathrina sulphurea Carter [1871 A], fide Haeckel [1872].

Ascetta coriacea Haeckel [1872].

. DARWINIL Haeckel.

Leucosolenia darwintt Haeckel [1870].

Ascaltis darwintt Haeckel [1872].

. DECIPIENS Haeckel.

Ascaltis cerebrum var. decipiens Haeckel [1872].

. DEPRESSA Dendy.

Leucosolenia depressa Dendy {1891 A}.

. picrvowEs Haeckel.

Leucosolenia dictyoides Haeckel [1870].

Ascetta primordialis var. dictyoides Haeckel [1872 /.

. FALKLANDICA breitfuss.

Leucosolenia falklandica Breitfuss [1898 EK}.

. FLEXILIS Haeckel.

Ascettu flewilis Haeckel [1872].

Leucosolenia gardinert Dendy [1913].

. GEGENBAURI Haeckel.

Leucosolenia gegenbauri Haeckel [1870].

? Nardoa spongiosa Kolliker [1864], fide Haeckel [1872].

Ascaltis gegenbaurt Haeckel [1872].

. GoETHEL Haeckel.

Leucosolenia goethet Haeckel [1870].

Ascaltis goether Haeckel [1872].

. cractuis Haeckel.

Ascilla gracilis Haeckel [1872].

. GRANTIL Haeckel.

Leucosolenia grantii Haeckel [1870].

Ascaltis solanderti (=Ascaltis botryoides var. solanderit)

Haeckel [1872], fide Haeckel [1872].

: AR

726

71.

(DR,

73.

74.

75.

76.

17.

78.

79.

80.

Sill

82.

83.

PROF. A. DENDY AND MR. R. W. H. ROW ON

. HIMANTIA Johnston.

Grantia botryoides var. himantia Johnston [1842].

Ascetta coriacea var. himantia Haeckel [1872].

. JAPONICA Haeckel.

Ascilla japonica Haeckel [1872].

. LAMARCKIL Haeckel.

Leucosolenia lamarckii Haeckel [1870].

Aulorhiza intestinalis Haeckel [1870], fide Haeckel oe

Ascaltis lamarckii Haeckel [1872].

. LocULOSA Haeckel.

Ascetta primordialis var. loculosa Haeckel [1872].

. MACLEAYI von Lendenfeld.

Ascetta macleayt von Lendenfeld [1885 A].

. MINORICENSIS Lackschewitsch.

Leucosolenia minoricensis Lackschewitsch [1886 ].

. MULTIFORMIS Greitfuss.

Leucosolenia multiformis Breitfuss [1898 B].

. osScULUM Carter.

Clathrina osculum Carter [1885-1886].

. PEDUNCULATA von Lendenfeld.

Leucopsis pedunculata von Lendenfeld [1885 B].

. PELLICULATA Dendy.

Leucosolenia pelliculata Dendy [1891 A].

_ PHILLIPINA Haeckel.

Ascetta blanca var. phillipina Haeckel [1872].

. POTERIUM Haeckel.

Ascetia primordialis var. poterium Haeckel [1872].

Ascandra conulata von Lendenfeld MS., jide Breitfuss

[1897].

. PRIMORDIALIS Haeckel.

Prosycum primordiale Haeckel [1870].

Olynthus simplex Haeckel [1870], fide Haeckel [1872].

? Grantia pulchra O. Schmidt [1862], fide Haeckel [1872].

Nardoa arabica Michlucho-Maclay MS., fide Haeckel

[1872].

Ascetta primordialis Haeckel [1872].

. PROTOGENES Haeckel.

Ascetta primordialis var. protogenes Haeckel [1872].

Ascetta procumbens von Lendenfeld [1885 B], fide Dendy

[1891 Aj,

' CALCAREOUS SPONGES. TG

84. L. proxima Dendy.

Leucosolenia proxima Dendy [1891 A}.

85. L. PSAMMOPHILA Row.

Leucosolenia psammophila Row [1913 MS.].

86. L. putcHERRIMA Dendy.

Leucosolenia pulcherrima Dendy [1891 A}.

87. L. RosEA Kirk.

Leucosolenia rosea Kirk [1895].

88. L. saarrrarta Haechel.

Ascetia sagittaria Haeckel [1872].

89. L. sceprrum Haeckel.

Ascetta sceptrum Haeckel [1872].

90. L. spinosa von Lendenfeld.

Ascetta spinosa von Lendenfeld [1891].

91. L. stiprrata Dendy.

Leucosolenia stipitata Dendy [1891 A}.

92. L. vestcuta Haeckel.

Ascetta vesicula Haeckel [1872].

93. L. virrea Row.

Leucosolenia vitrea Row [1913 MS.].

94. L. witsont Dendy.

Leucosolenia wilsont Dendy [1891 A].

The following species are of doubtful value :—

95. L. LAMINOCLATHRATA Carter.

Clathrina laminoclathrata Carter [1885-1886].

Too imperfectly described to be recognisable.

96. L. putcHRA O. Schmidt.

Grantia pulchra O. Schmidt {1865}.

Possibly identical with LZ. primordialis Haeckel, fide Haeckel

[1872].

97. L. sponeiosa Kolliker.

Nardoa spongiosa Kolliker [1864].

Possibly identical with LZ. gegenbauri Haeckel, fide Haeckel

[1872].

98. L. SULPHUREA Gray.

Clathrina sulphurea J. Ki. Gray (non Carter) [1867 ].

Possibly identical with LZ. clathrus O. Schmidt, fide Haeckel

[1872].

728 PROF. A. DENDY AND MR. R. W. H. ROW ON

Genus 2. Denpya Bidder [1898].

Diagnosis. Sponge colony consisting of a large central individual

lined by collared cells, from which radially arranged diver-

ticula are given off. Skeleton composed of equiangular

triradiates to which quadriradiates may be added. Subgastral.

sagittal radiates never present. Nuclei of collared cells

probably always basal.

For illustrations of this genus see Dendy [1891 A].

Carter’s Clathrina tripodifera was included by Dendy [1891 A]

in the genus Lewcosolenia, as the sole representative of the

“ Radiate” section of that genus. Bidder [1898] proposed for

its reception a new genus, Dendya, and the recent discovery by

the ‘Sealark’ Expedition of a closely allied, but quite distinct,

species in the Indian Ocean seems to justify the retention of

Bidder’s genus. Unfortunately Bidder associated his genus with

Poléjaefi’s Heteropegma (= Leucaltis) in a new family Hetero-

pegmidx, of which Dendya was made the type genus. While

admitting a certain degree of relationship between Dendya and

Leucaltis, we cannot agree that this is so close as to justify

placing them in the same family, for not only is Lewcaltis

corticate, while Dendya is non-corticate, but Leucaltis is also

heteroceel, while Dendya is homoceel.

The chief interest attaching to the genus Dendya lies in its

radiate structure, which, at first sight, seems to suggest a possible

starting point for the Sycettid as well as for the Leucascid-

Leucaltid line of descent. We no longer consider, however, that

Dendya stands very near the origin of the Sycettide, from the

simplest of which it differs widely in the structure of the skeleton,

especially in the absence of subgastral sagittal radiates, in the

fact that the radial tubes tend to cinastbonn ase. and in the basal

position of the nuclei of the collared cells. The tendency of the

radial tubes to form reticulations is indeed a difficulty in the way

of separating the genus sharply from ZLeucosolenia, a fact well

illustrated by Carter’s Leucetta clathrata (= —Leucosolenia clath-

rata), which is intermediate between the two as regayds the

canal system while resembling Dendya in the presence of the

characteristic dermal tripod spicules. [ Row, 1913 MS. ]

We recognise the following species as belonging to this

genus :—

1. D. prouirerA Dendy.

Dendya prolifera Dendy [1913].

2. D. TRIPODIFERA Carter. Type species of the genus.

Olathrina tripodifera Carter |1885— 1886].

Leucosolenia tripodifera Dendy [1891 A}.

Dendya tripodifera Bidder [1898].

CALCAREOUS SPONGES. 729

Genus 3. ASGUTE nov.

Diagnosis. The diverticula of the gastral cavity, if any, never

radially arranged around a central tube. With a uteoid

dermal skeleton of colossal longitudinal oxea. Nuclei of the

collared cells (? always) basal.

As the presence of a uteoid dermal skeleton is considered to

form a good generic character in other families, we see no reason

why it should not be used in the same way amongst the Homo-

ceelidee, and therefore propose this genus for Dendy’s Lewcosolenia

uteoides, with which Carter’s Aphroceras asconoides is doubtless to

be associated. The nucleus of the collared cells is basal in the

former species, but its position is not known in the latter.

In both the known species of this genus the sponge has the

form of a group of simple ascon persons, attached to one another

by their bases, and without any anastomoses in the colony.

We recognise the following species as belonging to this

genus :—

1. A. AsconorbEs Carter.

Aphroceras asconoides Carter | 1885-1886 }.

2. A. urEoIDES Dendy. ‘Type species of the genus.

Leucosolenia uteoides Dendy [1892 C}.

Genus 4. Ascyssa Haeckel [1872].

Diagnosis. Diverticula of the gastral cavity, if any, never radially

arranged around a central tube. Skeleton consisting entirely

of oxea.

For illustrations of this genus see Haeckel [1872].

We consider the entire absence of radiate spicules as affording

sufficient reason for the generic separation of Haeckel’s two

species of Ascyssa from all the other Homoccelide.

We assign the following species to this genus :—

1. A. acurera Haeckel.

Ascyssa acufera Haeckel [1872].

2. A. TRoGLODYTES Haeckel. Type species of the genus.

Ascyssa troglodytes Haeckel [1872].

Family 2. LEUCASCIDAL Dendy [1892 B] (emend.).

Diagnosis. Sponge typically forming a massive colony, usually

with several or many oscula, but sometimes integrated into

a single individual with definite external form. Without

any large central gastral cavity lined by collared cells, but

with an exhalant canal system devoid of collared cells.

730 PROF. A. DENDY AND MR. R. W. H. ROW ON

Flagellate chambers ranging from long and possibly branched,

with a tendency to radial arrangement round the exhalant

canals, to small, approximately spherical, and scattered.

With a distinct and independent dermal membrane (or

cortex) pierced by true dermal pores. Skeleton consisting

mainly of equiangular and equiradiate spicules, which may

become sagittal at the oscular margins. Radiates of the

chamber! ayer without definite arrangement, but irregularly

scattered in the walls of the elongated chambers, or between

the small, scattered chambers. No subgastral sagittal

radiates. Nuclei of collared cells probably always basal.

This family was provided by Dendy [1892 B] for the reception

of the genus Leucascus with its two species, L. simplex and

L. clavatus. Minchin [1900] refused to recognise either the

genus or the family, and included the two species in his Clathrina,

apparently ignoring the fact that none of the known species of

Clathrina, or indeed any other homoccel sponge, possess an

independent dermal membrane or cortex. It is true that many

Clathrinas develop a pseudoderm, but this is invariably formed

from the outermost tubes of the reticulation, and therefore

includes a layer of endoderm (gastral layer). In Lewceascus, on

the other hand, the dermal membrane is formed exclusively of

ectoderm and mesoglea (dermal layer), and does not consist

merely of the outer tubes of the reticulation. Moreover, the

radiate and non-reticulate arrangement of the elongated chambers

in Leucascus indicates a relationship with Dendya rather than

with the Clathrinoid Leucosolenias.

We here extend our conception of the family Leucascide to

include, not only the genus Leucascus and allied genera with a

similar type of canal system, but also a number of species with

a leuconoid type of canal system, which we have hitherto assigned

to the genera Leucandra and Leucilla. The skeleton of these

species, in the absence of all traces of syconoid ancestry such as

subgastral sagittal triradiates, clearly imdicates a wide phylo-

genetic separation from the typical Leucandras and Leucillas,

such as Leucandra aspera and Leucilla amphora, and closely

resembles that of Lewcascus.

It will be remembered that Haeckel, in “ Die Kalkschwimme ”

(vol. 11. p. 122, [1872]), placed his Leucetta primigenia (one of the

species which was formerly assigned to Leucandra, but now placed

in the Leucascide) at the beginning of his Leucones, which he

derived directly from an Ascon ancestry. He says: “ Leucetta

primigenia, als die wahrscheinliche Stammform der Leuconen,

steht in ihre Skeletbildung der gemeinsamen Stammform aller

Kalkschwimme, der Ascetta primordialis, so nahe, dass man sie

unmittelbar von der letzteren ableiten kann.” We agree with

these views so far as the relationship to the ‘‘ Ascones”’ is con-

cerned, but we can no longer agree that the more advanced types

of ‘‘ Leucones” (such as Leweandra) have had a leucettid ancestry.

CALCAREOUS SPONGES. 731

There is certainly no indication whatever of the genus Leucetta

having passed through a syconoid stage in its evolution, as we

believe to be the case with the true Leucandras and Leucillas,

and we therefore propose to re-establish this genus for certain

leuconoid Leucascidee.

We have also placed in this family the genus Pericharax

Poléjaeff [1883], on account of the similarity of the skeleton and

the basal position of the nucleus in the collared cells, while the

genus Leucomalthe can only be placed here provisionally.

It should perhaps be mentioned here that von Lendenfeld’s

genus Leucopsis [1885 B] was also supposed by its author to

represent a ‘Transition form between Asconide and Leuconide.”

The genus, with its single species, Leucopsis pedunculata, was,

however, so imperfeetly ‘described that it is impossible to form

any definite opinion as to its systematic position; it is very

possibly merely a reticulate Lewcosolenia with well developed

mesoglea and pseudogaster, an opinion which is supported by an

examination of some of von Lendenfeld’s type specimens, which

one of us was able to make in Berlin. At all events, it appears

to differ widely from any of our Leueascide, and it has been

placed by us among the Leucosolenias.

We have been able to determine the position of the nucleus in

no less than ten species of this small family, including five species

of Leucetta, and in all cases have found it basal, a fact that affords

strong support to our view that the genus Leucetta should be very

widely separated from Lewcandra, in which the nucleus is, so far

as we are aware, always apical.

Genus 5. Leucascus Dendy [1892 B].

Diagnosis. Flagellate chambers greatly elongated, tubular, and

sometimes copiously branched.

For illustrations of this genus see Dendy [1893 A].

We recognise the following species in this genus :—

Section A. Without oxea.

1. L. rnstenis Row.

Leucascus insignis Row [1913 MS. ].

2. L. stmpLeEx Dendy. Type species of the genus.

Leucascus simplex Dendy | 1892 B}.

Secrion B. With large radially arranged oxea.

3. L. cravatus Dendy.

Leucascus clavatus Dendy [1892 B}.

CBX PROF. A. DENDY AND MR. R. W. H. ROW ON

Genus 6. Leucomaurie, Haeckel [1872] (emend.).

Diagnosis. Colony individualised, with definite external form

and large central gastral cavity opening by a large single

osculum. Flagellate chambers greatly elongated, tubular,

copiously branched. Skeleton consisting of regular radiates,

large longitudinally placed oxea scattered throughout the

sponge body and not confined to the cortex, and minute,

irregularly hastate microxea.

For illustrations of this genus see Haeckel [1872].

The name Lewcomalthe, originally applied to one of Haeckel’s

subgenera, 1s retained for his Lewcandra bomba, which presents

many peculiarities distinguishing it from the rest of the species

of that genus. Of these, the most important from the point of

view of our present classification is the very unusual type of canal

system figured by Haeckel, which shows a number of large and

very much branched flagellate chambers radiating from a central

gastral cavity. This is very different from the normal leuconoid

canal system, and is fairly similar to that of Zewcascus. On this

account, and on account of the regular triradiates of the skeleton,

we have placed this species among the Leucascide, though, it

must be acknowledged, on somewhat doubtful grounds, and the

peculiarities of its spiculation have necessitated the provision of a

special genus to receive it.

The only known species is :—

1. L. sompa Haeckel.

Leucandra bomba Haeckel [1872].

Genus 7. Leucerra Haeckel [1872] (emend.).

non Leucetta Poléjaetf [1883].

non Leucetéa von Lendenfeld [1891].

Magnosis. Canal system leuconoid, with small, spherical or sub-

spherical flagellate chambers irregularly scattered through

the chamber layer.

For illustrations of this genus see Haeckel [1872] and

Dendy [1913].

Haeckel [1872] proposed the genus Leucetta for calcareous

sponges with a leuconoid canal system and a skeleton composed

of triradiate spicules only, taking Z. primigenia for his type

species.

Poléjaeff [1883] abandoned Haeckel’s classification, but re-

tained the name Lewcetta in an entirely different sense, equivalent

to our Leucettusa, taking one of Haeckel’s species, LZ. corticata,

for the type species of his genus.

CALCAREOUS SPONGES. TBO

Von Lendenfeld [1891] used the genus Leucetta to include

“‘Teuconide with triacts or tetracts or both.”

Dendy [1893 A] included Haeckel’s species of Leucetta in the

genus Leucandra, on the ground that the mere absence of quadri-

radiate or oxeote spicules, or both, could not be regarded as of

generic significance.

Asa result of further study of the Calearea, we are convinced

that the genus ZLewcandra, as used hitherto by Dendy, is not a

natural one, but must be subdivided, although on lines different

from any that have previously been suggested, It appears to us

that certain species, namely those which we now assign to

Leucetta, have originated, quite independently of the remainder,

from the homoccl sponges through a leucascid ancestry, and

have consequently never passed through a Sycon stage in their

phylogeny. The species in question are characterised by thei

equiangular triradiates, and by the absence of any trace of the

typical Sycon skeleton.

As regards the canal system, it appears that the same course of

evolution, from a condition with elongated and more or less

radially arranged flagellate chambers, to a condition with small

scattered spherical chambers, has taken place in both cases, so

that there has been a very complete convergence between the

genera Leucettw and Leucandra as now understood by us. The

true Leucandras, however, are distinguished by more or less

distinct traces of the skeletal structure exhibited by their

syconoid ancestors.

This view, so far as our present information goes, is supported

by histological evidence ; for, as already indicated, the position

of the nucleus of the collared cells in Leucetta is basal, while in

typical Leucandras it appears to be apical.

We also include in the genus Lewcetta, as now conceived,

certain species which we have hitherto regarded as belonging

to the genus Leucilla. These species are characterised by the

development to a varying extent of an inwardly pointing apical

ray on some of the trivadiates of the dermal cortex. They are to

be distinguished from the true Leucillas on the same grounds as

those on which the other Leucettas are distinguished from the

true Leucandras, and we do not consider the mere presence of

such apical rays by itself as a sufficient justification for generic

separation.

As we are taking Haeckel’s type species of Leucetta as the

type of the genus as understood by us, we consider that we ave

justified in retaining his name for the genus, although that name

has been used in different senses by subsequent writers.

We allocate the following species to this genus :—

Secrion A. Without oxea.

1. L. cuacosEnsis Dendy.

Leucetta chagosensis Dendy [1913].

734 PROF. A. DENDY AND MR. R. W. H. ROW ON

2. L. rroripana Haeckel.

Leucaltis floridana Haeckel [1872].

3. L. INFREQUENS Row.

Leucetta infrequens Row [1913 MS. ].

4, L. microrapuis Haeckel.

Leucetta primigenia var. microraphis Haeckel [1872].

Leuconia dura Poléjaeff [1883], fide Dendy [1892 Bj.

5. L. primicenta Haeckel. Type species of the genus.

Sycothamnus fruticosus Haeckel [1870], fide Haeckel

1872}.

ae clausu Haeckel [1879], fide Haeckel [1872].

Leucetta primigenia Haeckel [1872].

Leuconia fruticosa Poléjaeff [1883 }.

Haeckel’s two earlier names are nomina nuda, as they were

not accompanied by diagnoses, and we have rejected them,

although Poléjaeff has revived one of them.

6. L. pyrtrormis Dendy.

Leucetta pyriformis Dendy [1913].

7. L. PROLIFERA Carter.

Teichonella prolifera Carter | 1878].

Leucilla prolifera Dendy [1892 B].

Leucetta prolifera Row [1913 MS. |}.

8. L. souipa O. Schmidt.

Grantia solida O. Schmidt [1862].

Leuconia nivea J. HK. Gray [1867], fide Haeckel [1872].

Leucaltis solida Haeckel [1872].

9. L. tricgona Haeckel.

Leucetta trigona Haeckel [1872].

Section B. With large, usually radially arranged oxea,

but without microxea.

10. L. cArtERI Dendy.

Leucaltis floridana var. australiensis Carter [1885-1886].

Leucandra cartert Dendy [1892 B].

Section C. With both large oxea and microxea.

11. L. expansa Low.

Leucetta expansa Row [1913 MS.}].

The following species is doubtfully assigned to Leaucetéa :—

12. L. nomorapuis Poléjaeff.

Pericharax carteri var. homoraphis Poléjaeff [1883].

The name cartert cannot be used for this species, as it is

already employed in this genus,

CALCAREOUS SPONGES. 735

Genus 8. PEricHarax Poléjaeff [1883].

Diagnosis. Sponge colony individualised, with large central

cavity (probably a pseudogaster) opening by a wide vent

and surrounded by a very thick wall. Canal system leuco-

noid, with sub-spherical, scattered, flagellate chambers, and

with subdermal cavities whose walls are supported by a

special skeleton derived partly from the inturned rays of

tangential dermal triradiates. Skeleton of the chamber

layer confused, composed of equiangular triradiates of two

very different sizes.

For illustrations of this genus see Poléjaeff [1883] and Dendy

[1913].

This genus was proposed by Poléjaeff [1883] for a species,

Pericharax carter, from Tristan da Cunha, but recent authors,

for the most part, have not accepted it. The discovery by the

‘Sealark’ Expedition of the original species, and of a new one

closely allied to it, has enabled us to make a careful study of

the question, and we are convinced that the genus is valid.

The position which we assign to it is justified, not only by the

character of the skeleton, but also, as in the case of Leucetta, by

the basal position of the nucleus of the collared cells in both

species.

The dermal triradiates in both species are very curious spicules,

with a strong tendency to irregular curvature of the rays, often

resulting in one or more of them dipping down deeply between

the subdermal cavities.

Pericharax heteroraphis (Poléjaett’s P. carteri var. heteroraphis)

must be taken as the type of the genus. The same author's

P. carteri var. homoraphis, as we have noted above, appears to be

quite distinct, and is not a Pericharax at all, but probably a

Leucetta. Pericharax poléjaevi Breitfuss [1896 A], placed by its

author in this genus on account of the presence of subdermal

cavities, is really a typical Leucandra, for the presence of sub-

dermal cavities without a special supporting skeleton cannot be

regarded as of generic importance. Poléjaeff also included in the

genus Haeckel’s Lewcandra cucumis, the position of which is

discussed by us under the genus Paraleucilla.

We recognise the following as species of this genus :—

1. P. HETERORAPHIS Poléjaef. ‘Type species of the genus.

Pericharax carteri var. heteroraphis Poléjaeft [1883].

2. P. pezizA Dendy.

Pericharax pexiza Dendy [1913].

736 PROF, A. DENDY AND MR. R. W. H. ROW ON

Family 3. LEUCALTIDA‘ nov.

Diagnosis. Sponge colony tubular and ramified, or even anas-

tomosing, with many oscula, or individualised with large

central cavity and single osculum. Wall of colony composed

of at least two distinct layers, namely, a dermal cortex with

strongly developed skeleton of tangential radiates, and a

chamber layer with a skeleton greatly reduced or even

absent. A thin gastral cortex or membrane may or may

not be present. Skeleton composed, mainly at any rate, of

equiangular radiates. No sub-gastral sagittal radiates.

Nuclei of collared cells probably always basal.

The members of this family appear to have been derived from

a Dendya-like ancestor by the development of a thick dermal

cortex with a strongly developed cortical skeleton, and the con-

sequent more or less complete reduction of the no longer necessary

skeleton of the chamber layer. As in other families of Calcarea,

the flagellate chambers range from greatly elongated and even

branched, and more or less radially arranged, to small, sub-

spherical and scattered.

Bidder [1898] has already pointed out that the nuclei of the

collared cells in Lewcaltis clathria Haeckel (Heteropegma nodus-

gordit Poléjaeff) are basal in position, and we are able to confirm

this observation and to add that they are basal also in Leucetiusa

dictyogaster Row [1913 MS. ].

The reduction of the skeleton of the chamber layer, correlated

with the development of a thick dermal cortex with a special

cortical skeleton, finds its parallel in the genus Grantiopsis

amongst the Grantiide, but in that case the syconoid ancestry

is very clearly indicated in the remains of an articulate tubar

skeleton, while in Lewcaltis the vestigial skeleton of the chamber

layer shows no indications whatever of an articulate origin.

The characters above mentioned, In our opinion justify the

close association of Leucaltis with Leucettusa, and the wide

separation of these genera from both Leucandra and Leucilla

in our scheme of classification.

We prefer to derive the Leucaltide directly from a Dendya-

like ancestor, rather than indirectly through Leucascus, because

Leucaltis still preserves the more primitive type of radial colony

formation with what we presume to be a true central gastral

cavity, while Leweascus has adopted a massive type of colony

formation in which the exhalant canals are possibly to be regarded

as pseudogastral in nature.

CALCAREOUS SPONGES. 737

Genus 9. Leucanris Haeckel [1872] (emend.).

Heteropegma Polejaeff [1883].

Diagnosis. Sponge colony tubular, ramified and anastomosing,

with many oscula. Flagellate chambers elongated and

branched, more or less radially arranged round the central

gastral cavities of the tubes.

For illustrations of this genus see Poléjaeff [1883] and Dendy

[1893 A].

Dendy [1892 B, 1893 .A] placed this genus in the family Am-

phoriscide, on account of the large subdermal quadriradiates

possessed by the only known species. He also regarded the

vestigial triradiates of the skeleton of the chamber layer as the

remnants of an articulate tubar skeleton, being misled by the

radial arrangement of the flagellate chambers. We are now

convinced that in both these respects he was wrong, and that

the genus is probably, as Bidder [1898] maintained, related

to Dendya, from which it has been directly derived without

passing through an intermediate syconoid stage.

In arriving at this conclusion we lay great stress upon the

regular and equiangular form of the triradiates of both the

chamber layer and the dermal cortex, and also upon the basal

position of the nuclei of the collared ‘cells. The genus may, in

fact, almost be regarded as a Dendya with a thick dermal cortex.

Dendy [1913] has shown that only one species can be re-

cognised in the genus. A re-investigation of the type specimen

of Haeckel’s Leucaltis clathria has convinced us that it is not

only generically, but also specifically identical with Poléjaeft’s

Heteropegma nodus-gordu, and the latter name thus becomes a

synonym of Leucaltis clathria Haeckel. As the other species of

Haeckel’s genus Leucaltas must be removed to older genera,

L. clathria must be taken as the typical species, and we are

therefore unable to retain Poléjaefi’s name Heteropegma. Again,

Carter’s Clathrina latitubulata is only a variety of Leucaltis

clathria, differing in some slight details of spiculation. Mr.

Carter seems to have been led into provisionally placing his

species in the genus Clathrina by the external form of the whole

colony, inh resembles a reticulate Clathrina on a gigantic

scale. The reticulation, however, is not composed of simple

ascon tubes as in Clathrina, but of a colony of a higher order,

with numerous true ascon tubes lying in the thickness of the

wall. Finally, Ridley’s Leucaltis bathybia var. mascarenica is

evidently, from his description, and from the slides which we

have been able to examine, nothing but the same species.

The extent to which apical rays are developed on the tangential

radiates of the dermal cortex varies greatly in different in-

dividuals,

738 PROF. A. DENDY AND MR. R. W. H. ROW ON

We recognise only the following species in the genus :—

1. L. craruria Haeckel.

Leucaltis clathria Haeckel | 1872).

Heteropegma nodus-gordii Poléjaeff [1883], fide Dendy

[1913].

Clathrina latitubulata Carter [1885--1886], jide Dendy

(L913s]:

Leucaltis bathybia var. mascarenica Ridley [1884], fide

Dendy [1913].

Leucaltis clathria Dendy [1913].

Genus 10. Leucerrusa Haeckel [1872] (emend.).

Leucetta Poleéjaeff [1883].

Diagnosis. Sponge colony individualised, with definite external

form and large central cavity opening by a large single

osculum. Canal system leuconoid.

For illustrations of this genus see Poléjaeff [1883] under the

name Leuceita, and Row [1913 MS. ].

The remarkable reticulate type of colony formation found in

Leucaltis prevents us from regarding that genus as directly

ancestral to Leucettusa, but as regards canal system the two are

related in the same way as Grantia and Leucandra.

Within the limits of the genus Leucettusa, however, we find

considerable variation with regard to the form of the flagellate

chambers. Poléjaeff’s figure of the canal system of Leucettusa

(Leucetia) vera shows the flagellate chambers in the outer part of

the chamber layer elongated and radially arranged, while those

in the inner part are subspherical and scattered, so that this

species appears to be intermediate between Lewcaliis and the

more typical Leucettusas in this respect.

Leucettusa (Leucetta) haeckeliana, on the other hand, has the

flagellate chambers all small and subspherical, and lying in the

irregular trabecule of the chamber layer, which are separated by

very wide, irregular exhalant lacune. In Leucettwsa dictyogaster

Row [1913 MS8.] the trabecule bearing the flagellate chambers

form a network which almost completely blocks up the central

gastral cavity as an altogether askeletal layer. A section of this

askeletal chamber layer, isolated from the cortex, would be almost

indistinguishable from a similar section of Oscarella, which also

has large collared cells with basally placed nuclei, so that the

possibility presents itself that Oscarella may be nothing but a

calcareous sponge which has lost the whole of its skeleton.

We also find in the genus Lewcetiusa various stages in the

development of the subdermal quadriradiate spicules, which are

entirely absent in L. corticata and L. dictyogaster, very sparse

in L. haeckeliana and large and very numerous in LJ. vera. As

- CALCAREOUS SPONGES. 739

already indicated, we no longer consider the presence of sub-

dermal quadriradiates, taken by itself, to be a character of generic

importance, being convinced that the addition of an apical ray

to a triradiate spicule may take place whenever and wherever it

may be required.

Tt will have been noticed that our genus Leucettusa is identical

in scope with Poléjaeft’s Leucetta [1883], but as we have found it

necessary to retain Haeckel’s name Leucetta for another genus,

as previously explained, we have been obliged to adopt his

subgeneric name for the group of species which Poléjaeff quite

rightly separated out.

We recognise the following species as belonging to this genus:—

Suction A. Without oxea.

1. L. corticara Haeckel. Type species of the genus.

Leucetta corticata Haeckel [ 1872).

2. L. HAECKELIANA Poléjaeff.

Leucetta haeckeliana Poléjaeft [1883].

3. L. mprrFecta Poléjaeff.

Leucetta imperfecta Poléjaeft [1883].

4, L. samBucus Preiwisch,

Leucetta sambucus Preiwisch [1904].

5. L. vera Poléjaeff.

Leucetta vera Poléjaeff [1883].

Srcrron B. With microxea but without large oxea.

6. L. DICTYOGASTER Low.

Leucettusa dictyogaster Row [1913 MS8.}.

Family 4. MINCHINELLIDA/ nov.

Lithonina Doderlein [1892].

Lithonine Kirkpatrick [1911 A}.

Diagnosis. Canal system leuconoid (in all known forms and

presumably always so). Main skeleton composed of quadri-

radiates cemented together in various ways by calcareous

cement. Apparently without subgastral sagittal radiates.

Nuclei of collared cells (probably always) basal.

We have been able, owing to the kindness of Mr. Kirkpatrick,

to examine preparations of Minchinella and Murrayona, in which

the collared cells are sufficiently well preserved to enable us to

determine the position of the nucleus. We find that this is basal

in both cases, whereas in Lelapia we find 1% to be apical. As

Proc. Zoou. Soc.—1913 No XULIX 49

740 PROF, A, DENDY AND MR. R. W. H. ROW ON

Lelapia differs widely in other characters also from both Min-

chinella and Murrayona, we are forced to the conclusion that the

so-called Pharetronid sponges are at least diphyletic in origin,

and we have removed the Lelapiide to a position in the Sycettid

line of descent. We shall, however, discuss the question further

under the head of phylogeny.

The genera Minchinella, Petrostroma and Plectroninia all

agree in the possession of a stony skeleton composed of fused

quadriradiates, and thus differ widely from Murrayona. We

therefore unite them in one family under the name Minchinellide,

while relegating Murrayona to a special family of its own.

Genus 11. Mincurnetyia Kirkpatrick [1908].

MNagnosis. Sponge lamellar, with pore-bearing chimneys on one

side and oscular chimneys on the other. The quadriradiates

of the main skeleton cemented together.into a compact

network and completely embedded in the enveloping cement.

Dermal skeleton of radiates, including tuning-fork spicules,

and oxea.

For illustrations of this genus see Kirkpatrick [1908].

The only known species of the genus is :—

1, M. LAMELLOSA Kirkpatrick.

Minchinella lamellosa Kirkpatrick [1908].

Genus 12. Perrostroma Doderlein [1892].

Diagnosis. 'The quadriradiates of the skeleton of the chamber-

layer fused together laterally by calcareous cement into a

network. Dermal skeleton of separate quadriradiates and

triradiates and bunches of tuning-fork spicules.

For illustrations of this genus see Déderlein [1897].

The only known recent species of this genus is :—

1. P. scHutzer Doderlein.

Petrostroma schulzei Doderlein [1892].

Genus 13. PLecrroninta Hinde [1900].

Diagnosis. Quadriradiates of the main skeleton with their facial

rays truncated or expanded terminally and fused end to end

with the facial rays of adjacent spicules, while the apical

rays remain free and pointed. Dermal skeleton of separate

radiates, including tuning-fork spicules, and oxea.

For illustrations of this genus see Kirkpatrick [1900 B].

CALCAREOUS SPONGES, 7A)

‘The only known recent species of this genus are :—

1. P. peansit Kirkpatrick.

Plectroninia deanstt Kirkpatrick [1911 A}.

2. P. winpE1 Kirkpatrick.

Plectroninia hinder Kirkpatrick [1900 B].

The type of the genus is P. halli Hinde [1900], a fossil species

from the Eocene of Victoria.

‘Family 5. MURRAYONIDN nov.

Murrayonine Kirkpatrick [1911 A].

Diagnosis. Canal system presumably always leuconid. Skeleton

of the chamber layer a rigid calcareous network, not com-

posed of spicules. No subgastral sagittal radiates. Dermal

skeleton composed chiefly of overlapping calcareous scales.

Nuclei of collared cells basal.

This family seems to mark the culminating point of the

Leucascid-Leucettid line of evolution, and there is no other

known calcareous sponge with a skeleton so highly specialised as

WMurrayona.

It is at any rate possible that the aspicular main skeleton of

Murrayona is derived from the cement-covered fibres of spicules

which are known to occur in many of the fossil forms, by a

gradual disappearance of the spicular core, analogous to what we

find among the Chalinine, during the evolution of the group.

It is, however, a far cry from the one to the other, and we do not

doubt that there are many other ways in which the Murrayonid

skeleton may have arisen.

Genus 14. Murrayona Kirkpatrick [1910].

Diagnosis. With a definite pore-zone in which the dermal skeleton

consists of small trivadiates. Tuning-fork spicules present

beneath the dermal scales.

For illustrations of this genus see Kirkpatrick [1910].

The only known species of this genus 1s :—

1. M. pHanoeris Kirkpatrick.

Murrayona phanolepis Kirkpatrick [1910].

4g*

742, PROF, A. DENDY AND MR. R. W. H. ROW ON

Family 6. SYCETTIDA Dendy [1892 B}.

Diagnosis. Flagellate chambers elongated, arranged radially

around a central gastral cavity, their ends projecting more

or less on the dermal surface and not covered over by a

continuous dermal cortex strengthened by tangential dermal

spicules. “Tubar skeleton articulate, with subgastral sagittal

radiates. Collared cells usually confined to the radial

chambers in the adult, and probably always with apical

nuclei.

The Sycettide, in our opinion, form the starting point of a

distinct line of evolution, embracing the great majority of the

recent heteroccel Calcarea. This view is supported by the fact

that in all the 44 species of Sycettide, Heteropiide, Grantiide,

Amphoriscidee and Lelapiidee in which it has been determined,

the position of the nuclei of the collared cells is apical.

It would appear therefore that this line of evolution must

have originated from homoccel ancestors with apical nuclei. Such

forms are known to occur amongst the simple (7. e. non-reticulate)

species of Leucosolenia, as for example L. lucasi and-L. bella.

Why the transition from the simple homoccel to the radial

heteroccel condition originally took place we can only guess.

That it was by the outgrowth of radial buds is clearly indicated,

however, by the ontogeny of the genus Sycon (compare Schulze

[1875]). No real intermediate forms are known, however, for we

can no longer consider that Dendya is on the same line of

ancestry, while von Lendenfeld’s Homoderma sycandra is already

a highly specialised Sycon with the typical Sycon skeleton, and

differing from other Sycons only in the persistence of the collared

cells in the central gastral cavity. According to Minchin the

Sycettidz have arisen from his homoccel family Leucosoleniide,

but we have already endeavoured to shew that in the present state

of our knowledge it is impracticable to distinguish this family

from his Clathrinide.

The presence of sagittal radiates in which the oral angle is

wider than the lateral angles appears to be a very constant

character of the Sycettide and their derivatives. It would seem,

further, that the occurrence of such spicules in the articulate

tubar skeleton is correlated with the development of very definite

radial tubes, but it is extremely difficult, at any rate in practice,

to distinguish such spicules individually from equiangular

radiates which have become sagittal by bending of the oral rays.

The presence of subgastral sagittal radiates is, however, especially

characteristic of this line of evolution. The oxeote type of

spicule also tends to assume more importance in the skeleton of

this family than in the Leucascidee and Leucaltide, in which

families it but rarely occurs.

CALCAREOUS SPONGES. | 143

Genus 15. Sycerra Haeckel [1872] (emend.).

Diagnosis. The radially arranged flagellate chambers always

completely separate from one another, and never possessing

tufts of oxea at their distal ends. With no properly defined

inhalant canals leading to the prosopyles.

For illustrations of this genus see Haeckel [1872] under

Sycaltis conifera and Sycetta primitiva.

This is the simplest of all the genera with syconoid canal

system, its primitive nature being shown by the absence of fusion

between the flagellate chambers and by the absence of tufts

of oxea at their distal ends, unless, indeed, this absence is due to

the disappearance of ancestral oxea.

Sycetta asconoides Breitfuss [1896 B], to judge by the published

description, apparently occupies an intermediate position between

Sycetta and Sycon, as these genera are understood by us. The

description states that the flagellate chambers are fused together

laterally, but that they have no tufts of oxea at their distal ends.

During a recent visit to Berlin, however, one of us (Row) had

the opportunity of examining one of the type slides of this

species, and found that Breitfuss had overlooked the presence of

a few tangentially placed dermal triradiates. The occurrence of

these spicules, of course, places the species in the genus Grantia in

the family Grantiide, of which it is one of the simplest forms.

We include the following species in this genus :—

1. S. contFERA Haeckel.

Sycaltis conifera Haeckel [ 1872).

2. S. primitiva Haeckel. ‘Type species of the genus

Sycetta promitiva Haeckel [1872].

3. 8. sacirriFERA Haeckel.

Sycetta sagittufera Haeckel [1872].

Genus 16. Sycon Risso [1826] (emend.).

Diagnosis. Radial chambers usually more or less united at places

where they come into contact with one another, and always

crowned distally with tufts of oxeote spicules. Properly

defined inhalant canals usually present, the outer ends of

which may be covered by a thin pore-bearing dermal

membrane without special skeleton.

For illustrations of the structure of this genus see Schulze

[1875] and Dendy [1893 A].

As pointed out by Dendy [1893 A] the most characteristic

feature of this genus is afforded by the tufts of oxeote spicules

which crown the distal ends of the radial chambers, taken in

744 PROF. A. DENDY AND MR. R. W. H. ROW ON

conjunction with the absence of a dermal cortical, as distinct:

from a tubar, skeleton. In the more specialised species a pore-

bearing dermal membrane stretches between the distal ends of the

radial chambers, covering over the ends of the inhalant canals, but

this contains no special skeleton of its own. The flagellate

chambers may also exhibit a considerable amount of branching

towards their distal extremities, but they never lose thei

elongated character and radial arrangement.

The genus is sharply distinguished from Sycetta by the presence

of the tufts of oxea at the distal ends of the chambers, less.

sharply from Grantia by the absence of a special dermal cortical

skeleton.

We include in our conception of the genus Sycon von

Lendenfeld’s genera Homoderma and Sycantha, and Jenkin’s.

Tenthrenodes antarcticus, Streptoconus australis and Hypodictyon

longstafi. We have already mentioned that Homoderma is.

merely a Sycon with persistent collared cells in the central

gastral cavity. Jenkin [1908 B] has shown conclusively that

Sycantha tenella is a typical Sycon, but has erected a new genus.

Tenthrenodes for ‘‘Sycettidee with linked chambers,” an almost

identical character with that on which the genus Sycantha was

founded; and although Dendy [1893 A] retained Sycantha on

these grounds, we no longer consider that such “linking” can be

regarded as of generic importance. We may point out here that.

Tenthrenodes scotti, the other species included by Jenkin in his

genus, is placed by us in the genus (’rantia, on account of the

presence of tangential triradiates in the dermal cortex. Strepto-

conus australis and Hypodictyon longstaffi are “ chiact”’-bearing

forms which were placed by Jenkin in his family Chiphoride.

We assign the following species to the genus :—

1. 8S. aLopecurus Haeckel.

Sycum alopecurus Haeckel [1870].

Sycandra ampulla var. alopecurus Haeckel [1872].

2. S. ampuLLA Haeckel.

Sycariun ampulla Haeckel [1870].

Sycon petiolatus O. Schmidt MS., fide Haeckel [1872].

Sycum petiolatum Haeckel [1870], fide Haeckel | 1872).

Sycandra ampulla Haeckel [1872].

3. 8. ANTARCTIOUM Jenkin.

Tenthrenodes antarcticus Jenkin [1908 B].

4, 8. arcticum Haeckel.

Sycum arcticum Haeckel [1870].

Sycon raphanus O. Schmidt [1870], fide Haeckel [1872].

Sycandra arctica Haeckel [1872].

5. S. ASPERUM Gibson.

Sycandra aspera Gibson | 1886}.

10.

Mae

16.

18.

CALCAREOUS SPONGES. 745

. AUSTRALE Jenkin.

Streptoconus australis Jenkin [1908 B].

. BARBADENSE Schuffner.

Sycandra barbadensis Schuffner [1877 ].

. BOOMERANG Dendy.

Sycon boomerang Dendy [1892 B].

. BOREALE Schuffner.

Sycandra borealis Schuffner [1877].

. CAMINATUM Thacker.

Sycon caminatum Thacker [1908].

. CARTERI Dendy.

Sycon carteri Dendy [1892].

Sycantha tenella von Lendenfeld MS., fide Breitfuss

[1897].

Sycon carter’ Row [1913 MS. ].

. CILIATUM Labricius.

Spongia ciliata Fabricius [1780].

Sycum gigantewm Haeckel [1870], fide Haeckel [18721.

Sycocystis oviformis Haeckel [1870], jide Haeckel

[1872].

Sycodendrum ramosum Haeckel [1870], jide Haeckel

[1872].

Sycandra ciliata Haeckel [1872].

. coactum Urban.

ycandra coacta Urban [1905].

. comMuTATUM Haeckel.

Sycandra coronata var. commutata Haeckel [1872].

. compactum Lambe.

Sycon compactum Lambe [1893].

. cononaAtTUM “Hillis and Solander.

Spongia coronata Ellis and Solander [1786].

Grantia ciliata Bowerbank [1864-1882], fide Haeckel

[1872}.

Sycandra coronata Haeckel [1872].

Sycon coronatum Dendy [1892 B].

- EGLINTONENSIS Lambe.

Sycon eglintonensis Lambe [1900 B}.

. ELEGANS Bowerbank.

Dunstervillia elegans Bowerbank [1845].

Dunstervillia lanzerote Haeckel [1870], jide Haeckel

[1872].

Sycandra elegans Haeckel [1872".

746

5 Se

PROF. A. DENDY AND MR. R. W. H. ROW ON

. ENSIFERUM Dendy.

Sycon ensiferum Dendy [1892 B].

Sycon ensiferum Row [1913 MS. ].

rormosuM Haeckel.

Dunstervillia formosa Haeckel [1870].

Sycandra elegans vay. formosa Haeckel [1872].

. GELATINOSUM de Blainville.

Alcyoncellum gelatinosum de Blainville [1834-1847 ].

Grantia virgultosa Bowerbank MS8., fide Haeckel [1872].

Sycandra alcyoncellum Haeckel [1872], fide Dendy

[1892 B].

Sycandra arborea Haeckel [1872], fide Dendy [1892 B].

Sycon gelatinosum Dendy [1892 B}.

. GIGANTEUM Dendy.

Sycon giganteum Dendy [1892 B].

. HELLERI von Lendenfeld.

Sycandra helleri von Lendenteld (1891).

. HUMBOLDTIL Lisso. Type species of the genus.

Sycon humboldtii Risso [1826].

Dunstervillia corcyrensis O. Schmidt [1862], fide

Haeckel [1872].

Dunstervillia schmidti Haeckel [1870], fide Haeckel

[1872].

Sycandra humboldtii Haeckel [1872].

. IMPLETUM /aeckel.

Artynas villoswm Haeckel [1870], fide Haeckel [1872].

Sycandra villosa var. impletum Haeckel [1872].

. INCONSPICUUM von Lendenfeld.

Sycandra inconspicua von Lendenfeld [1885 B].

. INCRUSTANS brettfuss.

Sycon incrustans Breitfuss [1898 E}.

. KARAJAKENSE Breitfuss.

Sycon karajakense Breitfuss [1897 }.

. KERGUELENSIS Urban.

Sycon kerguelensis Urban [1908].

. LAMBEL, sp. 2.

Sycon asperum Lambe [1896].

The new specific name has been given to the above

species on account of the fact that the name asperwm is

already occupied in this genus. (See above.)

31. S. LANCEOLATUM Haeckel.

Sycum lanceolatwm Haeckel [1870].

Sycandra ciliata var. lanceolata Haeckel [1872].

AO.

4].

44,

45,

A6.

47.

48.

op)

CALCAREOUS SPONGES.

. LENDENFELDI Row.

Sycon lendenfeldi Row [1913 MS.}.

. LInGuA Haeckel.

Sycortis lingua Haeckel [1872].

. LONGSTAFFI Jenkin.

Hypodictyon longstagi Jenkin [1908 B].

. MAxImMuUM Haeckel.

Sycandra arctica var. maxima Haeckel [1872].

. MINUTUM Dendy.

Sycon minutum Dendy [1892 B].

. MUNDULUM Lambe.

Sycon mundulum Lambe [1900 B}.

. MUNITUM Jenkin.

Sycon munitum Jenkin [1908 A].

. ORNATUM Kirk.

Sycon ornatum Kirk [1897].

. ovatum Haeckel.

Sycum ovatum Haeckel [1870}.

Sycandra ciliata var. ovata Haeckel | 1872).

. PARVULUM Preiwisch.

Sycandra parvula Preiwisch [1904}.

. PEDICELLATUM AUrk.

‘ycon pedicellatum Kirk {1897}.

. PeTIoLATUM Haeckel [1870].

Sycum petiolatum Haeckel [1870].

Sycandra ampulla vay. petiolata Haeckel [1872].

. POLARE Haeckel.

Sycandra arctica var. polaris Haeckel {1872}.

. PROBOSCIDEUM Haeckel.

Syconella proboscidea Haeckel [1870].

Sycandra raphanus vay. proboscidea Haeckel [1872]

. PROCUMBENS Haeckel.

Sycum procumbens Haeckel [1870].

Sycandra raphanus var. procumbens Haeckel [1872].

. PRoTECTUM Lambe.

Sycon protectum Lambe [1896].

. QUADRANGULATUM O. Schmidt.

Syconella quadrangulata O. Schmidt {1868}.

Sycandra quadrangulata Haeckel [1872].

747

748

50.

51.

52.

53.

54.

55.

_ 56.

57.

58.

59.

60.

PROF. A. DENDY AND MR. R&R. W. H. ROW ON

. S. RAMosUM Haeckel.

Sycandra ramosa Haeckel [1872].

Leuckartea natalensis Michlucho-Maclay MS., fide Haeckel

[1872].

S. RAMSAYI von Lendenfeld.

Sycandra ramsayt von Lendenfeld [1885 A }.

S. RAPHANUS O. Schmidt.

Sycon raphanus O. Schmidt [1862].

Spongia inflata Delle Chiaje [1828], fide Haeckel [1872].

Sycarium vesica Haeckel [1870], fide Haeckel [1872].

Sycandra raphanus Haeckel [1872].

S. scumiptit Haeckel.

Sycandra schmidtii Haeckel [1872].

This species must be distinguished from Dunstervillia

schmidtii Haeckel, a synonym of Sycon humboldti Risso.

S. SCHUFFNERI, sp. 1.

Sycandra quadrata Schuffner [1877].

We propose this new name in order to avoid confusion

with Haeckel’s variety quadrata of Sycon quadrangulatum

(O. Schmidt).

S. serosum O. Schmidt.

Sycon setosum O. Schmidt [1862].

Sycandra setosa Haeckel [1872].

S. STAURIFERUM Preiwisch.

Sycandra staurifera Preiwisch [1904].

S. SUBHISPIDUM Carter.

Grantia subhispida Carter [1885-1886].

S. sycaAnpDRA von Lendenfeld.

Homoderma sycandra von Lendenfeld [1885 A].

Leucosolenia (2) sycandra Dendy [1891 A}.

See also Row [1913 MS.], under Sycon lendenfeldi.

S. TABULATUM Schuffner.

Sycandra tabulata Schuftner [1877].

Very probably identical with Haeckel’s variety tabulata of

Sycon elegans Bowerbank.

S. TENELLUM von Lendenfeld.

Sycantha tenella von Lendenfeld [1891].

Sycon tenellum Jenkin [1908 B].

S. TeRGESTINUM Haeckel.

Sycum tergestinum Haeckel [1870].

Sycandra raphanus var. tergestina Haeckel [1872].

CALCAREOUS SPONGES. . 749

61. S. rEssELLATUM Bowerbank.

Grantia tessellata Bowerbank [1864-1882 ].

Sycandra elegans var. tessellata Haeckel [1872].

62. S. resserARIUM Haeckel.

Sycandra quadrangulata var. tesseraria Haeckel [1872}-

63. S. ruBaA von Lendenfeld.

Sycandra tuba von Lendenfeld (1891).

64. S. ruBuLosum Haeckel.

Sycandra coronata var. tubulosa Haeckel [1872].

65. S. veruM Row.

Sycon verum Row [1913 MS.].

66. S. vittosum Haeckel.

Sycarium villoswm Haeckel [1870].

Sycum clavatwm Haeckel [1870], fide Haeckel [1872].

Sycandra villosa Haeckel {1872}.

67. S. vircuLtosum Haeckel. i

Sycandra aleyoncellum var. virgultosa Haeckel [1872].

Genus 17. Sycanpra Haeckel [1872] (emend.).

Diagnosis. The radially arranged flagellate chambers more or less

united where they come into contact with one another.

Gastral cavity traversed by strands of tissue containing

bundles of parallel oxea and forming a more or less strongly

developed endogastric network. Radially arranged dermal

oxea present.

For illustrations of this genus see Haeckel [1872].

We propose to use this generic name for O. Schmidt’s Ute

utriculus (=Sycandra utriculus Haeckel), which is sufficiently

sharply distinguished by its skeletogenous endogastric network.

There is only one other species in which this character is known

to occur, namely Leucettaga loculosa, a member of the family

Grantiide.

The speciés of Sycandra which precede S. utriculus in Haeckel’s

monograph having been relegated to earlier genera such as Sycon,

Ute and Grantia, this species becomes the type of the genus.

The only known species is :—

1. S. urricutus O. Schmidt.

Ute utriculus O. Schmidt [1870].

Sycandra utriculus Haeckel [1872].

750 PROF. A. DENDY AND MR. R. W. H. ROW ON

Family 7. HETEROPIIDA) Dendy [1892 B].

Diagnosis. With a distinct and continuous dermal cortex covering

over the chamber-layer and pierced by inhalant pores.

Subgastral sagittal and subdermal pseudosagittal radiates

are present. Flagellate chambers varying from elongated

and radially arranged to spherical and irregularly scattered.

With or without an articulate tubar skeleton. Nuclei of

collared cells probably always apical.

This family is identical in scope with the family as originally

proposed by Dendy [1892 B], and the difference now made in the

diagnosis is due to the fact that our conception of the subdermal

triradiates has changed. Up to the present we have considered

the characteristic subdermal spicules in this family as being truly

sagittal, with the basal ray centripetally directed. We have now

convinced ourselves, however, by a careful examination of a

number of species, that this is not the case, but that the

inwardly pointing ray is really one of the oral rays, and that

the original basal ray has taken on the appearance and position

of an oral ray. In other words, we find the clearest evidence

that these spicules are derived from ordinary distally situated

triradiates of the articulate tubar skeleton, which have undergone

rotation followed by the acquisition of a secondary pseudo-

symmetry. We therefore propose for them the name of

pseudosagittal. It will be remembered that Poléjaeff [1883]

recognised, in the case of Grantessa (Anphoriscus) poculum and

G. flamma, that the subdermal triradiates are not ordinary

sagittal spicules and that the centripetal ray is really one of the

lateral (=oral) rays and not the basal ray. He, however, con-

sidered that they are triradiates of the dermal cortex which

have undergone re-orientation, and not, as we maintain, tubar

triradiates.

Various species of the genus Grantessa show quite clearly how

the change has taken place. In Grantessa hirsuta we have a

primitive type with long chambers and an articulate skeleton of

many joints. At the distal ends of the chambers are tufts of

oxea, towards which the basal rays of the triradiates of the distal

joint of the tubar skeleton are inclined, as indeed occurs also in

the genera Sycon and Grantia. Moreover, the whole spicule has

become tilted until in some cases one of the original oral rays has

assumed a position at right angles to the surface, while the other

has come to lie nearly parallel to the surface, where it probably

serves to guard the entrance to the inhalant canal. In more

advanced cases, such as Grantessa sacca and G. hispida, the great

elongation of the now inwardly directed oral ray increases the

resemblance to an ordinary sagittal spicule, but a characteristic

asymmetry of the outwardly directed (apparent oral) rays,

accompanied by a definite kink or angulation in one of them,

. CALCAREOUS SPONGES. OD

(which appears to be due to change of position during individual

growth), affords a clear indication of what has really taken place.

Finally, in the most advanced types, such as Grantessa intusarti-

culata, we find the pseudosagittal subdermal spicules assuming

great dominance, almost to the exclusion of the typical articulate

tubar skeleton, so that we arrive at the so-called inarticulate

type. |

The development of these characteristic spicules appears to

antedate the appearance of a definite dermal cortex, for we

find in Sycon ensiferwm Dendy a similar canting of certain of

the distal tubar triradiates, which renders this species almost

indistinguishable from Grantessa. Indeed, it is this out-turning

of one of the rays of the distal tubar trivadiates that has, in our

opinion, led to the formation of a dermal cortex, probably by the

drawing out of the soft tissues of the sponge with the rays in

question. Thus the origin of the dermal cortex in this family

would be intimately connected with the development of these

subdermal pseudosagittal triradiates. In the Grantiide, on the

other hand, the dermal cortex appears to have originated in the

development of tangentially placed trivadiates in a previously

aspicular pore-bearing dermal membrane.

These views undoubtedly tend to bridge over the gap between

the Sycettide and the Heteropiide, and ‘radioed the more primitive

species of Grantessa are differentiated from Sycon and Grantia by

very slight characters, and difficult to separate from them, but.

the rotation of the trivadiates in question appears to have formed

the starting point of a new line of skeletal evolution which seems

to us to deserve recognition as marking a distinct family.

We consider the views here put ferward as to the origin of the

subdermal pseudosagittal spicples of the Heteropiide to be more

in accordance with observed facts than those previously suggested

by one of us (Row 1909) in regard to the ‘“‘subdermal secondary

sagittal triradiates ” of Grantilla, which seem to be pseudosagittal

spicules really similar to those of Grantessa.

We have changed the spelling of the name of the family from

Heteropide to Heteropiide, the latter being more in accordance

with the usual practice.

Genus 18. GranteEssA von Lendenfeld [1885 B] (emend.).

Diagnosis. Canal system syconoid. No colossal longitudinally

placed oxea.

For illustrations of this genus see von Lendenfeld [1885 B]

and Dendy [1893 A].

The tubar skeleton in this genus ranges from articulate, with

very numerous joints, as in Grantessa sacca, G. erinaceus, G. hirsuta

and G, hispida, to inarticulate or nearly so, as in G. glabra and

752 PROF. A. DENDY AND MR. R. W. H. ROW ON

G. polyperistomia. This character might imdeed be used as a

basis for the subdivision of the genus, were it not for the

impossibility of drawing a satisfactory line between the two types

of tubar skeleton,

We recognise the following species as belonging to this

genus :—

Secrion A. With large, usually radially arranged oxea

but without microxea.

1. G. compressa Carter.

Heteropia compressa Carter [1885-1886 }.

2. G. ERECTA Carter.

Heteropia erecta Carter [1885-1886].

3. G. ERINACEUS Carter.

Leuconia erinaceus Carter [1885-1886 |

4, G. rtamMa Poléjaeff.

Amphoriscus amma Poleéjaeft [1883].

5. G. GLABRA Low.

Grantessa glabra Row [1909].

6. G HASTIFERA Low.

Grantilla hastifera Row [1909].

Grantessa hastifera Dendy [1913].

7. G. nirsuta Carter.

Hypograntia hirsuta Carter [1885-1886].

Grantessa hirsuta Row [1913 MS.].

8. G. HIsPIDA Dendy.

Grantessa hispida Dendy {1892 B).

9, G. LANCEOLATA Breitfuss.

Lbnerella lanceolata Breitfuss [1898 B].

10. G. niTIDA Arnesen.

Ebnerella nitida Arnesen [1901).

11. G. paiaaica Ridley.

Nardoa pelagica Ridley [1881 }.

12. G. PLURIOSCULIFERA Carter.

Heteropia pluriosculifera Carter [1885-1886].

13. G. pocutum Polejaeff.

Amphoriscus poculum Poléjaeff [1883 ].

Heteropia patulosculifera Carter [1885-1886], fide Dendy

[1892 B].

Grantessa poculum Dendy [1892 B].

15.

16.

CALCAREOUS SPONGES. 753

. POLYPERISTOMIA Carter.

Heteropia polyperistonia Carter [1885-1886 ].

Grantessa polyperistonua Row [1913 MS. ].

. sacca von Lendenfeld. Type species of the genus.

Grantessa sacca von Lendenfeld [1885 B].

G. SYCILLOIDES Schuffner.

Sycortis sycilloides Schuftner [1877].

Section B. Without large oxea, but with microxea,

. G. INTUSARTICULATA Carter.

Hypograntia intusarticulata Carter [1885-1886 ].

Hypograntia medioarticulata Carter [1885-1886], fide

Dendy [1892 B].

Grantessa intusarticulata Dendy [1892 B].

Section C. With large, usually radially arranged oxea

and with microxea.

18. G. KiKentTHALI Breitfuss.

Hbnerella kikenthali Breitfuss [1896 A].

19. G, PREIWISCHI, sp. n.

Ebnerella compressa Preiwisch [1904].

This new specific name has been given to the species, as

compressa is already occupied. (See above.)

20, G. spissa Carter.

Heteropia spissa Carter [1885-1886].

21. G. rHompsont Lambe.

Amphoriscus thompsoni Lambe [1900 B].

Section D. Without any oxea.

22. G. ciactatis Haeckel.

Sycaltis glacialis Haeckel [1872].

23. G. MURMANENSIS Greitfuss. |

Amphoriscus murmanensis Breitfuss [1898 B].

24, G. stmpLex Jenkin.

Grantessa simplex Jenkin [1908 A}.

25. G, srauRiDEA Haeckel.

Sycetta stauridea Haeckel [1872].

Djeddea violacea Michlucho-Maclay MS., fide Haeckel

[1872].

26. G. ZANZIBARENSIS Jenkin.

Grantessa zanzibarensis Jenkin [1908 A].

754 PROF. A, DENDY AND MR. R. W. H. ROW ON

Genus 19. Hermropia Carter [1885-1886] (emend.).

Diagnosis. Canal system syconoid. Dermal cortex with colossal

longitudinal oxea.

For illustrations of this genus see Row [1913 MS.].

This genus stands in precisely the same relation to Grantessa

that Ute does to Grantia. It is noteworthy that, in all known

species of this genus, as in the more highly developed species of

Grantessa, the tubar skeleton has been reduced to the subgastral

sagittal triradiates, supplemented by the subdermal pseudosagittal

triradiates, and has thus become ‘‘ inarticulate.”

The genus Heteropia was diagnosed by Carter in July 1886 as

follows :—‘‘ Caleareous sponges in which the wall is simply

composed of sarcode supported on large sagittiform triradiates,

whose heads are fixed in opposite sides of it respectively, and

whose long shafts, extending perpendicularly across it, more or

less overlap each other.”

Most of the species placed by Carter in this genus belong

to the earlier genus Grantessa of von Lendenfeld. There is,

however, one of his species, Heteropia ramosa, which is dis-

tinguished by the presence of colossal longitudinal dermal oxea,

and which may be regarded as the type of Carter’s genus. It is

curious that Mr. Carter himself [1886] described it under the

name of Aphroceras ramosa, whilst saying at the same time that

it belonged to his genus /eteropia.

We recognise the following species as belonging to this

genus :— :

Section A. Without microxea.

1. H. GLomErosa Bowerbank.

Leuconia glomerosa Bowerbank [1872-1876].

2. H. rAmosa Carter. Type species of the genus.

Aphroceras ramosa Carter [1886].

3. H. stmpLex Low.

Heteropia simplex Row [1913 MS. ].

Section B. With microxea.

4. H. roperert Lambe.

Heteropia rodgerit Lambe [1900].

Genus 20. Ampuiutr Hanitsch [1894].

Diagnosis. Canal system syconoid. Both gastral and dermal

cortices with colossal longitudinal oxea.

For illustrations of this genus see Hanitsch [1895].

CALCAREOUS SPONGES. 755

This genus may be regarded as derived from some more

primitive type of Grantessa by the addition of colossal longitudinal

oxea to both dermal and gastral cortices. Those in the gastral

cortex are probably to be regarded as having been derived from

the oxea of the oscular fringe, by downward extension. In the

only known species microxea are present, and the articulate tubar

skeleton still persists.

The only known species is :—

1. A. pautint Hanitsch.

Amphiute paulint Hanitsch [1894].

Genus 21. VosmArropsis Dendy [1892 B}.

Diagnosis. Canal system sylleibid (or leuconoid ?). Skeleton of

the chamber layer composed of the centrifugally directed

rays of subgastral sagittal triradiates and the centripetally

directed rays of subdermal pseudosagittal triradiates, which

may be supplemented or partially replaced by confused

triradiates. No colossal longitudinal oxea.

For illustrations of this genus see Dendy [1892 B].

In all the known species of Vosnaeropsis we find that the

canal system has not developed beyond the sylleibid condition,

and there can still be distinguished in the chamber layer very

clear indications of an inarticulate tubar skeleton ; in fact, the

genus seems to have reached almost exactly the same level of

evolution as JJegapogon in the Grantiide, when allowance is

made for the different type of skeleton in the two families.

This would at any rate seem to suggest that the family Heteropiide

is of comparatively recent origin, and that more complex forms,

comparable to the higher types of the Grantiide, have not yet

made their appearance, unless, indeed, they have merely escaped

observation.

We recognise the following species in this genus :—

Section A. With large oxea and microxea.

1. V. pepressa Dendy.

Vosmaeropsis depressa Dendy [1892 B}.

2. V. MAcERA Carter. Type species of the genus.

Heteropia macera Carter [1885-1886 }.

Vosmaeropsis macera Dendy [1892 B}.

3. V. witsont Dendy.

Vosmaeropsis wilsont Dendy [1892 B].

Proc. Zoou. Soc.—1913, No. L. 50

756 PROF. A. DENDY AND MR. R. W. H. ROW ON

Srecrion B. With large, usually radially arranged oxea,

but without microxea.

Although the author’s description does not conform to the

above diagnosis, we include Poléjaefi’s Leucilla connexiva in this

section of the genus, for the figures given by him show oxea

present, though no reference is made to them in the text, and

our own examination of the type specimen revealed the presence

of occasional trichoxea.

4. V. connEXIvA Poléjaeff.

Leucilla connexiva Poléjaetf [1883].

5. V. cyaruus Verrill.

Leucandra cyathus Verrill [1873].

6. V. DENDYI How.

Vosmaeropsis dendyi Row [1913 MS8.].

7. V. primitiva Row.

Vosmaeropsis primitiva Row [1913 MS.].

8. V. spericatum Lidley.

Aphroceras sericatum Ridley [1884].

This species has been placed in Vosmaeropsis as a result of an

examination of the type specimen made by us at the Natural

History Department of the British Museum, which revealed the

existence of typical subdermal pseudosagittal triradiates, though

the author’s original description does not mention them.

Genus 22. GranrittA Row [1909] (emend.).

Diagnosis. Canal system syconoid, 'Tubar skeleton ( ? always) in-

articulate, composed of subdermal pseudosagittal triradiates

and subgastral sagittal triradiates, supplemented by

subdermal quadriradiates. No colossal longitudinal oxea.

For illustrations of this genus see Row [1909].

This gens was originally proposed by Row for two species,

G@. quadriradiata and G. hastifera, which were supposed to possess

certain features that necessitated the provision of a new family,

Grantillide. We now consider, however, that the characters in

question do not represent any fundamental peculiarities of

structure, and we have therefore abandoned the family, as already

stated in the Introduction.

One of the two species originally assigned to Grantilla, G.

qguadriradiata, however, presents an association of subdermal

quadriradiates with subdermal pseudosagittal triradiates, which is

not known in any other species of calcareous sponge, and we

therefore retain the name Grantilla for this species with an

emended diagnosis. The development of subdermal quadri-

radiates has evidently taken place repeatedly in the phylogeny of

CALCAREOUS SPONGES. 757

the Calearea. We have seen it already, for example, in Lewcetta,

Leucaltis and Leucettusa, and have not in those cases considered

the presence of such spicules as of generic value. In Grantilla,

however, they seem to assume more importance, and to take a

larger share in the formation of the skeleton of the chamber layer.

Nevertheless, had the genus not been already in existence, we

should have hesitated to propose it on this character alone.

The only known species of the genus is :-—

1. G. QUADRIRADIATA Row.

Grantilla quadriradiata Row [1909].

Family 8. GRANTIID Ai Dendy [1892] (emend.).

Diagnosis. With a distinct dermal cortex and a proper cortical

skeleton of tangential radiates, sometimes supplemented by,

and occasionally replaced by, oxea. Flagellate chambers

ranging from elongated and radially arranged to small,

spherical and irregularly scattered. Skeleton of the chamber

layer ranging from regularly articulate to irregularly

scattered. Typically with subgastral sagittal radiates. No

subdermal pseudosagittal triradiates. Subdermal quadri-

radiates, if present, always associated with a chamber-layer

skeleton containing confused triradiates. Nuclei of collared

cells probably always apical.

It must frankly be admitted that the boundary line between

the Sycettide and the Grantiide is by no means sharply defined.

The great distinguishing feature is the presence in the latter of a

distinct dermal cortex with its own proper skeleton. The develop-

ment of such a cortex appears to have formed the determining

condition for the further evolution of both the canal system and

the skeleton, and it must therefore be regarded as of great system-

atic importance.

The first commencement of such a cortex is, however, so slight

as to be almost indistinguishable from the mere pore-bearing

dermal membrane of the most highly specialised Sycons. In

Grantia compressa the cortex is so feebly developed that Dendy, in

his early work [1892 B], included this species in the genus Sycon,

laying more stress upon the presence of dermal tufts of oxea than

we are now inclined to do in this connection. It appears to us

that the line between Sycon and Grantia, and therefore between

the Sycettidee and Grantiide, must be drawn at the appearance

of a dermal cortical skeleton of tangential radiates distinct from

the skeleton of the radial chambers, and in accordance with these

views Grantia compressa is excluded from the genus Sycon.

Moreover, it must be pointed out that G. compressa is not the

only member of this family in which dermal tufts of oxea occur,

‘as they are present also in Sycute dendyi Kirk.

; 50*

758 PROF, A. DENDY AND MR. R. W. H. ROW ON

With the transition from the syconoid to the leuconoid type o

canal system in this family, and the correlated replacement of th

articulate tubar skeleton by irregularly scattered radiates, we ge,

a close approach to the more advanced Leucascide, such at

Leucetta and Pericharax, and we have here one of those cases o

convergence which are so frequently met with amongst sponges»

but we have already laid sufticient emphasis upon this point.

Even in the genus Leucandra, however, subgastral sagittal tri-

radiates are usually present, and when they are absent their

absence must be regarded as secondary.

So far as our experience goes the nucleus of the collared cells

is always apical in position in this family. We have been able to

determine it in 17 species, as enumerated in an earlier section of

this paper.

The family is a very large one, comprising no less than 25 out

of the 51 genera of recent calcareous sponges which we recognise,

and containing a great diversity of structural types within it.

There are, however, very great difficulties in the way of dividing

it into subfamilies, the chief of these being the fact that the

possible methods of deriving the various genera from one

another within the family are manifold, and it is impossible

to determine satisfactorily which are the true lines upon which

evolution has proceeded. We might, for example, place all

those genera which have a syconoid canal system and colossal

longitudinal oxea in the dermal cortex together in a subfamily

Uteine; or we might separate the genus Uteopsis from the

others, and unite it with Achramorpha and Anamiwilla in a

subfamily characterised by the reduction of the tubar skeleton to

a single joint. But neither of these two possible subfamilies

would seem to be very sharply defined, and moreover, the cha-

racters in question are not confined to members of the Grantiidee.

In short, we feel that in the present state of our knowledge it is

impossible to decide which method of grouping would express

most correctly the real affinities of the genera concerned. ‘This

is the case with almost all the possible methods of grouping the

genera, and we have therefore decided not to attempt to split up

the family, but merely to indicate the approximate relationships

of the genera, so far as this is possible in a linear series, by the

order in each we have arranged them,

Although it seems probable that the majority of the genera in

this family are descended from the genus Sycon, yet it is quite

possible that some of them may be descended independently from

Sycetta, and therefore that the family may be of diphyletic

origin.

We have changed the name of the family from Grantide to

Grantiide in accordance with the usual practice of systematic

zoologists.

CALCAREOUS SPONGES. 759

Genus 23. Grant1A Fleming [1828] (emend.).

Diagnosis. Canal system syconoid. Colossal longitudinal oxea,

if present, projecting from the surface. Tubar skeleton

articulate, composed of radiate spicules, which may or may

not be supplemented by oxea.

For illustrations of this genus see Dendy [1893 A ].

It has beeri conclusively shown by Minchin [1896] that the

type species of the genus Grantia is G'. compressa, and that the

name Grantia must always be given to the group of species

associated with G. compressa. We have already pointed out that

this species has a definite, though slight, dermal cortex, and

that its true position is in the present family, and in fact, in the

genus Grantia as defined by Dendy in 1892.

We may point out that G. intermedia Thacker stands alone in

the genus, as far as is at present known, in the presence of apical

rays on the tangential cortical radiates, and although we do not

attach much importance to such spicules, we feel that their

presence in this species indicates at any rate a possible starting

point for the family Amphoriscidee.

We recognise the following species as belonging to this

3 § Sp ging

genus :—

Section A. With large, usually radially arranged oxea,

but without microxea.

1. G. artantica Ridley.

Grantia atlantica Ridley [1881].

2. G. BrevipiLis Haeckel.

Sycandra cupillosa var. brevipilis Haeckel [1872].

3. G. CANADENSIS Lambe.

Grantia canadensis Lambe | 1896].

4, G. caprntosa O. Schmidt.

Ute capillosa O. Schmidt [1862].

Sycandra capillosa Haeckel [1872].

5, G, CHARLACEA Jenkin.

Dermatreton chartaceum Jenkin [1908 B].

6. G. CLAVIGERA O. Schmidt.

Sycinula clavigera O. Schmidt [1870].

Sycandra clavigera (Sycandra compressa var. clavigera)

Haeckel [1872].

7. G. comoxensis Lambe.

Grantia comoxensis Lambe [1893].

760

10.

11.

13.

14.

18.

19,

20.

2l.

PROF. A. DENDY AND MR. R. W. H. ROW ON

compressa Fabricius. Type species of the genus.

Spongia compressa Fabricius [1780].

Sycum lingua Haeckel [1870], fide Haeckel [1872].

Sycarium rhopalodes Haeckel [1870], fide Haeckel [1872].

Sycandra compressa Haeckel {1872}.

Sycon compressum Dendy [1892 B].

. FOLIACEA Montagu.

Spongia foliacea Montagu [1812].

Sycandra foliacea (Sycandra compressa var. foliacea)

Haeckel [1872].

. GENUINA Row.

Grantia genuina Row [1913 MS.]}.

. GRACILIS von Lendenfeld.

Vosmaeria gracilis von Lendenfeld [1885 B].

. HODGSONT Jenkin.

Dermatreton hodgsoni Jenkin [1908 B}.

.INtTERMEDIA Zhacker.

Grantia intermedia Thacker [1908].

. LOBATA Haeckel.

Sycandra lobata (Sycandra compressa var. lobata) Haeckel

[1872].

. toners Haeckel.

Sycandra capillosa vax. longipilis Haeckel (1872).

. MonsTRUOSA Dreitfuss.

Grantia monstruosa Breitfuss [1898 B].

PENNIGERA Haeckel.

Sycandra pennigera (Sycandra compressa var. pennigera)

Haeckel [1872].

scorrt Jenkin.

Tenthrenodes scotti Jenkin [1908 B].

. tenuis Urban.

Grantia tenuis Urban [1908].

. VOSMAERL Dendy.

Grantia vosmaeri Dendy [1892 B].

Section B. Without any oxea.

AscoNoIbDES bretifuss.

Sycetta asconoides Breitfuss [1896 B].

Our reasons for placing this species under Grantia rather

than under Sycetta have been stated when discussing the

latter genus.

CALCAREOUS SPONGES. 761

22. G. cuputa Haeckel.

Sycetta cupula Haeckel [1872].

23. G. iInvENuSTA Lambe.

Grantia invenusta Lambe [1900 B].

24, G. strositus Haeckel.

Sycetta strobilus Haeckel [1872].

Section C. With large, usually radially arranged oxea,

and with microxea.

25. G. acuLEATA Urban.

Grantia aculeata Urban [1908].

26. G. EXTUSARTICULATA Carter.

Hypograntia extusarticulata Carter [1885-1886].

Grantia extusarticulata Dendy [1892 B].

27. G. inpica Dendy.

Grantia indica Dendy [1912].

28. G. MIRABILIS [ristedt.

Ascandra mirabilis Fristedt [1887].

Grantia mirabilis Lundbeck [1909].

29. G. tuBEROSA Poléjaeff.

Grantia tuberosa Poléjaeff [1883].

Section D. With microxea, but without large oxea.

30. G. pavieata Haeckel.

Sycortis levigata Haeckel [1872].

Sycortusa levigata von Lendenfeld [1885 B].

31. G. painiipsir Lambe.

Grantia phillipsii Lambe [1900 B].

The following are doubtfully assigned to this genus :—

32. G. sINGULARIS Breitfuss.

Sphenophorina singularis Breitfuss [1898 B].

The genus Sphenophorina is discussed at some length in the

list of rejected genera.

33. G. uRcEOLUS Miller.

Spongia urceolus Miller [1788-1796].

Stated by Johnston [1842] to be very probably a variety of

Grantia compressa.

Genus 24. T'kICHONOPSIS nov.

Diagnosis. Sponge consisting of a single stipitate person with

enormously expanded gastral cavity and thin, much folded

162 PROF. A. DENDY AND MR. R. W. H. ROW ON

wall, whose convoluted edge represents the oscular margin.

Canal system syconoid. Tubar skeleton articulate. Without

colossal longitudinal oxea.

For illustrations of this genus see Dendy [1891 B].

We propose this genus for the reception of the remarkable

species 7’. labyrinthica, usually known as Grantia labyrinthica,

which forms the subject of a special memoir by one of us, Dendy

[1891 B]. We now consider that the very peculiar external form

is of sufficient importance to justify generic separation. The

species was originally placed by Carter in his genus Zeichonella,

on account of some superficial resemblance to his 7. prolifera;

but although the name TZeichonella has now been universally

abandoned even for Leucetta (Teichonella) prolifera, we do not

consider ourselves justified in reviving it for Grantia labyrinthica,

for Mr. Carter himself subsequently dissociated this species from

Teichonella and placed it in the genus Grantia [1885-1886].

The only known species is :—

1, T. LABYRINTHICA Carter.

Teichonella labyrinthica Carter | 1878].

Grantia labyrinthica Dendy [1891 B].

Genus 25. Grantropsis Dendy [1892 B].

Diagnosis. Canal system syconoid. Dermal cortex as thick as

the chamber layer, with many layers of tangential triradiates.

Tubar skeleton articulate, the proximal joint being composed

of subgastral sagittal quadriradiates (? or triradiates), the

other joints of sagittal triradiates practically reduced to

the basal ray by suppression of the paired rays. Without

colossal longitudinal oxea.

For illustrations of this genus see Dendy [1893 A] and Row

[1913 MS.]}.

This genus was first proposed by Dendy [1892 B] for his

Grantiopsis cylindrica, and was considered by him to be a sub-

genus of Grantia. Jenkin [1908 Bj placed it as a distinct genus

in his family Staurorrhaphide, on the ground that the subgastral

spicules were “‘chiactines.” As we cannot accept the chiact

theory, we again transfer the genus to the Grantiide, but

consider it sufficiently distinct from Grantia to deserve generic

recognition.

We have recently discovered, as the result of our study of

Mr. Carter’s MS. illustrations, in the possession of one of us,

that that author’s “ Hypograntia infrequens (incertze sedis)” is

undoubtedly a species of Girantiopsis, and the same species has

recently turned up again in the collection made by the Hamburg

CALCAREOUS SPONGES. 763

South-Western Australian Expedition of 1905. As neither

Mr. Carter’s Hypograntia nor the species H. infrequens were

ever recognisably diagnosed, we do not consider it necessary to

abandon the generic name Grantiopsis. The question will be

more fully dealt with in the forthcoming report on the above-

mentioned collection [Row 1913 MS.].

We recognise the following species in this genus :-—

1. G. cyninprica Dendy. Type species of the genus.

Grantiopsis cylindrica Dendy [1892 B].

2. G. INFREQUENS Carter.

Hypograntia infrequens Carter [1885-1886 }.

Grantiopsis infrequens Row [1913 MS.].

Genus 26. SycurTE nov.

Diagnosis. Canal system syconoid. Dermal cortex provided with

colossal longitudinally arranged oxea. ‘Tubar skeleton

articulate. Distal ends of the flagellate chambers crowned

with tufts of oxea lying between the colossal longitudinal

oxea.

For illustrations of this genus see Kirk [1894].

This genus has been provided for Kirk’s Sycon dendyi, a species

which is curiously intermediate in character between Sycon and

Ute, retaining the well-defined tufts of oxea which are charac-

teristic of Sycon and at the same time possessing the colossal

longitudinal oxea characteristic of Ute.

The only known species is :—

1 S. penpy1 Kirk.

Sycon dendyi Kirk [1894].

Genus 27. Urs O. Schmidt [1862] (emend.).

Diagnosis. Canal system syconoid. Tubar skeleton articulate.

Dermal cortex well developed, containing colossal longi-

tudinal oxea. No tufts of oxea at the distal ends of the

flagellate chambers.

For illustrations of this genus see Dendy [1895 A].

We allocate the following species to this genus :—

Srcrion A. Without microxea.

1. U. ensata Bowerbank.

Grantia ensata Bowerbank [1864-1882].

Sycandra glabra var. ensata Haeckel [1872].

764 PROF. A. DENDY AND MR. R. W. H. ROW ON

2. U. atapra O. Schmidt. Type species of the genus.

Ute glabra O. Schmidt [1864].

Ute capillosa J. K. Gray [1867], fide Haeckel [1872].

Sycandra glabra Haeckel [1872].

3. U. rniaipa Haeckel.

Sycandra glabra var. rigida Haeckel [1872].

4, U. SYCONOIDES Carter.

Aphroceras syconoides Carter [1885-1886].

Section B. With microxea.

dD. U. spENCERI Dendy.

Ute spencert Dendy [1892 B].

6. U. sprcunosa Dendy.

Ute spiculosa Dendy [1892 B].

Genus 28. Synure Dendy [1892 A].

Diagnosis. Sponge consisting of many Ute-like individuals com-

pletely fused together, and invested with a common cortex

containing colossal longitudinal oxea.

For illustrations of this genus see Dendy [1893 A].

This genus represents the highest known type of integration

met with amongst syconoid sponges.

The only known species is :—

1. 8. puncHELLA Dendy.

Synute pulchella Dendy [1892 A].

Synute pulchella Row [1913 MS.].

Genus 29. Sycoporus Haeckel [1872] (emend.).

Utella Dendy [1892 B}.

Diagnosis. Canal system syconoid. ‘Tubar skeleton articulate.

Gastral cortex with a layer of large longitudinally arranged

oxea, but no oxea in the dermal cortex.

For illustrations of this genus see Haeckel [1872].

Dendy proposed the genus Utella in 1892 for the reception of

Haeckel’s Sycandra hystrix, and suggested that O. Schmidt’s

Ute utriculus might also be included in it. As we feel that the

laws of priority necessitate our using Haeckel’s subgeneric names,

where possible, in preference to later ones, we propose to substitute

Sycodorus for Utella, the species which precede S. hystria in the

subgenus in Haeckel’s monograph having been assigned to earlier

genera. For Ute utriculus we have retained the generic name

Sycandra.

CALCAREOUS SPONGES, 765

The only known species is :—

1. 8. nystrix Haeckel.

Sycandra hystrix Haeckel [1872].

Genus 30. AcHrAMorPHA Jenkin [1908 B]| (emend.).

Diagnosis. Canal system syconoid. Skeleton of the chamber

layer reduced to the basal rays of the subgastral sagittal

triradiates (which may become quadriradiates by the addi-

tion of an apical ray), with radial oxea lying between the

chambers and projecting from the surface. No colossal

longitudinal oxea.

For illustrations of this genus see Jenkin [1908 B]; and Breitfuss

[1898 D] under Hbnerella schulzer.

This genus was proposed by Jenkin for the three species

glacialis, grandinis and nivalis, which resemble one another

closely, and which all possess the so-called chiactines of his

supposed family Staurorrhaphide. The fact that another species,

Breitfuss’s Hbnerella schulzei, differs in no essential point except

the absence of chiactines, affords strong evidence for our view

that the latter are nothing but subgastral sagittal triradiates

that have developed apical rays, and therefore not even of generic

importance, since such spicules are known to occur in other

Grantiide. These species, however, form a well-defined group,

and we accordingly retain the generic name Achramorpha, with

an emended diagnosis based upon what we believe to be more

important characters.

We assign the following species to the genus :—

Section A. With microxea.

1. A. enacianis Jenkin.

Achramorpha glacialis Jenkin [1908 B}.

2. A. GRANDINIS Jenkin.

Achramorpha grandinis Jenkin [1908 B}.

3. A. NIVALIS Jenkin. Type species of the genus.

Achramorpha nivalis Jenkin [1908 B].

4. A. SCHULZEL Greitfuss.

Hbnerella schulzei Breitfuss [1896 A].

Secrion B. Without microxea.

5. A. rRuNcATA Topsent.

Grantia truncata Vopsent [1907].

766 PROF. A. DENDY AND MR. R. W. H. ROW ON

Genus 31. UTropsis nov.

Diagnosis. Canal system syconoid. Tubar skeleton reduced to

the basal rays of subgastral sagittal radiates, supplemented

distally by radially arranged oxea. Dermal cortex well

developed, and containing colossal longitudinal oxea,

For illustrations of this genus see Poléjaeff [1883].

We propose this genus for Poléjaeff’s Ute argentea, which

obviously differs widely from the other species of the genus Ute.

The replacement of the distal portion of the tubar skeleton

by oxea is a very unusual feature, and, from the analogy of

Grantiopsis, we think it possible, but not probable, that these

oxea are really radiates whose paired rays have been completely

lost. It seems more probable that they are to be compared to

the radial oxea of Achramorpha.

The ‘‘tubar” quadriradiates referred to by Poléjaeff presumably

belong to the exhalant canals of the chambers, and not to the

chambers themselves.

The only known species of the genus is :—

1. U. arcentea Poléjaeff.

Ute argentea Poléjaeff [1883].

Genus 32. ANAMIXILLA Poléjaeff [1883].

Diagnosis. Canal system syconoid. Tubar skeleton reduced to

the outwardly directed basal rays of the subgastral sagittal

radiates. Skeleton of the chamber layer otherwise con-

sisting of large triradiate spicules, arranged without regard

to the direction of the chambers. Dermal cortex well

developed, but without colossal longitudinal oxea.

For illustrations of this genus see Poléjaeff [1883].

As Dendy has previously pointed out [1893 A], this genus may

be looked upon as a Grantia in which the ordinary tubar skeleton

has been almost entirely replaced by the invasion of large tri-

radiates from the dermal cortex. Thus the genus is of interest

as indicating one method by which the confused chamber-layer

skeleton of Lewcandra may have arisen.

The only known species is :—

1. A. rorrest Poléjaeff

Anaminilla torresi Poléjaeff [1883].

CALCAREOUS SPONGES. 767

Genus 33. Sycyssa Haeckel [1872].

Diagnosis. Canal system syconoid. Skeleton entirely composed

of oxea. Dermal cortex well developed, but without colossal

longitudinal oxea. Gastral cortex with a subgastral layer of

oxea, arranged longitudinally.

For illustrations of this genus see Haeckel [1872].

This genus is highly remarkable for the complete suppression

of the radiate spicules. An analogous condition is met with in

Ascyssa, Leucyssa, Trichogypsia and Kuarrhaphis.

The only known species is :—

1. S. suxtEey1 Haeckel.

Sycyssa huxleyi Haeckel [1872].

Genus 34. Mrcaprocon Jenkin [1908 B] (emend.).

Magnosis. Canal system sylleibid or leuconoid. Skeleton of the

chamber layer retaining clear traces of the original articulate

character and not confused ; composed chiefly of subgastral

sagittal quadriradiates, with their apical rays projecting into

the gastral cavity; with a few sagittal trivadiates arranged

as usual. No gastral skeleton of tangentially placed radiates,

except round the osculum. No colossal longitudinal oxea.

For illustrations of this genus see Jenkin [1908 B].

Jenkin included in this genus five species, W/. cruciferus,

M. villosus, M. raripilus, M. pollicaris and M. crispatus, and

placed it in the family Staurorrhaphide on account of the

presence of so-called chiactines. His figure of J/. villosus,

however, 1s alone sufficient to indicate that the “ chiactines”

are merely subgastral sagittal radiates which have developed

apical rays, as In so many other cases, and we find it necessary

to base the genus, which we believe to be a natural one, on other

characters.

The absence of tangentially arranged gastral radiates, combined

with the presence of the so-called ‘“chiactines,” might be used as

an argument for the validity of the chiact theory, on the sup-

position that all the gastral tangential radiates had been converted

into chiactines, but we must remember that in one species at any

rate, J. raripilus, the so-called chiactines are associated with

subgastral sagittal triradiates, which differ from them only in

the absence of an apical ray, and there is no ground for supposing

that the chiactines have any special significance. It is quite

possible that all these subgastral sagittal spicules have been

rotated into their present positions, as already pointed out in the

Introduction, but this fact does not justify us in distinguishing

768 PROF. A, DENDY AND MR. R. W. H. ROW ON

the chiactines as fundamentally different from other subgastral

sagittal radiates.

Megapogon villosus, with its sylleibid canal system and almost

syeonoid skeleton, exhibits a very interesting stage in the

evolution of the leuconoid type, and the same is perhaps true

of MW. pollicaris.

We place the following species in this genus :—

1. M. crispatus Jenkin.

Megapogon crispatus Jenkin [1908 B].

2. M. crucirerus Poldjaeff. Type species of the genus.

Leuconia crucifera Poléjaeff [1883].

3. M. PoniicaRris Jenkin.

Megapogon pollicaris Jenkin [1908 B].

4, M. rAripiLus Jenkin.

Megapogon raripilus Jenkin [1908 B}.

5. M. vinLosus Jenkin.

Megapogon villosus Jenkin [1908 B}.

Genus 35. Leucanpra Haeckel [1872] (emend.).

Diagnosis. Sponge usually a single person, or a colony of such

persons in which the component individuals are readily

recognisable. Canal system leuconoid. Skeleton of the

chamber layer more or less confused, but frequently with

vestiges of an articulate tubar skeleton in the form of sub-

gastral or other sagittal trivadiates. Dermal skeleton of

tangentially placed triradiates, which may sometimes develop

an apical ray. Colossal longitudinally placed oxea, when

occurring in the dermal cortex, never forming a smooth

layer, but always projecting conspicuously from the surface.

For illustrations of this genus see Vosmaer [1880] and Dendy

[1893 A].

The genus Leucandra as here defined is much more narrowly

circumscribed than it was by Dendy previously [1892 B]. In

fact Dendy’s genus is here represented by no less than 10 genera,

namely, Lewcandra, Baeria, Leucopsila, Aphroceras, Leucettaga,

Lamontia and Hilhardia in the family Grantiide, and Leuco-

malthe, Pericharax and Leucettusa in other families, while

certain species have been transferred to Leucetta. On the other

hand, we include in the present genus certain species which

possess subdermal quadriradiates, and which on that account

were placed by Dendy in the genus Leweilla; for, as we had

occasion to point out with regard to both Leucetta and Grantia,

CALCAREOUS SPONGES. 769

we do not consider that the mere presence or absence of a fourth

ray on a radiate spicule in the dermal cortex can be regarded as

of generic import. We shall discuss the true characteristics of

Leucilla when dealing with that genus.

The genus Leucandra as now defined may be regarded as

derived from a Grantia-like ancestor by the conversion of

the syconoid canal system into a leuconoid one, with the

simultaneous replacement of the articulate tubar skeleton by

an irregularly scattered skeleton of the chamber layer (compare

Anamixilla). Indications of the syconoid ancestry can, however,

frequently be detected in the skeleton (compare d/egapogon),

while as regards canal system such species as L. australiensis

Carter and ZL. infesta sp. n. (Leucilla intermedia Row [1909 )),

which are of the so-called sylleibid type, form connecting links

between Grantia and Leucandra.

We recognise the following species as belonging to this

genus :—

Section A. With large, usually radially arranged oxea,

but without microxea.

1. L, ananas Montagu.

Spongia ananas Montagu [1812].

Spongia pulverulenta Grant [1826], fide Haeckel [1872].

Scypha ovata 8. F. Gray [1821], fide Haeckel [1872].

Sycinula penicillata O. Schmidt [1870], fide Haeckel

[1872].

Leucandra ananas Haeckel [1872].

. ANGUINEA Lidley.

Leucortis anguinea Ridley [1884].

3. L. anomMALA Haeckel.

Leucetta pandora var. anomala Haeckel {1872}.

4, L. anmata Urban.

Leuconia armata Urban [1908].

5. L. aspera O. Schmidt.

Sycon asperum O. Schmidt [1862].

? Spongia panicea Esper [¢], fide Haeckel [ 1872}.

¢Spongia inflata Delle Chiaje [1828], fide Haeckel

[1872].

Leucandra aspera Haeckel [1872].

6. L. AUSTRALIENSIS Carter.

Leuconia fistulosa var. australiensis Carter | 1885-1886].

Leucandra austrahiensis Dendy [1892 B].

7. L. caminus Haeckel.

Dyssyconella caminus Haeckel [1870].

Leucandra caminus Haeckel (1872).

770 PROF. A. DENDY AND MR. R. W. He ROW ON

8. L. capiniavra Poléjaeff.

Leuconia multiformis var. capillata Poléjaeft [1883 ].

. CIRRATA Jenkin.

Leucandra cirrata Jenkin [1908 B].

10. L. crrrsosa Urban.

Leuconia cirrhosa Urban [1908].

11. L. cuavirormis Schuffner.

Leucandra claviformis Schuftner [1877].

12. L. compacta Carter.

Leuconia compacta Carter [1885-1886].

13. L. cramBessa Haeckel.

Leucandra crambessa Haeckel [1872].

14. L. crusracea Haeckel.

Leucaltis crustacea Haeckel [1872].

15. L. cuUMBERLANDENSIS Lambe.

Leucandra cumberlandensis Lambe | 1900 B).

16. L. ponnant Dendy.

Leucandra donnant Dendy [1905].

17. L. ecurnara Schuffner.

Leucandra echinata Schuftner [1877].

Leuconia echinata Carter [1885-1886], fide Dendy

[1913].

Leucandra echinata Dendy [1913].

18. L. ncepm O. Schmidt. Type species of the genus.

Sycinula egedit O. Schmidt [1870].

Leucandra egedii Haeckel | 1872].

19. L. rancienra Schugfner.

Leucandra falcigera Schuftner [1877].

20, L. risruLosa Johnston.

Grantia fistwlosa Johnston [1842].

Leucandra fistulosa Haeckel [1872].

21. L. cemmrpara Thacker.

Leucandra gemmipara Thacker [1908].

22. L. gosser Bowerbank.

Leucogypsia gosset Bowerbank [1864-1882].

Leucandra gosset Haeckel [1872].

23. L. HIBERNA Jenkin.

Leucandra hiberna Jenkin [1908 B}.

24. L. uirsuta Topsent.

Leucandra hirsuta Topsent [1907].

CALCAREOUS SPONGES. Con

25. L. HisPipA Carter.

Leuconia hispida Carter [1885-1886].

26. Li. INFESTA, sp. 7.

Leucilla intermedia Row [1909].

The new name is given to this species as intermedia is already

occupied,

27. LL. KERGUELENSIS Urban.

Leucandra kerguelensis Urban [1908].

28. L. LENDENFELDI Greitfuss.

Leuconia lendenfeldi Breitfuss [1897].

Leucortis elegans von Lendenfeld, MS., fide Breitfuss

[1897].

29. L. tunuuatTa Haeckel.

Leucandra lunulata Haeckel [1872].

30. L. MASATIERRE Breitfuss.

Leuconia masatierre Breitfuss [1898 E].

31. L. MEANDRINA von Lendenfeld.

Leucandra meandrina von Lendenfeld [1885 B].

32. L. minima Low.

Leucandra minima Row [1913 MS. }.

33. L. mutrirormis Poléjaeff.

Leuconia multiformis Poléjaeff | 1883}.

34, L. pHItiirensis Dendy.

Leucandra phillipensis Dendy [1892 B}.

35. L. pauyipa Row.

Leucandra pallida Row [1913 MS. ].

36. L. punyinar Haeckel.

Sycolepis pulvinar Haeckel {1870}.

Mlea dohrnii Michlucho-Maclay, MS8., jide Haeckel

[1872].

Leucortis pulvinar Haeckel | 1872).

37. L. rHULAKOMORPHA Low.

Leucandra thulakomorpha Row [1913 MS.].

38. L. vactnata von Lendenfeld.

Leucandra vaginata von Lendenfeld [1885 B].

39. L. vauipa Lambe.

Leucandra valida Lambe [1900 B].

40, L. vituosa von Lendenfeld.

Leucandra villosa von Lendenfeld {1885 B].

Proc. Zoou. Soc.—1913, No. UL. 5]

772 PROF. A. DENDY AND MR. R. W. H. ROW ON

41. L. wastnensis Jenkin.

Leucilla wasinensis Jenkin [1908 A].

Leucandra wasinensis Dendy [1913].

Secrion B. With large, usually radially arranged oxea,

and with microxea.

42, L. amorpHa Poléjueff.

Leuconia multiformis var. amorpha Polejaeff [1883].

43. L. anrracta Urban.

Leuconia anfracta Urban [1908].

44, LL, aprcatis Urban.

Leucandra apicalis Urban [1905].

45. L. BaLEARICA Lackschewitsch.

Leuconia balearica Lackschewitsch [1886 }.

46. L. BuLtBosa Hanttsch.

Leucandra bulbosa Hanitsch [1895].

47. L. coImBr& Greitfuss.

Leuconia coimbre Breitfuss [1898 C}.

48. L. conica von Lendenfeld.

Leucandra conica von Lendenfeld [1885 B].

49. L. catnHA Haeckel.

Leucandra crambessa var. callea Haeckel [1872].

90. L. crossLanpi Thacker.

Leucandra crosslandi Thacker [1908].

D1. L. cytinprica Fristedt.

Leucandra cylindrica Fristedt [1887 ].

02. L. FERNANDENSIS Breitfuss.

Leuconia fernandensis Breitfuss [1898 EK].

53. L. euapiator Dendy.

Leucandra gladiator Dendy [1892 B].

54. L. neatuit Urban.

Leucandra heathii Urban [1905].

55. L. sousrnt Topsent.

Leucandra joubini Topsent [1907 |.

56. L. toricata Polejaeff.

Leuconia loricata Poléjaeff [1883].

57. L. minor Urban.

Leucoma minor Urban [1908].

58. L. puater Breitfuss.

Leuconia platei Breitfuss [1898 EF].

09.

60.

66.

Ms. Ay,

CALCAREOUS SPONGES. 173

. PYRIFORMIS Lambe.

Leuconia pyriformis Lambe [1893].

. RoDRIGUEZIL Lackschewittsch.

Leuconia rodriguezti Lackschewitsch [1886].

. RUDIFERA Poléjaeff.

Leuconia rudifera Polejaeft [1883].

Leucandra rudifera Thacker [1908].

spissA Urban.

Leuconia spissa Urban [1908].

. TAYLORI Lambe.

Leucandra taylori Lambe [1900 A}.

. tyPicA Poléjaeff.

Leuconia typica Poleéjaeff [1883].

. vitREA Urban.

Leuconia vitrea Urban [1908].

Secrron C. With microxea, but without large oxea.

. JOHNSTON Carter.

Grantia nivea var., Johnston [1842].

Leuconia johnston Carter [1871 B}.

Leucandra johnstonii Haeckel [1872].

. LOBATA Carter.

Leuconia lobata Carter [1885-1886].

. MULTIFIDA Carter.

Leuconia multifida Carter [1885-1886 ].

. NIVEA Grant.

Spongia nivea Grant [1825-1826].

Leucandra nivea Haeckel [1872].

. ovata Poléjaeff.

Leuconia ovata Polejaeff [1883].

. PRAVA Breitfuss.

Leucona prava Breitfuss [1898 C}].

Secrion D. Without oxea of any kind.

. BATHYBIA Haeckel.

Dyssycum periminum Haeckel [1870], jfide Haeckel

(1872).

Leucaltis bathybia Haeckel [1872].

Grantia arabica Michlucho-Maclay, MS., fide Haeckel

eas],

The earlier of Haeckel’s names for this species, periminum, is

a nomen nudum, as it never was accompanied by a diagnosis

51%

V74 PROF, A. DENDY AND MR. R. W. H. ROW ON

73. L. BRUMALIS Jenkin.

Leucandra brumalis Jenkin [1908 B].

74, L. curva Schuffner.

Leucandra curva Schuftner [1877].

75. L. Fricipa Jenkin.

Leucandra frigida Jenkin [1908 B].

76. L. aeuatinosa Jenkin.

Lencandra gelatinosa Jenkin [1908 B].

77. L. HELENA von Lendenfeld.

Leucaltis helena von Lendenfeld [1885 B].

78. L. impressa Hanitsch.

Leucaltis impressa Hanitsch [1890].

79. IL. INNOMINATA, sp. 2.

Leucilla crosslandi Row {1909}.

The new name is necessitated by the fact that the name:

crosslandi is already occupied in this genus.

80. L. iwrermepIA Haeckel.

Leucetta pandora var. intermedia Haeckel {1872}.

81. L. Levis Poléjaeff’.

Leuconia levis Poléjaeff [1883].

82. L. naustcaAm Schuffner.

Leucaltis nausicae Schuftner [1877].

83. L. panporA Haeckel.

Leucetta pandora Haeckel [1872].

84. L. pumita Dowerbank.

Leuconia pumila Bowerbank | 1864-1882].

Leucaltis pumila Haeckel [1872].

85. L. saarrrara Haeckel.

Leucetta sagittata Haeckel [1872].

86. L. sCHAUINSLANDI Prewwisch.

Leucetta schawinslandi Preiwisch [1904].

87. L. rELuM von Lendenfeld.

Polejna telwm von Lendenfeld [1891].

88. L. verpEnsiIs Zhacker.

The following species are doubtfully assigned to this genus :—

Leucandra verdensis Thacker | 1908}.

89. L. mnFLata Delle Chiaje.

Possibly identical with Lewcandra aspera, fide Haeckel

Spongia inflata Delle Chiaje [1828].

[1872].

CALCAREOUS SPONGES. 175

90. L. paANicEA Esper.

Spongia panicea Esper, fide Haeckel [1872].

Ksper’s original reference to this species has not been

found by us, but Haeckel [1872] states that the species 1s

possibly identical with Leucandra aspera.

Genus 36. Banria Michlucho-Maclay [1870] (emend.).

Diagnosis. Canal system leuconoid. Skeleton of the chamber

layer composed almost exclusively of irregularly scattered

colossal quadriradiates. Microxea present in large numbers,

and of very characteristic form, being almost always pierced

with a small hole towards one end.

For illustrations of this genus see Haeckel [1872].

The very characteristic ‘‘needle-eye ” spicules of this genus are

really triradiates, in which two of the rays are very much reduced

and have come to le approximately side by side, being actually

fused at their distal ends. In this way we get a linear spicule

very slightly swollen at one end, and in the centre of the swelling

a small hole, the remnant of the space between the two originally

separate rays. That this is the true explanation of these spicules

was made abundantly clear from an examination by one of us

(Row) of a microscopical preparation of the species preserved at

Jena, for while most of the spicules were found to correspond

exactly to the type described above, a few of them had the

reduced rays not fused together but widely open, thus maintaining

the triradiate condition. Exactly similar spicules occur in

Kuarrhaphis cretacea (q. v.).

It may perhaps be pointed out here that these spicules indicate

a possible way in which all the calcareous monaxon spicules may

have originated. At any rate their occurrence adds probability

to the presumption that all calcareous oxea have been derived

from triradiates In some way or other.

The only known species of the genus 1s :—

1. B. ocuorensis Wichlucho-Maclay.

Baeria ochotensis Michlucho-Maclay [1870].

Leucandra ochotensis Haeckel [1872].

Genus 37. LEuUCOPSILA nov.

Diagnosis. Canal system leuconoid. Skeleton of the chamber

layer composed almost exclusively of irregularly scattered

colossal quadriradiates. Gastral cortex well developed, but

without any radiate spicules, the whole of the gastral skeleton

being formed of a dense layer of microxea,

776 PROF. A. DENDY AND MR. R. W. H. ROW ON

For illustrations of this genus see Haeckel [1872].

The very peculiar character of the skeleton of the gastral cortex

seems to justify the erection of a new genus for O. Schmidt's:

Leuconia stylifera.

The only known species is :—

1. L. srynirera O. Schmid.

Leuconia stylifera O. Schmidt [1870].

Leucandra stylifera Haeckel [1872].

Genus 38. APHROCERAS Gray [1858].

Diagnosis. Sponge usually a single person or a colony of such

persons in which the component individuals are readily re-

cognisable. Canal system sylleibid or leuconoid. Skeleton

of the chamber layer more or less confused, but frequently

with vestiges of an articulate tubar skeleton in the form of

subgastral or other sagittal radiates. Dermal skeleton of

tangentially placed triradiates supplemented by colossal oxea

placed longitudinally and not projecting from the surface

sufficiently to render it hispid.

For illustrations of this genus see von Lendenfeld [1891],

under Vosmaeria corticata.

The genus Aphroceras was originally proposed by Gray [1858]

to receive a sponge from Hong-Kong, whose chief characteristics,,

at any rate from our point of view, were the leuconoid canal

system and the colossal longitudinal oxea of the dermal cortex.

Since then other species which combine these characters have

been described, and, although recent authors have not seen fit to

accept this genus, we feel that these species form a very natural

eroup, and we consider that the characters distinguishing it are

sufficiently well defined to render it possible to separate it from

its nearest ally, Lewcandra. In fact, almost the only species that

presents any difficulty is Haeckel’s Leweandra crambessa and its

varieties, in which the colossal oxea are not arranged longi-

tudinally, but lie scattered quite irregularly over the surface of

the sponge. ‘This condition is somewhat intermediate between

that of some Leucandras and that of Aphroceras, and we prefer

to place this species in the genus Leucandra.

It may be advisable to state that we do not consider Aphroceras

to have been derived from U¢e or a Ute-like form, but directly

from an ancestral Leucandra.

We recognise the following species as belonging to this.

genus :—

CALCAREOUS SPONGES. COU

Section A. Without microxea.

1. A. auctcornis Gray. Type species of the genus.

Aphroceras alcicornis Gray [1858].

Cyathiscus actinia, Haeckel [1870], fide Haeckel [1872].

Leucandra aleicornis Haeckel [1872].

2. A. carapHracta Haeckel.

Leucandra cataphracta Haeckel [1872].

3. A. ELoNGATA Schuffner.

Leucandra elongata Schuftner [1877].

Section B. With microxea.

4. A. cusprrosaA Haeckel.

Leucandra aleicornis var. cespitosa Haeckel [1872].

5. A. CLIARENSIS Stephens.

Leucandra cliarensis Stephens [1912].

6. A. corricAta von Lendenfeld.

Vosmaeria corticata von Lendenfeld [1891].

Genus 39. Leucerraca Haeckel [1872] (emend.).

Diagnosis. Canal system leuconoid. Skeleton almost entirely

composed of a confused mass of triradiates, which are mostly

irregular and which form the dermal cortical skeleton as

well as the skeleton of the chamber layer. Gastral cavity

traversed by numerous endogastric septa, which possess a

special skeleton of their own in the form of minute radiates.

For illustrations of this genus see Haeckel [1872].

This genus is proposed for the reception of Haeckel’s Leucetta

pandora var. loculifera, which is the only known species. It

affords an example amongst leuconoid Calcarea of that remarkable

development of endogastric septa which occurs also in certain

Leucosolenias (e. g. ZL. wilsoni) among the Homoceelide, in

Leucetiusa among the Leucaltide, and in Sycandra among the

Sycettide. We have, however, only considered it necessary to

attribute generic importance to this character when it is combined,

as in the present instance, with the presence of an endogastric

skeleton.

We have revived this name, which was applied by Haeckel to

one of the subgenera of his Lewcetta, for this genus.

The only known species is :—

1. L. nocunirera Haeckel.

Leucetta pandora var. loculifera Haeckel [1872].

778 PROF, A. DENDY AND MR. R. W. H. ROW ON

Genus 40. ParatEuctLia Dendy [1892 B].

Diagnosis. Canal system leuconoid. Skeleton of the chamber

layer composed of confused triradiates. Subdermal cavities

present, supported by an inner and an outer layer of quadri-

radiates, whose apical rays cross each other in opposite

directions. Dermal cortex with tangentially placed tri-

radiates, between which lie large, longitudinally placed oxea.

For illustrations of this genus see Haeckel [1872].

This genus was proposed by Dendy [1892 B] to receive Haeckel’s

Leucandra cucumis, but abandoned by him in 1893 on the ground

that it was not sufliciently distinct from Leucilla. He also

pointed out that Poléjaeff had previously proposed the name

Pericharax for the same sponge. Further consideration has,

however, convinced us that the dermal: quadriradiates of

Leucandra cucumis are not really comparable to the subdermal

quadriradiates of Leweilla and other Amphoriscide, being related

solely to the cortex and not to the chamber layer at all. We

therefore transfer the species to the family Grantiide, with which

it has much more in common than with any other family of

Calearea. We know nothing, however, of the position of the

nucleus of the collared cells. The first mentioned species of

Pericharax, on the other hand, is P. cartert Poléjaeff [ 1883, p. 19}

which we have now shown to belong to the Leucascid-Leucaltid

line of descent. We cannot therefore associate Leucandra

cucumis with either Zewcilla or Pericharax, and it therefore

appears to us that as it is clearly distinguished from other

Grantude by the presence of subdermal cavities with a special

skeleton, it is necessary to revive the genus Paraleucilla for its

reception. Haeckel, it is true, placed his Lewcandra cucwmis in

the subgenus “ Leucogypsa,’ but Bowerbank’s name ‘ Leuco-

gypsia,” if revived at all, which we think very undesirable, would

have to be reserved for Leucandra.

The only known species is :-—

1. P. cucumis Haeckel.

Leucandra cucumis Haeckel [1872].

Paraleucilla cucumis Dendy [1892 B}.

Genus 41. Lamontia Kirk [1894].

Diagnosis. Sponge consisting of a single person with a specialised

pore-zone below the terminal osculum. Canal system

leuconoid. Skeleton of the chamber layer consisting of small

scattered oxea. Dermal cortex with triradiates in addition

to oxea. Gastral quadriradiates present.

For illustrations of this genus see Kirk [1894].

CALCAREOUS SPONGES. 779

The only species of this curious genus is Kirk’s Lamontia zona

from New Zealand, which perhaps forms a transition from the

genus Leucandra to the genus Leucyssa.

1. L. zona Kirk.

Lamontia zona Kirk {1894}.

Genus 42, Luucyssa Haeckel [1872] (emend.).

Diagnosis. Canal system leuconoid. Skeleton entirely composed

of smooth oxea.

¥or illustrations of this genus see Haeckel [1872].

We can only suppose that this genus, which has only been

observed by Haeckel, owes its peculiar skeleton to the complete

suppression of ancestral radiates.

The only known species is :—

1. L. sponemtia Haeckel.

Leucyssa spongilla Haeckel [1872].

Genus 43. Tricnocypsi1A Carter [1871 B].

Diagnosis. Canal system leuconoid. Skeleton entirely composed

of spined oxea.

For illustrations of this genus see Haeckel [1872].

In 1871 Carter proposed the genus 77richogypsia for his species

T. willosa. In 1872, in ‘* Die Kalkschwimme,” Haeckel regarded

this species as a variety of his Leucyssa inerustans, the specific

name incrustans having been given by him without description in

1870 under the genus Sycolepis. The diagnosis of Sycolepis also

contained no reference to the essential peculiarities of the species

in question, and was subsequently abandoned by its author.

The name Z'richogypsia has therefore priority in our opinion

over both Sycolepis and Leucyssa, and we have retained it here

for species which, like 7’. villosa, have spined oxea, while using

Haeckel’s name Leucyssa for those with smooth oxea.

We consider that Haeckel’s variety lichenoides, which is the

first variety of his Leweyssa incrustans, 1s specifically distinct from

Carter’s Trichogypsia villosa, and as the name villosa has priority

over incrustans, we confine the latter to the form termed by

Haeckel var. lichenoides.

We recognise the following species of this genus :—

1. T. rycrusrans Haeckel.

Leucyssa incrustans var. lichenoides Haeckel [1872].

780 PROF. A. DENDY AND MR. R. W. H. ROW ON

2. T. vittosa Carter. Type species of the genus.

Trichogypsia:villosa Carter [1871 B].

Leucyssa incrustans var. villosa Haeckel [1872].

Tt is doubtful whether Haeckel’s Sycolepis incrustans is really

a synonym of Zrichogypsia villosa or of 7’. incrustans.

Genus 44. KUARRHAPHIS nov.

Diagnosis. Canal system presumably leuconoid. Skeleton com-

posed exclusively of perforated ‘ needle-eye ” spicules.

For illustrations of this genus see Haeckel [1872].

We propose this name for Haeckel’s Lewcyssa cretacea, which

differs from all the other species included by him in the genus

Leucyssa in the remarkable perforation of the ‘“needle-eye”

spicules. Spicules of practically identical form are found in the

genus Baeria, and we must refer the reader to that genus fora

diseussion of their nature and origin. In Baeria they are still

associated with trivadiates and colossal quadriradiates. Whether

Kuarrhaphis is to be regarded as derived from a Baeria-like

ancestor by the complete suppression of the radiate spicules,

or whether the remarkable ‘“ needle-eye” spicules have arisen

independently in the two cases, it is impossible to decide.

The only known species is :—

1. K. creracea Haeckel.

Leucyssa cretacea Haeckel [1872].

Genus 45. Hinnarptia Poléjaeff [1883].

Diagnosis. Sponge calyciform, with pores on the inner and

oscula on the outer surface of the cup. Canal system leu-

conoid. Skeleton of the chamber layer confused, composed

of triradiates of various shapes and sizes, and of microxea.

Cortex of inner surface with microxea and sagittal trivadiates,.

cortex of the outer surface with large oxea and sagittal

triradiates.

For illustrations of this genus see Poléjaeff [1883].

The only known species of the genus is Poléjaeft’s Lilhardia.

schulzet, a highly remarkable sponge in many ways, the distri-

bution of the pores and oscula being exactly the reverse of the

usual condition in cup-shaped sponges, and indicating that the

sponge cannot be regarded as a single leuconoid person with

expanded osculum, in the same way as the calyciform Pericharax

pexiza.

CALCAREOUS SPONGES. 781

Although the genus Hilhardia was abandoned by Dendy

[1892 5B}, we now consider that it is fully entitled to recog-

nition.

The only known species is :—

1. HE. scnuuze1 Poléjaeff.

Eilhardia schulzei Poléjaeff [1883].

Family 9. AMPHORISCID# Dendy [1892 B] (emend.).

Diagnosis. Flagellate chambers ranging from elongated and

radially arranged to small, spherical and irregularly scattered.

With a distinct dermal cortex supported by a skeleton of

tangentially placed radiates to which oxea may be added.

Some or all of the dermal radiates with large apical rays,

which project inwards through the chamber layer to a greater

or less extent, and form the principal part of its skeleton.

No articulate tubar skeleton, but sometimes, in the leuconoid

forms, a confused skeleton of quadriradiates in the chamber

layer. Nuclei of collared cells probably always apical.

The most conspicuous feature of this family lies in the large

dermal or subdermal quadriradiates with centripetally directed

apical rays. Such spicules may indeed be present in certain

species of Lewcandra, but in such cases they are always associated

with a confused chamber-layer skeleton of scattered triradiates,

which is never the case in the Amphoriscide. If there be a

confused chamber-layer skeleton in this family it is found to be

composed of quadriradiates, which presumably have been derived

from the subdermal and subgastral quadriradiates themselves by

immigration.

The evidence seems to indicate that Zeucandra and Leucilla,

though difficult to separate in practice, owe their resemblance

largely to convergence, and that each has been independently

evolved from some syconoid ancestor, in the one ease directly from

some such form as Grantia, in the other through some such form

as Amphoriscus.

In some Amphoriscide large subgastral quadriradiates are

present, and in others, or even in the same, subgastral sagittal

triradiates (or quadriradiates) resembling the sagittal radiates

of the first joint of an articulate tubar skeleton. Whether the

centrifugally directed ray of the large subgastral quadriradiates

is homologous with the basal ray of the subgastral sagittal tri-

radiates, or whether it is an apical ray added to a tangential

triradiate of the gastral cortex, is a question which we cannot

decide without further evidence.

782 PROF, A. DENDY AND MR. R. W. H. ROW ON

Genus 46. AmpHoriscus Haeckel [1870] (emend.).

Diagnosis. Canal system syconoid. Without any special root-

tuft of anchoring spicules.

For illustrations of this genus see Haeckel [1872] under

Sycaltis and Sycilla, and Poléjaeff [1883].

We recognise the following species in this genus :—

Section A. Without oxea.

1, A. curysauis O. Schinidt.

Ute chrysalis O. Schmidt [1864]. Type species of the

genus.

Sycilla chrysalis Haeckel [1872].

. cyaTuiscus Haeckel.

Amphoriscus cyathiscus Haeckel [1872].

Sycilla cyathiscus Haeckel [1872 .

3. A. cyLtinDrus Haeckel.

Sycilla cylindrus Haeckel [1872].

4, A. Kryprorapuis Urban.

Amphoriscus kryptoraphis Urban [1908].

5. A. oviparus Haeckel.

Sycaltis ovipara Haeckel [1872].

6. A. semont Breitfuss.

Amphoriscus semoni Breitfuss [1896 C].

7. A. TestipaRus Haeckel.

Sycaltis testipara Haeckel [1872].

8. A. uRNA Haeckel.

Amphoriscus urna Haeckel [1870].

Sycilla urna Haeckel [1872].

SEecTION B. With microxea, but without large oxea.

9. A. BUCCICHIE von Hbner.

Amphoriscus buccichii von Ebner [1887 ].

10. A. ELoncatus Poléjaef.

Amphoriscus elongatus Poléjaeff [1883].

11. A. erucorit von Lendenfeld.

Hbnerella gregorii von Lendenfeld [1891 }.

12. A. opiatus Row.

Amphoriscus oblatus Row [1918 MS8.].

CALCAREOUS SPONGES. 783

Genus 47, Sycutmis Haeckel [1872] (emend.).

Diagnosis. Canal system syconoid. With a root-tuft of oxea and

anchoring quadriradiates.

For illustrations of this genus see Haeckel [ 1872}.

This is a highly specialised genus of a single species, but had it

not already been proposed by Haeckel, we should hardly have felt

justified in distinguishing a special genus on the characters

available.

The only known species is :—

1. 8. synapra Haeckel.

Syculmis synapta Haeckel [1872].

Genus 48, Leucinna Haeckel 1872 (emend.).

Diagnosis. Canal system sylleibid or leuconoid. Skeleton of the

chamber layer typically composed of the centripetally and

centrifugally directed apical rays of subdermal and sub-

gastral quadriradiates, but subgastral sagittal triradiates and

confused chamber-layer quadriradiates may be present, while

the subgastral quadriradiates may be absent.

For illustrations of this genus see Haeckel [1872] and Dendy

[1893 A].

The resemblance of this genus to some species of Leucandra

has already been pointed out. It also resembles by convergence

some species of the genus Leucetta, but may be distinguished by

the fact that traces of syconoid ancestry are still to be met with

in the skeleton (e. g., the presence in some species of subgastral

sagittal triradiates), while the triradiates are not of the cha-

racteristic regular type occurring in the Leucascide. The position

of the nucleus of the collared cells, as determined in Leucilla

australiensis and L. princeps, is, moreover, apical, instead of

basal as in the Leucascide.

We recognise the following species as belonging to this

genus :—

Section A. Without oxea.

1. L. AmpHora Haeckel. Type species of the genus.

Leucilla amphora Haeckel [1872].

2. L. AUSTRALIENSIS Carter.

Leuconia johnston var. australiensis Carter [1885-1886].

Leucilla australiensis Dendy [1892 B].

784 PROF, A. DENDY AND MR. R. W. H. ROW ON

3. L. carsuta Haeckel.

Lipostomella capsula Haeckel [1870].

Leucilla capsula Haeckel [1872].

Section B. With large radially arranged oxea or

trichoxea, but without microxea.

4, L. ecuinus Haeckel.

Leuculmis echinus Haeckel [1872].

5. L. OXEODRAGMIFERA Row.

Leucilla oxeodragmifera Row [1913 MS.].

6. L. princers Row.

Leucilla princeps Row [1913 MS.].

. PROTEUS Dendy.

Leucilla protews Dendy [1913].

8. L. urer Poléjaeff.

Leucilla uter Poléjaett [1883].

Section C. Without large oxea, but with microxea.

9. L. Nurriner Urban.

Rhabdodermella nuttingi Urban [1902].

10. L. sAccHarata Haeckel.

Leucandra saccharata Haeckel [1872].

The following species apparently also belongs to this genus, but

was very inadequately diagnosed :—

ll. L. Levconipes Bidder.

ycaltis lewconides Bidder [1891].

Family 10. LELAPIIDA# nov.

Didlytine Kirkpatrick [1911 A].

Diagnosis. Canal system presumably always leuconoid. Skeleton

of the chamber layer containing fibres or bundles of modified

sagittal triradiates placed side by side, but not cemented

together. Nuclei of collared cells (presumably always)

apical.

This family appears to bea highly specialised offshoot from the

leuconoid Grantiide. The presence in Lelapia of distinct sub-

gastral sagittal triradiates and the apical position of the nuclei of

the collared cells afford very strong evidence in support of this

CALCAREOUS SPONGES. 785

view, and necessitate, as we have already pointed out, its wide

separation from the other so-called Pharetronid sponges.

The presence of tuning-fork spicules in Lelapia cannot be taken

as indicating close affinity with the latter, for, as is well known,

such spicules occur in Haeckel’s Lewcandra (Leucortis) pulvinar

and L. (Lewcetta) pandora, while they are replaced in Kebira, the

only other known genus of Lelapiide, by radiates in which the

oral rays have been practically suppressed.

Genus 49, LeLapia Gray [1867].

Diagnosis. Skeleton of the chamber layer composed of large

scattered oxea and loose fibres of tuning-fork spicules.

Dermal skeleton of tangential triradiates and microxea.

Gastral skeleton of tangential triradiates and quadriradiates.

For illustrations of this genus see Dendy [1893 B].

This genus was originally based by Gray on figures published

by Bowerbank of the characteristic tuning-fork spicules. Carter

really first described the sponge, adopting Gray’s name, Lelapia

australis, for the species which he studied. There is, of course,

no guarantee that Carter’s species is either generically or

specifically identical with that which furnished the spicules

figured by Bowerbank. There is some probability, however, from

the locality, and from the fact that the spicules are stated by

Bowerbank to have been “loosely fasciculated,” that the two

species are really identical, and in any case the genus and species

may conveniently be retained for the sponge described by Carter.

The only known species is :—

1. L. austrauis Gray.

‘A new species of sponge” Bowerbank [1858-1862].

Lelapia australis Gray [1867].

Lelapia australis Carter [1885-1886].

Lelapia australis Dendy | 1893 B).

Genus 50. Kepira Row [1909].

Diagnosis. Skeleton of the chamber layer composed of large

longitudinally arranged oxea, and of loose fibres of sagittal

triradiates whose paired rays are vestigial. Dermal and

gastral skeleton of tangential trivadiates.

For illustrations of this genus see Row [1909].

The only known species is :—

1. K. ureorpes Low.

Kebira uteoides Row [1909].

786 PROF. A. DENDY AND MR. R. W. H. ROW ON

GENERA AND SPECIES “INCERT# SEDIS.”

Genus 51. Sycanris Haeckel [1872] (emend.).

Diagnosis? Canal system syconoid (?). Skeleton of the chamber

layer confused.

The sense in which we employ this genus is obviously quite

different from that in which it was used by Haeckel, who based

it upon the presence of triradiate and quadriradiate spicules and

the absence of oxea, Most of the species assigned to it by him

have been relegated to other genera, but his description of the

sundermentioned species appears to indicate that it possesses

characteristics separating it widely from any syconoid sponge

of normal structure, and therefore, although there seems to be

some similarity between this sponge and Leucascus, we prefer

to consider it as “‘incerte sedis” rather than to assign any

definite position to it in our classification. It may be reiated to

Ananiailla.

The only known species is :-—

1. S. pERForATA Haeckel.

Sycaltis perforata Haeckel [1872].

The following species are so inadequately known as to render:

all attempts to identify them ineffectual :—

Ute viridis O. Schmidt [1868].

Medon barbata Duchassaing and Michelotti [1864].

Medon imberbis Duchassaing and Michelotti [1864].

The following species have been referred to without de-

scription :—

Grantia striatula Bowerbank MS8., referred to by Bowerbank

[1864—1882, vol. i. p. 233].

Leucogypsia algoauensis Bowerbank MS., referred to by Bower-

bank [1864-1882, vol. i. p. 166].

Ute papillosum O. Schmidt, referred to by Gray [1867].

(We have not been able to discover any reference to a

species of this name in any of Schmidt’s papers, and think

that Gray’s reference is probably a misprint for Ute

capillosum O. 8.)

LIST OF REJECTED GENERIC NAMES.

The following list includes all the generic names that have, so

far as we are aware, been applied to calcareous sponges, but which

we have not made use of in this paper. Many of them have been

used in various senses by different authors, and in such cases all

CALCAREOUS SPONGES. 187

the instances which are of systematic importance have been

referred to, though the list of references is by no means com-

plete. Under each genus the date and type-species are given, in

order to simplify the work of determining the true names of

genera in accordance with the laws of priority, should any of

these names be revived for future use. Wealso give (in brackets)

the name under which the type species will be found in the present

paper.

ALCYONCELLUM Quoy et Gaimard [1833].

Type species, as regards calcareous sponges, 4. gelatinoswm

de Blainville, (=Sycon gelatinosum).

The name dAleyoncellum was originally proposed by Quoy and

‘Gaimard for certain hexactinellid sponges, but was subsequently

used by de Blainville [1834-1837] to include Sycon (Aleyoncellwm)

gelatinosum. Gray [1867] has also used the name for a genus of

calcareous sponges, and with the same type species. The name is

now regarded as a synonym of both Huplectella and Sycon.

AmpuHoripium Haeckel [1870].

Type species 4. viridis O Schmidt.

Schmidt’s Ute viridis is absolutely unrecognisable, even

generically.

AmpuHorvuLa Haeckel [1870].

' Type species A. solida O. Schmidt, (= Leucandra solida).

ArtyNAS Haeckel [1870].

Type species 4. compressus Fabricius, (=Grantia compressa).

ARTYNELLA Haeckel [1870].

Type species 4. compressa Fabricius, (= Grantia compressa).

ARTYNES Gray [1867].

Type species A. compressa Fabricius, (=Grantia compressa).

Artynium Haeckel [1870].

Type species 4. compresswm Fabricius, (=Grantia compressa).

ARTYNOPHYLLUM Haeckel [1870].

Type species 4. compresswm Fabricius, (=Grantia compressa).

Ascautis Haeckel [1872].

Type species dA. canariensis Michlucho-Maclay, (= Leuco-

solenia canariensis).

Ascanpra Haeckel [1872].

Type species 4. cordata Haeckel, (= Leucosolenia cordata).

The name Ascandra has been used in almost exactly Haeckel’s

sense by several subsequent writers, notably von Lendenfeld,

Breitfuss, Arnesen, and various systematists who have followed

Proc. Zoo. LION. No. LIT. 52

788 PROF. A. DENDY AND MR. R. W. H. ROW ON

the modification of Haeckel’s original system propesed by vor

Lendenfeld in 1891. The name has also been used by Minchin

[1896, 1900, &c.] in an entirely different sense for a genus whose

type, and only, species was A. falcata, placed by us in the genus

Leucosolenia. In the latter sense it is the equivalent of von Len-

denfeld's Homandra.

Ascetta Haeckel [1872].

Type species A. primordialis Haeckel, (= Leucosolenia

primordialis).

As employed by Haeckel, this genus includes only species

whose whole skeleton is composed of triradiates ; von Lendenfeld,

however, has used it [1891] to include all those ascon sponges

which do not possess oxea.

AscitLa Haeckel [1872].

Type species A. gracilistHaeckel, (= Leucosolenia gracilis).

AscometrRA Haeckel [1872].

Type species A. primordiale Haeckel, (= Leucosolenia

primordialis).

The name Ascometira was used by Haeckel, in the “ artificial

system” given at the end of his ‘Monographie,’ to replace the:

name Zhecometra used for exactly the same group in his earlier

‘Prodromus, and for no apparent reason save nomenclatorial

symmetry. He gave his new genus, however, a different type

species.

Ascortis Haeckel [1872].

Type species 4. horrida Haeckel, (= Leucosolenia horrida).

Ascutmis Haeckel [1872].

Type species A. armata Haeckel, (= Leucosolenia armata).

Ascuris Haeckel [1872].

Type species A. arrecife Haeckel, (= a variety of Leucoso-

lenia canariensis).

ASTROSCLERA Lister [1900].

Type species 4. willeyana Lister.

Although originally described as a member of the Calcarea, this

sponge is now known to be an aberrant member of the Non-

ealearea (vide Kirkpatrick [1912)).

AvLopLtecmMA Haeckel [1870].

Type species A. loculoswm Haeckel, (= Leucosolenia loculosa).

Avtorniza Haeckel [1870].

Type species A. witestinalis Haeckel, (= Leucosolenia

lamarckii).

CALCAREOUS SPONGES. 789

Caucisponeia de Blainville [1834-1837].

Type species C. compressa Fabricius, (=Grantia compressa).

This genus was proposed by de Blainyille in exactly the same

sense as Fleming’s Grantia [1828]. Since the latter genus takes

priority, Calcispongia becomes merely a synonym of it.

CiaTHRINA Gray [1867].

Type species C. clathrus O. Schmidt, (= Leucosolenia

clathrus).

This genus has been employed by Minchin [1896, 1900] as the

type genus of one of his families of homoccel sponges. We have

already (p. 718) given our reasons at length for not accepting his

conclusions.

CrystotyntHus Haeckel [1870].

Type species C. vesicula Haeckel, (= Lewcosolenia vesicula).

CaNostoMELLA Haeckel [1870].

Type species C. caminus Haeckel, (= Leucandra caninus).

Canostomium Haeckel [1872].

Type species C’. crambessa Haeckel, (= Leucandra crambessa).

Canostomus Haeckel [1872].

Type species C. primigenius Haeckel, (= Leucetta primigenia).

Cyatuiscus Haeckel [1870].

Type species C. actinia Haeckel, (= Aphroceras alcicornis).

DERMATRETON Jenkin [1908 B].

Type species D. chartacewm Jenkin, (=Grantia chartacea).

This genus was proposed by Jenkin for certain species of the

family Grantiide, which possess ‘‘ linked” chambers. We do not

consider that this character is of generic rank, and we have there-

fore included both these species in the genus Grantia.

Dseppra Michlucho-Maclay MS. (fide Haeckel [1872)]).

Type species D. violacea Michlucho-Maclay MS., fide Haeckel

[1872], (=Grantessa stawridea).

The generic name Djeddew is quoted by Haeckel in the

synonymy list attached to his Sycetta stawridea. This seems’ to

be the only authority for the name.

Dunstervitiia Bowerbank [1845].

Type species D. elegans Bowerbank, (=Sycon elegans).

Bowerbank’s genus was adopted by Haeckel as one of the

genera of his “artificial” system, but no other author seems to

have made use of it save Gray. Itis now considered to be merely

a synonym of Sycon.

52*

790 PROF. A. DENDY AND MR. R. W. H. ROW ON

Dyssycartum Haeckel | 1872}.

Type species D. egedii O. Schmidt, (= Leucandra egedii).

In proposing this genus Haeckel states that it is equivalent to

O. Schmidt’s Sycinula. Both these are considered by us to be

synonyms of Leucandra, but the question is discussed in more

detail under Sycinula.

DyssyconELLta Haeckel [1870].

Type species D. pumila Bowerbank, (=Leucandra pumila).

Dyssycum Haeckel [1870].

Type species D. fistuloswm Johnston, (= Leucandra fistulosa).

Dyssyous Haeckel [1872].

Type species D. primigenius Haeckel, (= Leucetta primigenia).

A genus of the artificial system, identical with Dyssycum of

the ‘ Prodromus’ of 1870. There does not seem to be any reason

for the change of spelling, but such changes, and sometimes more

radical ones, were made by Haeckel in several cases in his later

work.

EBNERELLA von Lendenfeld [1891].

Type species £. buccichti von Ebner, (= Amphoriscus

buccichit).

GuancuHa IMichlucho- Maclay | 1868).

Type species G. blanca Michlucho-Maclay, (= Leucosolenia

blanca).

HereropreeMa Poléjaeff [1883].

Type species H. nodus-gordit Poléjaeft, (= Leucaltis clathria).

As we have shown when discussing Lewcaltis, Poléjaeff’s name

is merely a synonym of the latter.

Homanpra von Lendenfeld {1891}.

Type species H. falcata von Lendenfeld, (= Leucosolenia

falcata).

This genus is the equivalent of Ascandra in Minchin’s sense

[1896, 1900, &e.}], but must not be confused with Ascandra in

the original sense of Haeckel. The latter genus was used by

yon Lendenfeld in almost exactly Haeckel’s sense, so that, since

he considered A. falcata to be worthy of generic separation, a

new name became necessary. Ata later date Minchin revised

the classification of the homoccel sponges, and relegated the

name <Ascandra to A. falcata, so that the name Homandra

was rendered unnecessary. (See, for details, Minchin [1896,

1897].) Wehave given above (p. 720) our reasons for considering

the separation of L. falcata from the genus Leucosolenia to be

inadvisable.

CALCAREOUS SPONGES. 791

Homoperma von Lendenfeld [1885 A].

Type species H. sycandra von Lendenfeld, (=Sycon sycandra).

This genus was proposed for a somewhat aberrant Sycon, and

was made by its author the only genus of a new family of Ascones,

the Homodermide. The question has been dealt with to some

extent above (p. 716), and is fully investigated in Row’s report

on the Calcarea of the Hamburg South-Western Australian

Expedition of 1905 (see Row [1913 MS.]).

Homerra von Lendenfeld {1891}.

A genus proposed by von Lendenfeld on hypothetical characters,

and without any species.

Hypopicryon Jenkin [1908 B].

Type species H. longstaffi Jenkin, (=Sycon longstaffi).

This genus was erected by Jenkin for the above species, and

placed by him in his supposed family Chiphoride, on account of

the presence of chiactines, being separated from Streptoconus on

account of the “linking” of the chambers. Neither of these

characters is considered by us as of generic value.

Hypoerantia Carter [1885-1886 ].

Type species H. infrequens Carter, (=Grantiopsis infrequens).

This genus was proposed by Carter for several species of diverse

nature, and the diagnosis was extremely unsatisfactory. Further,

the first (7.e. type) species was said by its author to be ‘“incertze

sedis.” Under these circumstances we do not feel that we can

allocate the name to any of our genera.

LeucKkarteA Michlucho-Maclay MS., fide Haeckel [1872].

Type species ZL. natalensis Michlucho-Maclay MB8., jide

Haeckel [1872], (=Sycon ramosum).

The only warranty for this name seems to be that it is included

in the synonymy list attached to Haeckel’s Sycandra ramosa as a

MS. name of Michlucho-Maclay’s.

Leucoeyrsia Bowerbank [1864-1882].

Type species LZ. gossei Bowerbank, (= Leucandra gosse?).

One of the four genera of Calcareous sponges proposed by

Bowerbank in his ‘ Monograph of British Sponges.’ It is now

usually considered to be a synonym of Leucandra, and although

perhaps it has right of priority over the latter, yet we feel that

the name Leucandra is so well known that it should be preserved.

Levucomerra Haeckel [1872].

Type species L. primigenia Haeckel, (= Leucetta primigenia).

Levconia Grant [1841].

A genus proposed by Grant and used by many subsequent

authors very nearly in the sense in which we use Lewcandra

792 PROF. A. DENDY AND MR. R. W. H. ROW ON

(e.g. Carter, Poléjaeff, Urban, etc.), but as shewn by Vosmaer

[1887] and Dendy [1893 A], the name is not valid, as it had been

previously applied to a genus of Montusca.

Leucopsis von Lendenfeld [1885 B].

Type species L. pedunculata von Lendenfeld, (= Leucosolenia

pedunculata).

This species was supposed by von Lendenfeld to be a transition

form between Haeckel’s Ascones and Leucones, but it seems to

us that the structure described is far more probably that of a

Clathrinoid Leucosolenia provided with a stalk.

Lervucortis Haeckel [1872].

Type species Z. pulvinar Haeckel, (= Leucandra pulvinar).

Leucutmis Haeckel [1872].

Type species LZ. echinus Haeckel, (=Leucilla echinus).

LipostoMELLA Haeckel [1870].

Type species LZ. clausa Haeckel, (= Leucetta primigenia).

Mepon Duchassaing et Michelotti [1864].

Type species VW. barbata Duchassaing et Michelotti (incerta

sedis).

A genus comprising two species, both of which are quite un-

recognisable, and may even not be calcareous sponges.

Merits Kirkpatrick | 1908}.

Type species M. normani Kirkpatrick.

This sponge was originally described as a member of the family

Pharetronide, but recently Kirkpatrick has shown that its true

place is among the Non-calcarea.

Mura Michlucho- Maclay MS., fide Haeckel [1872].

Type species M/. dohrnii Michlucho-Maclay MS., fide Haeckel

[1872], (=Leucandra pulvinar).

The only authority for this name seems to be Haeckel, who

published several MS. names in the synonymy lists attached to

various species In his Monograph.

Mosrusisponera Duncan [1880}.

Type species JZ. parasitica Duncan.

An organism originally described as a parasitic caleareous

sponge, but it seems very doubtful whether it belongs to the

sponges atall. We certainly do not feel inclined to recognise it,

without further evidence, as a member of the Calcarea.

Narpoa O. Schmidt [1862].

Type species WV. reticulum O. Schmidt, (= Lewcosolenia

reticulum).

Minchin [1896] has shown that this name was previously

CALCAREOUS SPONGES. 793

used for a genus of AsTERoIDEA, and is therefore permanently

unavailable.

Narpoma Haeckel [1872].

Type species WV. nitida Haeckel, (= Leucosolenia nitida).

Narvorpsis Haeckel [1870].

Type species WV. gracilis Haeckel, (= Leucosolenia gracilis).

Narporus Haeckel [1872].

Type species V. primordialis Haeckel, (=Leucosolenia pri-

mordialis).

OLYNTHELLA Haeckel [1872].

Type species O. coriacea Montagu, (= Lewcosolenia coriacea).

OtynrHium Haeckel [1870].

Type species O. nitidum Haeckel, (= Leucosolenia nitida).

Otyntuus Haeckel [1870].

Type species O. simplex Haeckel, (= Lewcosolenia pri-

mordialis).

This name, as an actual generic name, is considered by us to be

merely a synonym of Leucosolenia, but we may point out that its

author proposed it for what he considered to be the most primitive

adult sponge known, and, although we now believe that he

probably erred in considering specimens of this form to be adult,

yet the name is still retained for a hypothetical genus of an-

cestral Calcarea, and for an early stage in the ontogeny of the

individual. Even should adult Olynthus-forms occur, however,

we do not consider that they would be generically separable

from Leucosolenia.

Potesna von Lendenfeld [1891].

Type species P. uter Poléjaeff, (= Leucilla uter).

One of the genera based by von Lendenfeld on the presence of

a sylleibid canal system, a character which we do not consider of

generic importance.

Prosycum Haeckel [1870].

Type species P. simplicissimawm Haeckel, (= Lewcosolenra

primordialis).

RuHABDODERMELLA Urban [1902],

Type species 2. nwttingi Urban, (= Leucilla nutting:).

_ ScypHa S. J. Gray [1821].

Type species SX. coronata Ellis and Solander, (= Sycon

coronatum).

The genus Scypha actually has priority over Risso’s Sycon, but

194 PROF. A. DENDY AND MR. Rk. W. H. ROW ON

the latter has so long been in general use that it seems desirable:

to retain it. Moreover, the earlier name was proposed by a

botanist for organisms which he regarded as plants, and the-

generic diagnosis was hopelessly erroneous and misleading.

Sotenrpium Haeckel [1872].

Type species S. nitidum Haeckel, (= Leucosolenia nitida).

Soteniscus Haeckel [1870].

Type species S’. loculosus Haeckel, (= Leucosolenia loculosa).

SoLeNuLA Haeckel [1872].

Type species S. coriacea Montagu, (= Leucosolenia coriacea).

SPHENOPHORINA Breitfuss [1898 B].

Type species S. singularis Breitfuss, (=Grantia ? singularis)..

This name was originally proposed for a sponge showing certain

apparently great peculiarities in its skeletal structure and in

the form of its spicules. Possibly, if the structure described

really represents that of the sponge, it merits a distinct genus,

but the fact that only a fragment was found, and that that frag-

ment had been preserved in spirit for no less than 33 years before

it was examined by Breitfuss, led us to doubt whether there had

not been some corrosion of the spicules during that time. This

opinion was supported by an examination of type slides which one

ot us (Row) was able to make when in Berlin recently, for the

appearance of the specimens is just what might be produced by

the very slow eating away of the terminal portions of the spicule-

rays by very dilute acid. Under these circumstances we feel that

the genus had better be abandoned for the present, at any rate

until further material of the species assigned to 1t has been ob-

tained, and we have accordingly placed the only described species

provisionally in the genus Grantia.

SPHENOPHORUS Lreitfuss [1898 B].

Type species S. singularis Breitfuss, (=Grantia % singularis).

An earlier name for the previous genus, abandoned by its.

author as being preoccupied.

Sponera Linneus [1758-1759].

The name under which the earlier known species of Calcareous.

sponges were, In common with non-calcareous forms, described,

but now entirely abandoned.

Srreproconus Jenkin [1908 B}.

Type species S. australis Jenkin, (=Sycon australe).

One of the genera of Jenkin’s family Chiphoride, which has.

been abandoned by us for reasons given above.

CALCAREOUS SPONGES. 795

SycantHa von Lendenfeld {1891}.

Type species S. tenella von Lendenfeld, (=Sycon ener),

This genus was retained by Dendy [1892 B], but although the

species placed in it by von Lendenteld is certainly aberrant, we

agree with Jenkin [1908 B] that it does not merit generic

recognition.

Sycarium Haeckel [1870].

Type species S. ampulla Haeckel, (=Sycon ampulla).

Sycriprum Haeckel [1870].

Type species S. gelatinosum de Blainville, (=Sycon gelati-

NOsUnr).

Sycitua Haeckel [1872].

Type species S. cyathiscus Haeckel, (= Amphoriscus cyathiscus).

SYCINULA O. Schmidt [1868].

Type species S. aspera O. Schmidt, (= Lewcandra aspera).

This name, strictly speaking, may have priority over Leucandra,

but it was only mentioned casually by Schmidt, without diagnosis,

although he indicated S. aspera as type of the genus, and it has

never been accepted, except by Haeckel for one of his * artificial”

genera, whereas Leucandra has come into fairly general use.

And further, if the name Lewcandra were changed, it apparently

should be changed to Leucogypsia rather than to Sycinula.

Sycocystis Haeckel |1870].

Type species S. oviformis Haeckel, (=Sycon ciliatwim).

Sycopenprum Haeckel [1870].

Type species S. ramoswm Haeckel, (=Sycon ramosui).

SycotePis Haeckel [1870].

Type species S. inerustans Haeckel, (=either Zrichogypsi«

incrustans or T’, villosa).

Sycomerra Haeckel [1870].

Type species S. compressum Fabricius, (=Grantia compressa).

Syconetta O Schmidt [1868].

Type species S. quadrangulata O. Schmidt, (= Sycon quad-

rangulatun).

SycopHyiuum Haeckel [1870].

Type species S. lobatum Haeckel, (= Grantia lobata).

Sycorruiza Haeckel [1870].

Type species S. coriacea Montagu, (= Leucosolenia coriacea).

Sycortis Haeckel [1872].

Type species S. levigata Haeckel, (=Grantia levigata).

796 PROF. A. DENDY AND MR. R. W. H. ROW ON

Sycortusa Haeckel | 1872).

Type species S. levigata Haeckel, (=Grantia levigata).

One of the subgenera into which Haeckel divided his genus

Sycortis, raised by von Lendenfeld [1891] to generic rank.

Sycornamnus Haeckel [1870].

Type species S. fruticosus Haeckel, (= Leucetta primigenia).

Sycum Haeckel [1870].

Type species S. ciliatum Fabricius, (=Sycon ciliatwm).

A modification of Risso’s generic name Sycon for which there

does not seem to have been any adequate reason.

Sycurus Haeckel {1872}.

Type species S. primitivus Haeckel, (=Sycetta prinutiva).

Tarroma Haeckel [1870].

Type species 7’. canariense Michlucho-Maclay, (= Leuco-

solenia canariensis ).

Tarropsis Haeckel [1872].

Type species 7’. coriacea Montagu, (= Leucosolenia coriacea).

Tarrus Haeckel [1870].

Type species 7’, densus Haeckel, (= Leucosolenia densa).

TEICHONELLA Carter [1878}.

Type species 7’. prolifera Carter, (= Leucetta prolifera).

A genus proposed by Carter for two species, whose slight

similarity of external form misled him into thus associating them.

One of these has now been placed by us in the genus Lewcetta,

the other is the type of our genus Veichonopsis. For a criticism

of the genus Teichonella and its species see Dendy [1891 B].

TENTHRENODES Jenkin [1908 B].

Type species Z’. antarcticus Jenkin, (=Sycon antarcticum).

For a discussion of this genus and its species see under Sycon

(p. 744).

THecomerra Haeckel [1870].

Type species 7’. loculosa Haeckel, (= Leucosolenia loculosa).

Uvetia Dendy [1892 B).

Type species U. hystrix Haeckel, (=Sycodorus hystrix).

We have found it necessary to substitute Haeckel’s name

Sycodorus for the above.

Vosmaeria von Lendenfeld [1885 B}.

Type species V. gracilis von Lendenfeld, (=Grantia gracilis).

One of the genera based by von Lendenfeld on the presence of

CALCAREOUS SPONGES. 797

a ‘“sylleibid ” canal system, a character which we do not consider

to be of generic importance.

WaGNnERELLA Merejkowshi [1878]!

Type species W. borealis Merejkowski.

An organism originally described as a calcareous sponge, but

shewn by Mayer [1879] to be a Heliozoan.

In addition to the above, an enormous number of subgeneric

names, both ‘artificial’ and ‘natural,’ were proposed by

Haeckel in his two works on the group, but the list is sufficiently

swollen out with discarded generic names, without including

subgeneric ones also.

PHYLOGENY OF THE CALCAREA.

Our views as to the phylogeny of the Calcarea, elaborated in

the foregoing pages, may now be summarised as follows, and

illustrated by the accompanying phylogenetic tree. This tree

differs in certain important respects from that published by one

of us twenty years ago [Dendy, 1893 A], which is only to be

expected when we consider the great advances made in our

knowledge of the group in the interval. All the families of the

earlier scheme: and the general ideas of their relationships to

one another in the main lines of descent, are, however, retained

with but little alteration, but we recognise now four additional

families of recent Calcarea, the Leucaltide, the Minchinellide,

the Murrayonide and the Lelapiide, while several genera have

had to be transferred from one family to another.

One of the most important advances in the classification of

the group was made by Minchin [1896], in his demonstration

that even among the homoccel sponges two types of collared cells

are met with, with apical and basal nuclei respectively; a

suggestion which was followed up by Bidder [1898], who, it will

be remembered, proposed to divide, not only the Calcarea, but

the whole of the sponges into BAsINUCLEATA and APINUCLEATA

accordingly, or, confining the suggestion to the Calcarea, to

divide these into CALCARONEA with apical, and CancinEA with

basal, nuclei.

We think it quite likely that the latter of these two suggestions

will ultimately prove to be thoroughly sound. With regard to

the former, however, we consider it highly probable that several

distinct types of collared cells will be shewn to exist in the non-

calcareous sponges, though as yet we have very little information

on this point.

Continuing the observations of Minchin, we find that in the

798 PROF. A. DENDY AND MR. RB. W. H. ROW ON

genus Leucosolenia, which admittedly stands at the bottom of

the line of evolution of the Calearea, both types of collared cells

exist, but apparently not side by side in the same species; and

we find further, that the two principal lines of descent, which

Text-fig. 133.

“PHARETRONES-

PMINCHINELLIDA-

LELAPIIDA-

GRANTIID/AZ

EILHARDIA

neuen 7

SS LEUCETTAGA

ae LAMONTIA

LEUCYSSA

Sd ae

APHIS

Seed —TRICHOGYPSIA

pmTRicy

TEICHONOPS!S MEGAPOGON

GRANTIA

ANAMIXILLA

GRANTIOPSIS

SYCYSSA

LEUCILLA

[ee

! i Aa

ASCUTE

LEUCOSOLENIA

“OLYNTHUS”

HOMOCOELIDAZ

PHYLOGENY OF CALCAREOUS, SPONGES:

appear, on quite other grounds, to have sprung from the homoccel

sponges, are characterised respectively by the two types of

collared cells. It is interesting to note that these two main lines.

CALCAREOUS SPONGES. 799

of descent were clearly indicated twenty years ago, as represented

by the Leucascidee and Sycettide respectively.

It must, of course, be remembered that the real difference

between the two types of collared cells concerns, as Minchin has

shewn [1909], the relation of the flagellum, with its basal granule,

to the nucleus. “his relation has, of course, only been deter-

mined ina very few cases. In ZLeucosolenia coriacea, for example,

the basal granule is situated at the apex of the cell and the

nucleus at the base, while in Z. complicata the flagellum appears

to spring from the nucleus itself, which is apically situated.

There can be no doubt that the actual position of the nucleus

itself in the collared cell may vary temporarily under certain con-

ditions, but in good spirit-preserved material it appears always

to settle down into a characteristic position, which is either

basal or apical, and which may be determined without resort to

special methods of cytological investigation. We do not wish

to lay undue stress upon this character at present, and we

should not venture to use it were it not associated with other

distinctive features, but we have been surprised, in view of our

former opinion as to the systematic value of such a character, to

find how constant the position of the nucleus is in the two lines

of descent indicated. This will be sufficiently evident from

reference to the table given in the Introduction.

We have in vain attempted to split up the unwieldy genus

Leucosolenia into smaller groups. The utmost we have been

able to do has been to isolate from the main body of species three

well-marked types, Ascyssa, Ascute and Dendya. We do not

consider that Minchin’s proposal to divide the homoceel sponges

into two families, Leucosoleniide and Clathrinide, is at all

practicable in the present state of our knowledge, and if it be

true, as he himself has pointed out [1909], following Goldschmidt,

that the two types of relation of flagellum to collared cell may

occur in the same genus of Protozoa (JZastigina), we see no reason

for supposing that both may not occur in the genus Leucosolenia.

According to our view, this is a large and heterogeneous group

of primitive forms all closely related to one another and merging

into one another to a large extent, from which the two lines of

descent referred to have led the way to the evolution of the

higher Calcarea.

We will take the Dendya, or Leucascid-Leucaltid, line first,

in which the nucleus of the collared cells is basal. The starting

point of this lime seems to have been from some form closely

related to Dendya. The radiate arrangement of the colony in

this genus formerly misled Dendy [1893 A] into regarding it as

on the line of evolution of the Sycettide, but there are several

strong arguments against this view. The radiate arrangement

appears to be but a modification of a reticulate ‘‘ Clathrinid ”

character, and actual open anastomoses may occur between the

radial tubes, which, in spite of what has been said by more than

800 PROF. A. DENDY AND MR. R. W. H. ROW ON

one author, appears rarely if ever to be the case in true Sycettide:

or their descendants. To this must be added the primitive type

of skeleton, composed exclusively of equiangular radiates, which

do not exhibit the characteristic arrangement met with in the

syconoid sponges, with their differentiated gastral cortex and

articulate tubar skeleton. In this connection we may especially

note the absence of subgastral sagittal triradiates (or quadri-

radiates), which form such a constant feature of the Sycettide

and their derivatives.

The Dendya line seems to have given off two branches, repre-

sented by the Leucascide and Leucaltide respectively. The Leucas-

cide are undoubtedly the more primitive of the two. The genus

Leucascus itself, indeed, might very easily be mistaken for a

homoceel sponge were it not for the presence of a distinct and

independent pore-bearimg dermal membrane; it retains the

elongated, branched, and more or less radially arranged flagellate

chambers of its Dendya-like ancestors. Within the family evolu-

tion has led to the development of a more highly differentiated

dermal cortex in Lewcetta and Pericharax, accompanied by great:

reduction in the size of the flagellate chambers and complication

of the inhalant and exhalant canal systems. In this way

has arisen that remarkable convergence between Lewcetta and

Pericharax on the one hand, and the leuconoid Grantiide on the

other, which has for so long prevented the appreciation of the

fundamental distinction which really exists between these forms.

The remaining genus in the family, Leawcomalthe, is a highly

specialised and aberrant type, which is only included here

provisionally, until we know more of its minute anatomy and

histology.

In the Leucaltide the distinctive peculiarity has been the

enormous development of the dermal cortex with its special

skeleton, and the accompanying reduction of the skeleton of

the chamber layer to a more or less vestigial condition, or even

its complete disappearance. In this family, again, as regards

canal system, we meet with the customary transition from the

long chambers and radial arrangement of the more primitive

forms (ZLeuwcaltis) to the spherical chambers and _ scattered

arrangement of the highest (Lewcettwsa).

To this line of descent must also be relegated two out of the

three surviving families of ‘‘ Pharetrones,” namely, the Minchi-

nellidee and the Murrayonide. We found this conclusion upon

the basal position of the nucleus in the collared cells in

Minchinella and Murrayona ; but it must be borne in mind that

as regards their general organisation also the members of these

two families differ very widely from Lelapia and Kebira, the

only representatives of the Lelapiide, the third surviving family

of ‘‘ Pharetrones.”

We are therefore compelled to regard the so-called family

Pharetronidx as of diphyletic origin, and the resemblance, such

CALCAREOUS SPONGES. 801

as it is, between the Lelapiide on the one hand, and the

Minchinellide and Murrayonide on the other, as due to

convergence.

As to how many of the vast group of extinct ‘“‘ Pharetronid ”

sponges should be associated with the Minchinellide and

Murrayonide, and how many with the Lelapiidie, is a question

which we cannot attempt to decide, but we think there is evidence

to shew that the great majority belong to the basinucleate group,

though this opinion, of course, rests only on skeletal characters.

It seems highly probable that in past times the Dendya line of

descent led to the evolution of the dominant Pharetronid group,

while at the present day this group has dwindled away and has

been replaced chiefly by the now dominant Grantiide on the

apicinucleate line of descent.

The great antiquity of the Pharetronid group, considering its

high degree of organisation, is highly remarkable. It dates back

far into the Paleozoic Epoch, perhaps even to Silurian times

{| Ulrich, 1889], and almost certainly to Devonian [Zittel, 1878],

so that it seems that the Calearea had already reached one

of their highest states of evolution at the commencement of

the Paleozoic Epoch. Throughout the Secondary Period the

Pharetrones were dominant, and very numerous genera and

species have been described, whereas at the present day they

are almost extinct, though possibly a few more forms yet remain

to be discovered.

Of course it is quite possible that the predominance of the

Pharetrones over other types of Calcarea in past times is apparent

rather than real, owing to the fact that they alone, on account

of their coherent skeleton, had much chance of being preserved

in a fossil condition. Thus there may have been a kind of

‘“‘ ceological selection” of these forms in a fossil condition, but

it is indeed noteworthy that the apparently much more primitive

groups should predominate over these ancient and _ highly

specialised forms to such an extent as they do at the present day.

It is possible that a fresh outburst of evolutionary vigour on

the part of the more primitive persistent groups may have

occurred in comparatively recent times.

Turning now to the Sycettid line of descent, we must remind

the reader, in the first instance, that this appears to have given

rise to the vast majority of the recent Calcarea.

The most primitive genus on this line appears undoubtedly to

be Sycetta, with its radially arranged chambers standing entirely

separate from one another, with no trace of dermal cortex, and

without tufts of oxea at the distal ends of the radial chambers.

This genus already possesses a well-developed articulate tubar

skeleton, the first joint of which is composed of subgastral

sagittal trivadiates, which appear never to have been developed

along the Dendya line of descent, but which are remarkably

constant throughout the whole of the Sycettid line, although

802 PROF. A. DENDY AND MR. R. W. H. ROW ON

absent by suppression in a few cases where the skeleton has

undergone extreme modification. We are unable to indicate

any intermediate forms between the genus Sycetéa and the

Homocelide. It presumably arose from some homoccel ancestor

which formed colonies by radial budding, not unlike those of

Dendya, but the apical position of the nucleus of the collared

cells and the much more advanced type of skeleton shew that

the relationship to Dendya itself cannot be a close one, while the

interval to be bridged over between the most primitive Sycetta

and any Lewcosolenia is a very wide one. Moreover, Sycetta

itself seems to be in the nature of a cul-de-sac, for the entire

absence of the characteristic oxeote spicules of Sycon makes it

doubtful whether we can derive the latter genus directly from

the former, though both have probably sprung from some common

ancestor. If, however, Sycon derives its oxea from an ancestral

Leucosolenia, it is ditiicult to account for the absence of these

spicules in Sycetta, but the distribution of oxea in the Calearea

is an extremely difficult problem about which we have perhaps

said enough in an earlier part of this paper.

The fact that certain species of Sycon, for which von Lendenfeld

[1885 A] proposed his genus Homoderma, retain the collared cells

as a lining to at any rate a portion of the central gastral cavity

throughout life, certainly shews that one can draw no hard and

fast line of distimction between the Homoccelide and the old

group Heteroccela in this respect, but the forms in question have

such a highly specialised syconoid skeletal system that they

hardly help us to bridge over the interval between the Homoceelide

and the Sycettidee.

The family Sycettide is a very small one, the typical genus

being Sycon with a large number of species, while the only other

known genera are S7 ‘ycetta and Sycandra, each with a very small

number of species and each representing an offshoot which

probably leads no further. From the Sycettide two lines of

descent appear to lead to the Heteropiide and Grantiide

respectively.

In both these families the important step in further evolution,

as in the Leucascide and the Leucaltidee, has been the develop-

ment of a dermal cortex, but this cortex appears to have arisen

somewhat differently in the two cases. In the Heteropiidee it is

clearly associated with the out-turning of certain of the oral rays

of the distal tubar trivadiates so as to arch over the entrances to

the inhalant canals. We may assume that with these rays the

dermal tissues of the sponge have spread over the intercanals

and have given rise ultimately to the special cortical spicules

developed in situ. The rotation of the distal tubar trivadiates

in the manner indicated, and the preponderating development

of the now centripetally directed oral rays, have finally converted

these spicules into the ‘‘ pseudosagittal” triradiates which

r\ ey

CALCAREOUS SPONGES. 803

constitute the outstanding feature of the Heteropiide. They

are, so to speak, a new discovery, which the sponge utilizes to

the utmost, until finally their strong centripetally directed oral

rays, In association with the opposed basal rays of subgastral

sagittal triradiates, give rise to an “ inarticulate” tubar skeleton,

which replaces the articulate tubar skeleton of the ancestral Sycon.

The evolution of the canal system within the family appears

to have followed the usual lines up to a certain point. The

known species of Vosmaeropsis exhibit the type of canal system

described by von Lendenfeld as “ sylleibid,” intermediate between

syconoid and leuconoid, but a Heteropiid with a typical leuconoid

canal system has not yet been found.

Tn the Grantiide, on the other hand, the development of a

dermal cortex appears to have been inaugurated by the

appearance of a thin pore-bearing dermal membrane over the

ends of the inhalant canals in some syconoid ancestor, such as

is known to occur in some species of the genus Sycon at the

present day (e. g. S. boomerang).

The Grantiide must be regarded as the dominant family of

Calcarea at the present day, comprising, as they do, no less than

23 genera, and exhibiting a very wide range of structure both

as regards skeleton and canal system. It might be possible to

divide these genera into syconoid and leuconoid subfamilies, but

we should have no guarantee of the monophyletic origin of the

latter from the former. Nevertheless, the scarcity of sylleibid

forms, which might be regarded as connecting links between the

two types, is somewhat remarkable, and suggestive of a natural

cleavage. The known species of the genus Megapogon, however,

are sylleibid, and although most of the Leucandras have small

chambers, Z. australiensis Carter and L. infesta sp. n. (= Leucilla

intermedia Row [1909]) have very large ones, and might also be

considered as transitional forms. Also we must remember that

in a considerable number of cases we have no accurate information

as to the canal system.

The arrangement of the genera within the family is a very

difficult problem. They appear to group themselves around two

central types, Grantia and Leucandra, but as we have just

mentioned, our knowledge of the exact type of canal system in

many forms is very imperfect, while in others the only evidence

of their close relationship to Lewcandra is the canal system itself.

The appearance of this part of our phylogenetic tree will pro-

bably be greatly modified by subsequent investigations. Such

aberrant genera as Leucopsila, Baeria, Kuarrhaphis, Leucyssa

and Trichogypsia can only be included in the Grantiids pro-

visionally.

The difficulty of arranging the genera probably arises from the

fact that great gaps exist in the family owing to extinction

of intermediate forms. It might be argued that this family

Proc. Zoou. Soc.—1913, No. LILI. 53

804 PROF. A. DENDY AND MR. R. W. H. ROW ON

cannot be very ancient, because hardly any of thei fossil remains

have been discovered, but this may possibly be accounted for by

the fact that they do not possess a coherent skeleton, and

accordingly become disintegrated soon after death.

It is, however, quite possible that some of the very imperfectly

known extinct Pharetrones may really be offshoots from this

family, as we believe to be the case with the recent Lelapiide.

The customary association of the latter with the Pharetrones

is based upon the presence of a fibrous skeleton composed of

modified radiates, especially the tuning-fork spicule, and a

fibrous skeleton of this type 1s perhaps present in some of the

extinct genera. We have, however, purposely avoided discussing

the latter in this paper, as we do not know enough about them

to warrant us in drawing any but the most general conclusions.

As we have already pointed out, the apical position of the nucleus

of the collared cells in Lelapia, and the presence in it also of

subgastral sagittal triradiates, render the Grantiid origin of the

Lelapiide reasonably certain.

Tn addition to the Lelapiide, one other family, the Amphoriscide,

seems to have originated from the Grantiid stock. This family

derives its distinctive character from the development of strong,

centripetally directed apical rays on the tangential radiates of

the dermal cortex. Such apical rays have undoubtedly appeared

several times independently in the evolution of the Calcarea.

We find them, for example, in some species of Leuwcetta, in

Leucaltis, in some species of Leucettusa, in one species of

Grantia (G. intermedia), in some species of Leucandra, and m

Grantilla, as well as in this group. It may well seem doubtful

whether, in view of these facts, the character in question ought

to be regarded as of family significance in the Amphoriscide ;

but inasmuch as the latter appear to us to comprise a natural

assemblage of three closely related genera in which this character

is no longer casual but has assumed great importance in the

structure of the skeleton, we have decided to retain the group,

at any rate for the present. We have, however, considerably

reduced the size of the family by the removal of Lewcaltis

(Heteropegma), together with some of the species formerly

placed in Leucilla, but now divided between Leucetia, Leucettusa

and Leucandra, to which they seem to be more closely affiliated

by other characters, leaving in the genus Lewcilla a group of

species which are, we believe, all directly descended from

Amphoriscus.

The most primitive Amphoriscide, belonging to the genus

Amphoriscus, have a syconoid canal system and a somewhat

feebly devel6ped dermal cortex, and we accordingly consider the

family to be an offshoot from low down on the Grantiid stem.

Finally, we may point out that in this family again the canal

system has undergone its usual transformation from the syconoid

to the leuconoid type, with intermediate sylleibid forms.

CALCAREOUS SPONGES. 805

BIBLIOGRAPHY.

The following list of literature includes only those papers which

are actually referred to in the text, as it has been found impossible

to give anything like a complete list of the papers which deal with

calcareous sponges in the space at our disposal. However, there

will be found to be but few papers of real systematic importance

which are not mentioned, and, for obvious reasons, all papers

which describe new species are included, at any rate so far as we

are aware, and we have no reason to fear that our list is not

complete.

When more than one paper has been published by an author in

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1901. ArRNeEsSEN, Emily. <‘‘Spongier fra den norske kyst. I.

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bestemmelsestabel.” Bergens Mus. Aarbog, 1900 (1901),

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1834-1837. Buatnvitixn, H. de. ‘ Manuel d’Actinologie et de

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1845. BoweErsanx, J. 8. ‘Description of a new Genus of

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1858-1862. “On the Anatomy and Physiology of the

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Part I. Phil. Trans. Roy. Soc., London, vol. exlviii.

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Part Il. Phil. Trans. Roy. Soc., London, vol. clii.

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806 PROF. A. DENDY AND MR. R. W. H. ROW ON

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* Ascandra Hermesi, ein neuer homocceler Kalk-

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“ Amphoriscus semoni, ein neue Art heteroceeler

Kalkschwimme.” Zool. Anzeiger, Bd. xix. pp. 435-

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1898 B. —— “ Kalkschwammfauna des weissen Meeres und

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“ Die Kalkschwimme der Sammlung Plate. (Fauna

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1871 A. Carver, H. J. ‘On two undescribed Sponges and two

Esperiade from the West Indies; also on the nomen-

clature of the calcisponge Clathrina, Gray.” Ann. Mag.

Nat. Hist., (ser. 4) vol. vii. pp. 268-283.

“A description of two new Caleispongie (T'richo-

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Professor James-Clark’s discovery of the true form of

the Sponge-cell (Animal) and an account of the Polype-

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Prof. E. Haeckel’s view on the relationship of the

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Zool. Soc., London, 1858, pp. 113-114.

1867.

1821.

1870.

1872.

1877.

1908.

1890.

1894.

1895.

1886.

1900.

1869.

1908 A.

1908 B.

1842.

1893.

1894;

1895.

CALCAREOUS SPONGES. 809

Gray, J. E. ‘Notes on the Arrangement of Sponges,

with descriptions of some new genera.” Proc. Zool. Soc.,

London, 1867, pp. 492-558.

Gray,S.F. “A natural arrangement of British Plants.”

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Hazcken, E. ‘Prodromus eines Systems der Kalk-

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‘“* Die Kalkschwiimme, eine Monographie.” Berlin,

1872.

“‘ Die Physemarien (Haliphysema und Gastrophy-

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vol. xi. pp. 1-54.

Hammer, EH. ‘Neue Beitrige zur Kenntnis der

Histologie und Entwicklung von Sycon raphanus.”

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Haniiscu, R. “Third Report on the Porifera of the

L.M.B.C. district.” Proc. Liverpool Biol. Soc., vol. iv.

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—- “Amphiute, eine neue Gattung* heteroccler

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‘“‘ Notes on a collection of Sponges from the West

Coast of Portugal.” Trans. Liverpool Biol. Soe., vol. ix.

pp. 205-219.

Hiaern, T. ‘ Porifera of the L. M. B.C. District. First

Report upon the Fauna of Liverpool Bay.” Liverpool

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Hinpr, G. J. ‘‘On some Remarkable Calcisponges from

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JaMES-Cuark, H. “On the Spongie Ciliate as Infusoria

Flagellata; or, Observations on the Structure, Ani-

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Memoirs Boston Soc. Nat. Hist., vol. i. pp. 805-340.

JENKIN, C. F. “The Calcareous Sponges. (in) The

Marine Fauna of Zanzibar and British East Africa, from

Collections made by Cyril Crossland, M.A., in the

Years 1901 and 1902.” Proc. Zool. Soc., London, 1908,

pp. 434-456.

“The Calcarea of the National Antarctic Expe-

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Jounston, G. “A History of British Sponges and

Lithophytes.” Edinburgh, 1842.

Krrx, H.B. ‘Contribution to a knowledge of the New

Zealand Sponges.” Trans. New Zealand Lnstit., vol. xxvi.

pp. 175-179.

“Further Contribution to a knowledge of the

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vol. xxvil. pp. 287-392.

‘New Zealand Sponges. Third Paper.” Trans.

New Zealand Instit., vol. xxviii. pp. 205-210.

810

1895.

1900.

1908.

1910.

TOMAS

1911 B.

1912.

1864.

1886.

1893.

1896.

1900 A.

1900 B.

1885 A.

1885 B.

PROF. A. DENDY AND MR. R. W. H. ROW ON

Kirk, H. B. “ Notes on New Zealand Sponges. Fourth

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316.

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362.

“On two new Genera of recent Pharetronid

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pp. 003-514.

“On a remarkable Pharetronid Sponge from

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pp. 124-133.

“On a new Lithonine Sponge from Christmas

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“On Merlia normant, a sponge with a siliceous

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(n. s.) vol. lvi. pp. 657-702.

“Note on Astrosclera willeyana, Lister.” Proc.

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Konimiker, A. von. ‘‘ lcones Histologice, oder Atlas der

vergleichenden Gewebelehre.—I. Der feineren Bau der

Protozoen.” Leipzig, 1864.

Lackxscurwitscn, P. ‘ Ueber die Kalkschwimme Menor-

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530.

1758-1759. Linn mus, C. von. “Systema Nature.” (16th Edition.)

Holmie, 1758-1759.

1868.

CALCAREOUS SPONGES. 811

Lister, J. J. ‘ Astrosclera willeyana, the Type of a new

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pp. 70-77

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Museums der Kaiserlichen Akademie der Wissen-

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1869.

ON CALCAREOUS SPONGES.

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dition of H.M.S. Alert in the Straits of Magellan and

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1881, pp. 107-139.

‘““Spongida.” Reports on the Zoological Collections

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von Sycandra raphanus Waeckel.” Zettschr. wiss.

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No. 59.

THacker, A. G. ‘On Collections of the Cape Verde

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B.Se. (Lond.), F.Z.8. (ate of the Gatty Marine Labora-

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1904. The Calcareous Sponges.” Proc. Zool. Soc.,

London, 1908, pp. 757-782.

Toprsmnt, E. “ Eponges calcaires recueillis par le Francais

dans l’Antaretique (Expédition du Dr. Charcot).”

Bullet. Mus. Hist. Nat. Paris, 1907, pp. 539-544,

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Silurian Sponges.” American Geologist, vol. ii. pp. 233—

248,

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 813

1902. Urpan, fF. “ Rhabdodermella nuttingi, nov. gen. et nov.

spec.” Zeitschr. wiss. Zoologie, Bd. lxxi. pp. 268-275.

1905. “ Kalifornische Kalkschwimme.” Archiv fiir

Naturgesch., Jahrgang 72, Bd. i. pp. 33-76.

1908. “Die Kalkschwimme der deutschen Tiefsee-

Expedition.” Zoolog. Anzeiger, Bd. xxxili. pp. 247-252.

1909. “Die Calcarea.” Wissensch. Ergebnisse der

deutschen Tiefsee-Hapedition (Valdiia), Bd. xix. Jena,

1909.

1873. Verrinn, A. KE. ‘“ Exploration of Casco Bay by the U.S.

Fish Commission in 1873.” Proc. American Assoc.

Advane. Sci. 1873, Part 2, pp. 340-395.

1880. Vosmarr,G. C.J. “Ueber Leucandra aspera H., nebst

alleemeimen Bemerkungen ueber das Canalsystem der

Spongien.” Leiden, 1880. (dn wtaugural Dis-

sertation.)

‘“Porifera.” Die Klassen und Ordnungen des

Thierreichs, wissenschaftlich dargestellt in Wort und

Bild. Von Dr. H. G. Bronn. Bd. i. Leipzig und

Heidelberg, 1887.

1878. Zinrnn, K. A. von. “Studien ther fossile Spongien.

Ill. Monactinellide, Tetractinellide, und Calci-

spongiv.” Abhandl. Akad. Wissensch. Miinchen. vol. xiii.

part 2, pp. 1-48.

1887.

EXHIBITIONS AND NOTICES.

May 20, 1913.

Prof. HK. A. Mincnin, M.A., F.R.S., Vice-President,

in the Chair.

The Secretary read the following report on the Additions that

had been made to the Society's Menagerie during the month of

April, 1913 ==

The registered additions to the Society’s Menagerie during the

month of April were 205 in number. Of these, 95 were acquired

by presentation, 79 by purchase, 9 were received on deposit, | in

exchange, and 21 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 156.

Amongst the additions special attention may be directed to :—

1 White-handed Gibbon (Hylobates lar), from Penang, deposited

on April 10th.

814 ON DATES OF PUBLICATION OF TRANSACTIONS,

1 Maholi Galago (Galago maholi), from 8. Africa, presented by

Lady Yule, F.Z.8., on April 8th.

1 Bangs’s Agouti (Dasyprocta colombiana), 1 Bonda’s Squirrel

(Scturus saltuensis bonde), from Colombia, both new to the

Collection, presented by W. K. Pomeroy, Esq., F.Z.S., on April

21st.

A Collection of Birds, from North-West India, presented by

Major G. A. Perreau, F.Z.S., on April 1st, of which the following

are new to the Collection :—

1 Dark Grey Bush-Chat (Oreicola ferre), 1 Plumbeous Redstart

(Lhyacornis fuliginosa), 1 Red-flanked Bush- Robin (Janthia

rufilata), 2 White-tailed Blue Robins (WVotodela leucwra), 1 Great

Niltava (Niltava grandis), 2 Short-billed Minivets (Pericrocotus

brevirostris), 1 Larger Red - headed Crow - Tit (Scaorhynchus

rujiceps), 1 Cinnamon-bellied Nuthatch (Sitta cinnamomeiventris),

1 Grey-sided Laughing Thrush (Dryonastes cerulatus), 4 Rufous-

necked Laughing Thrushes (Dryonastes rujicollis).

1 Bushmaster (Lachesis mutus), from Trinidad, presented by

the Baron K. G. K. Leijonhufvud, F.Z.8., on April 28th.

1 large Anaconda (Hunectes murinus), from Trinidad, purchased

on April 28th.

The Rev. T. R. R. Srespsine, M.A., F.R.S., F.Z.8., drew

attention to Prof. F. K. Schulze’s important work on zoological

nomenclature, the ‘ Nomenclator Animalium,’ giving a_ brief

description of its objects and scope, and urging its claims for

assistance from British naturalists.

The LiprariaAn submitted the following list of dates of publi-

cation of the early parts of the Society’s “‘ Transactions,” which

had been drawn up from the records kept by Messrs. Taylor and

Francis, the Society’s printers.

The date of issue of Parts subsequent to Vol. VII. Part 1

appears at the foot of each sheet.

VouumeE I.

Part. Pages. Plates. Published.

il, 1-90. I.-XIT. Aug. 14, 1833.

2. 91-194. XITII.-XXVI. Apr. 23-25, 1834.

3, 195-800. XXVIT.-XXXVITI. Mar. 20; 1835.

4. 301-407. XXX VITI.-LIX. Dec. 3, 1835.

Vouume IT.

le 1-86. T-XVII. Oct. 2, 1836.

2 87-164. XVITI.-XXTX. May 4, 1838.

3: 165-248. XXX.-XLV. Dec. 5, 1838.

4, 249-342. XLVI.-LVIII. Apr. 6, 1840.

5. 343-420, LIX.-LXXT. May 15, 1841.

Part. Pages.

1-132.

So trea ae

1-30.

31-74.

75-88.

89-148.

GO IS OT G9 bo

1-32.

33-102.

ee Sh

1-48.

49-86.

87-124.

G0 I OTR Ge bo

I 1-144.

133-234.

235—276.

277-344.

345-380.

381-431.

149-182.

183-268.

269-352.

303-382.

103-242:

243-336.

337-421.

125-226.

227-308.

309-376.

377-494.

495-537.

ON GROWTH OF ANTLERS.

Vouume III.

Plates.

IL Wl

VIII.-X VIL.

XVIII-XxXX.

XXXI.-LI.

LIT-LVIL.

LVI1.-LXITITI.

Vouume LV.

I-VI.

IX.-XXV.

XXVI.-XXX.

XXXJ.-XLII.

XLIUI-LIT-~.

LIV .—-LXITI.

LXIV.-LX VII.

LXVIla.—-LXXVIT.

VoLuME V.

T—-XITI.

XIV .-XXVI.

XXVIT.-XLUIT.

XLIIL-LI.

LITT.-LX VII.

Vouume VI.

T-XIV.

XV.-XXIV.

XXV.-XXX.

XXXITL-XLI,

XLIV.-LIV.

LV.-LXII.

LXIIT-LXXXVIT.

UXXXVITT.-XCII.

Vouume VII.

= WIL,

815

Published.

June

Jan.

June

Dec.

16, 1842.

23, 1844,

5, 1844.

28, 1846.

. 22, 1848.

30, 1849.

1, 1851.

2, 1852.

, 29. 1853.

, 10, 1856.

. 28, 1858.

. 31, 1859.

e. 23, 1861.

. 15, 1862.

11, 1862.

97, 1863.

. 28, 1864,

5, 1865.

6, 1866.

. 15, 1866.

lu, WG.

, 28, 1867.

5, 1867.

>. 19, 1868.

4, 1868.

_ 15, 1868.

1, 1869.

. 20, 1869.

Mr. R. EK. Honpine exhibited a number of antlers, skulls, and

photographs illustrating variations in the growth of the Antlers of

Deer, and stated that antlers were liable to considerable variation

in form, due either to exuberance or proliferation of growth, or

to congenital defect in the embryonic stage as in the case of the

“Hummel” or Hornless Stag (text-fig. 134 A), which occasionally

made its appearance in Scottish and continental Deer forests.

816 MR. R. E. HOLDING ON

Text-fig. 134.

gat) P|

ea “ ‘

E 553

& \

yy i) Vy :

ie ‘i 4,

>I)

REX det

A. Upper portion of the skull of a “ Hummel” or Hornless Stag, aged 5 years.

B. Upper portion of the skull of Red Deer (Cervus elaphus), with deficient right

and arrest of left antler.

Ci. First antlers of young Wapiti Stag (Cervus canadensis), showing unusual

growth of nodules and “ burr.”

GROWTH OF ANTLERS. 817

Sometimes the arrest was on one side only, producing the Single-

horned Stag as shown (B). Occasionally a small supernumerary

horn might grow below the Burr or Coronel (6), as shown in the

figures of Muntjac (BE) and Red Deer (D), probably due, as in the

Four-horned Sheep, to accessory centres in the pedicle or horn

support. Other variations, and by far the most common, were due

to dichotomy, bifurcation, or segmentation of the beam or of

the tines. That favourable environment and ample food promoted

variation in the direction of exuberance or proliferation of points

was well demonstrated by the photographs from Warnham Court

and by illustrations in Mr. Winans’ book ‘“ Deer-breeding for

Fine Heads,” published by Messrs. Rowland Ward. On the other

hand, head-keepers and others in charge of Scottish Deer forests,

had observed that scarcity of food, hard weather, and exposure

have a marked effect on the antlers of Deer, causing deficiency

in growth, bad form, and degeneration.

It was not often, at least in Scotland, that a stag with a good

head escaped the attention of the owner or lessee of the Forest,

and therefore defects and variations due to extreme age were

seldom seen in the Red Stag, but in the Park Fallow Deer a

marked change took place in the form of the palmation of the

antler due to advancing age.

Malformations and variations due to accident, castration, or

wounds, were not referred to, as being pathological.

Of the specimens exhibited in illustration of his remarks,

Mr. Holding drew special attention to the antler of a Sambur

stag (J), showing excessive proliferation of points, and an

exceptionally fine pair of antlers lent by Mr. C. J. Lucas, of

Warnham Court.

Explanation (continued) of Text-fig. 134 (see opposite).

D. Head of Stag shot in Jura Forest, showing supernumerary right antler (a) growing

from a separate pedicle on the frontal bone.

Muntjac (Cervulus muntjac), showing supernumerary horns (4) springing from

the long pedicle.

F. Head of Fallow-buck (Cervus dama), showing dichotomy or bifurcation of beam

of right antler at the “burr.”

G. Antler of Axis Deer (Cervus axis), the bifurcation occurring above the burr.

H. Head of Red Stag from Warnham Park, having 42 points. From photograph by

J. G. Millais, Esq., F.Z.S.

I. Head of White-tailed Deer (Cariacus leucurus), showing excessive proliferation.

Shown by Capt. Frewen at a recent Exhibition of American trophies.

J. Antler of Sambur Deer (Cervus unicolor), showing profuse growth of “points ”

obliterating the normal character of the antler.

818 MR. R. LYDEKKER ON THE

June 3, 1913.

Prof. E. W. MacBrips, M.A., D.Sc., F.R.S., Vice-President,

in the Chair.

Mr. D. Sera-Suira, F.Z.S., Curator of Birds, exhibited the

egg and young of the Mikado Pheasant (Calophasis mikado), a

rare species, described first in 1906, from the mountains of

Formosa. Some living specimens had been imported in 1912 by

Mr. Walter Goodfellow, and the owners of these birds had

entrusted the eggs to the Zoological Society, where they were

being hatched.

The egg was cream-coloured and very large compared with

those of allied species of pheasants, measuring 57 x 41 mm.

The incubation period proved to be twenty-eight days, instead

of twenty-four as in the majority of pheasants, and the young

when newly hatched were very large, and had the quil-feathers

better developed than was the case in allied species.

PAPERS.

45. The ‘Transvaal Race of the Cape, or Khama, Hartebeest *.

By R. LypKexesr, F.R.S., F.Z.S. f

[Received April 1, 1913: Read April 8, 1913.]

(Text-figure 135.)

INDEX.

Page

Description of Bubalis caama selbornei ............ 819

In their excellent volume, ‘The Sportsman in South Africa,’

Messrs. Nicolls and Eglington describe (p. 45) the Cape, or Rooi,

Hartebeest (Lubalis caama), probably from fresh specimens, as

follows :—

“General colour reddish brown, with violet tinge throughout ;

dark plum-coloured saddle-patch, commencing at point of shoul-

der, extends over entire surface of back and ends in root of tail ;

a similar patch extends over each shoulder, downwards as far as

knees, and front of shin-bones of fore-legs; pale yellowish patch

on cheeks of rump; front of face, which is very long, almost

black, as is a stripe down back of neck.”

In their figure of the head (pl. iv. fig. 13) the broad face-blaze

is shown as extending uninterruptedly from the base of the horns

* [The complete account of this new subspecies appears here, but since the name

and a preliminary diagnosis were published in the ‘ Abstract,’ No. 119, 1918, it is

distinguished by being underlined.—Eprror. |

+ By permission of the Trustees of the British Museum.

CAPE HARTEBEEST. 819

to the muzzle; but, in many cases at any rate, this is divided by

a narrow tawny line at the level of the eyes. They omit te

mention a plum-coloured patch on the side of the thighs below

the light area, extending to the hocks.

In the ‘ Book of Antelopes,’ by Messrs. Sclater and Thomas, an

obviously over-coloured plate (no. iv.) of the entire animal is

given, in which the dark face-blaze, divided by an interocular

hght bar, and the dark markings on the fore and hind limbs are

clearly shown, although there is no sign of a plum-coloured

saddle-patch. The authors describe the general colour as brown-

ish fulvous, darker than in any other member of the genus.

The only specimen of an adult male Cape Hartebeest from

Cape Colony in the British Museum is one obtained by Sir

Andrew Smith, which has recently been dismounted and con-

verted into a flat skin. Although much faded by long exposure,

it serves to show that the type of colouring was originally much

the same as in the plate in the ‘ Book of Antelopes.

A few years ago Lord Selborne was good enough to offer to

endeavour to procure for the Museum specimens of such South

African Antelopes as might be required for public exhibition ;

and as a result of his lordship’s request the skin, skull, and horns

of an adult male of the Transvaal representative of the Cape

Hartebeest were received at the Museum in 1912, as a gift from

the De Beers Mining Company. The specimen w as in due course

set up by Rowland Ward, Ltd., and placed on exhibition in the

galleries in lieu of the old and faded example from Cape Colony

referred to above. At the time I was busy with other matters,

and consequently did not pay any attention to the details of the

new acquisition. Recently, however, I have had occasion to

review all the Hartebeests in the collection; and this survey has

left no doubt that the Transvaal Hartebeest, which was shot in

the neighbourhood of Kimberley, represents a very distinct

undescribed race of Bubalis caama.

From the typical Khama this race differs by its much paler

general colouring, which is yellowish fawn or tawny, not unlike

that of B. cokei, by the minor development and intensity of the dark

markings, and by the apparently less sharp definition of the white

on the sides and back of the lower part of the rump. The general

tawny tint tends to chestnut on the loins, as in B. cokei, but

elsewhere the back is of much the same colour as the flanks. The

nuchal stripe is indistinct ; and the face-blaze, instead of forming,

with the exception of the narrow light band between the eyes, a

continuous wholly black streak from the horns to the muzzle, is

much broken up, and everywhere mingled with fawn- coloured

hairs. It practically stops short of the horns, and is interrupted

in the neighbourhood of the eyes by a fawn area, mingled with

a few blackish hairs, for a length of about four inches, while the

nasal portion does not nearly reach the muzzle. ‘Then, again,

the dark patch on the shoulder and fore-leg is much less intense

than in the typical race, and is everywhere mingled with fawn

Proc. Zoo. Soc.—1913, No. LIV. 54

820 ON THE CAPE HARTEBEEST.

hairs, while it stops short of the knees, on which it forms a dark

patch, and on the shanks is represented solely by a very small

patch at about the middle of their length. The same lack of

intensity characterises the dark area on the thighs, which

appears to be smaller than in the typical southern race.

Text-fig. 135.

Transvaal Khama Hartebeest (Bubalis caama selbornei).

The Transvaal race, which is typified by the aforesaid mounted

buck, and may be named Dubalis caama selbornei [Abstract

PeZoSo UGNGs [Pl LAKIN.

G. Dollman, del. Henry Stone & Son, Ltd., Banbury

1. SOREX GRANTI. 2. S. ARANEUS CASTANEUS.

ON HEBRIDEAN MAMMALS. 821

P. Z.8. 1913, p. 19 (April 15)]|, cannot, indeed, be described

as a “ Rooi” (red) Hartebeest ; and is evidently a form

which is either losing or acquiring—I cannot say which—the

rufous body-colour and deep purplish-black markings of the

typical southern Khama. The two races may be defined as

follows :—

a. General colour rich rufous-brown; face-blaze black and ex-

tending, with the exception of a narrow fawn band between

the eyes, from horns to muzzle; limb-markings plum-colour,

and forming a continuous stripe on front of fore-legs.

B. caama typica.

&. General colour yellowish fawn; face-blaze mingled with tawny.

stopping short of horns and muzzle, and interrupted by a

long interval in region of eyes; limb-markings mingled with

tawny, interrupted above knees, on which they form a cap,

and represented by a small patch on front of shanks.

B. caama selbornet.

46. Ona Collection of Mammals from the Inner Hebrides.

By G. E. H. Barrerr-Hamitton, F.Z.S., and Marrin

A. C. Hinton.

[Received and Read April 8, 1913.}

(Plate LXXXITV.* and Text-figures 136-138.)

INDEX.

Systematic : Page

SOPH GIRUMGU, SDs Ts 2 e-cosdcsonercsaa4ab0 000 seauvac0n venues tery

JISOCOMGS GUSTO) Vo Vs sgnaceeadcaeceocopecncceccossuoce, SA

Microtus agrestis macgillivrati, subsp. n. ......... 831

The peculiar mammals of high interest yielded by investigation

of the peripheral regions of the British Islands, led Barrett-

Hamilton to believe that further valuable results might be

obtained by a closer study of the mammals of the Scottish Islands.

The despatch and management of collecters was, however, an

undertaking beyond his means and available time, and the

matter hung fire until taken up by Mr. W. R. Ogilvie-Grant,

whose enthusiasm as manager and collector of funds has made

it possible to amass the material on which this paper is based.

Karly in 1912 Mr. R. W. Sheppard was sent to the Inner

Hebrides, and commencing to collect at Bute, this young naturalist

gradually worked his way northwards through the group as far as

Coll, which he reached in July. As small mammals are difficult

* For explanation of the Plate see p. 839.

b4*

822 MAJOR G. BE. H. BARRETT-HAMILTON AND

to catch during the summer months, further investigation of the

islands was then postponed.

Among those who have assisted the work of exploration, either

financially or with local assistance, we may specially mention

the Duke of Argyll, the Marquis of Bute, Mr. A. R. Macgregor,

Mrs. Hugh Morrison of Islay, Mr. Colin Campbell of Jura, and

Mr. Harold Russell.

The present collection has come well up to expectations, since

it has resulted in the discovery of three quite distinct forms

belonging to the genera Sorex, Hvotomys, and Microtus*.

In working out the fauna of small islands, two distinct phe-

nomena must be disentangled. A peculiar mammal found on

a small island may represent a new development under insular

conditions, it may represent the survival of an ancient type else-

where extinct, and, as is often the case, it may combine both

features. We believe that Sorex grantii is an entirely new de-

velopment, and that Hvotomys alstonti and Microtus agrestis mac-

gillivrati are instances of ‘“ relics” surviving with little alteration

from the pleistocene epoch. The meaning of these three forms

has been partly discussed under the description of each of them ;

but it would be premature to enter upon an extended discussion of

the cause of variation or survival in islands. The influence of a

comparatively recent glacial period, and the extermination brought

about by man, now proved to be an ancient inhabitant of Britain,

are difficult to follow; and at all periods the British Islands have

been the scene of constantly shifting environments and altering

or variable climates, due principally to the proximity of a great

ocean and the frequent oscillations of sea-level. Many of our

common genera of mammals have inhabited the British Islands

since the deposition of the late pliocene Forest-Bed, but no one

has as yet ventured to guess how often they may have been

exterminated, to return again in fresh vigour on the resumption

of favourable conditions. The history of the British fauna

resolves itself into that of the various invasions or ‘‘ migrations ”

of mammals, which have been no less real, albeit more ancient

and more difficult to trace, than those of Kelt, Roman, Saxon,

Skandinavian, and Norman. ‘The severity of the struggle for

existence has in our area undergone profound variation, accord-

ing as the sea cut off or united our area with Europe, according

as severe climatic conditions were combined with insularity or

connection with the Continent. But, however frequently the

fauna may have been exterminated, renewed junctions with

the mainland have from time to time refreshed it with new

stock, and the geological records of what seem from a distance

almost kaleidoscopic changes are only now in process of

disentanglement.

* [The complete account of these new forms appear here, but since the names and

preliminary diagnoses were published in the ‘ Abstract,’ No. 119, 1913, they are

distinguished by being underlined.—Eprror. |]

MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS. | 823

SOREX ARANEUS CASTANEUS Jenyns. (Pl. LX XXIV. fig. 2.)

Three, GREAT CUMBRAE. Dimensions * : —

Head & Hind

body. Tail foot

No. 34, male, 28 March 1912 ...... 70 30 12

32, female, 27 e Feed ‘Nee 70 33 RS

BS) wine Oe Be A midds Soa 67 32 11

Average of 3 specimens of both sexes:— 69 32°6 12

Hight, Bure.

NowlSe males) 19eMaxeh:-[9i2) so. 70 Bie 12

Gomme SWAY eam eae 00 33 12

Osis Ona SOI a cat ee 70 32 12

(Als ae 20 ienee PaMsoos re 70 36 12

M4 temale, l9Miarch) 4) 222 65 32 ih;

15) ee oe * See NR AN 65 32 12

HORN ee.§ : Se wae iL ee 65 33 12

GOyeane. bi ANCA O EGE Corn ee. 73 32 12

Average of 8 specimens of both sexes:— 69°37 33°37 12

Three, ARRAN.

INowaSy malese NO esspril, LOZ es. Wi2 36 12

ie hemealen Mees ea TEES 70 36 12

CORO: one SEAT ee ( 36 12

Average of 3 specimens of both sexes:— 72°33 36 12

Four, Muu.

INos 143, male, 29-June 1912 |... 78 35 12

144, ,, SOM cs scin are 75 33 12

1I38\temale, Zilkume ys. .-..-- 68 40 12

1 ec Wee ep a ee 68 38 12

Average of 4 specimens of both sexes:— 72°25 36:5 12

Six, JURA.

Nos OSS males la aMiay TOMO Sees. 70 36 12

TON es Rid rid tee hegat at Boas 73 35 12

TPs | gp Gee haved: Sarin ee 76 36 12

11, temalesionys has! | tet ch 73 36 12

LAO: tibiae WAMU bem s -eussy) | meee 70 33 12

OU RONDE Wot Aas TAsiot Y3Ey 4 NO

Average of 6 specimens of both sexes:— 72°5 35:3 12

For cranial measurements see Table at p. 828.

The specimens from Great Cumbrae, Bute, Arran, and Mull do

* Collectors’ measurements: throughout hind feet without claws, tail without.

terminal hairs. All dimensions are given in millimetres.

824 MAJOR G. E. H. BARRETT-HAMILTON AND

not differ appreciably from examples of castaneus from the neigh-

bouring mainland. Skulls of castanews from South-eastern

England seem to attain slightly larger dimensions than those

from northern Britain, and this may indicate the existence of two

forms. The Jura Shrew is interesting in having a pelage inter-

mediate in character between that of castaneus and grantii de-

scribed below; in the four skulls seen the dental characters are

those of normal araneus, and therefore this form must be left

with the latter species.

Sorex Grant. (Pl. LX XXIV. fig. 1, & text-fig. 136.)

Abstract P. Z.S. 1913, p. 18 (April 15).

Twenty-two, Isnay. Dimensions :—

Head & Hind

body. Tail. foot

INosiiommaleye 24 sApril 1912) eee 36 13

(as 7 : Sig," tact 00 36 13

(ET 26 fae Ie 79 Br 13

SOs 95 BO on Pst bce 80 36 13

SOR ie igen He gan Tl 2 80 36 13

SB, 59 : ai a aa i ee 80 37 13

Sau 5 Yel bi seth. ae 78 38 il

OO. 2 Mays inn aaa 78 RD 12

DB. as Bana ah CS C® 35 12

OS os (ee eh irene: 75 35 12

100, 55 Su ve Pe aie TA ie 75 35 12

Ol,” 4 ae suites 78° 8B 12

NOS; 55 HO: ayes achat Aes BRE 75 36 13

i2mtemialles 3 Aspyr; tees naire 83 37 See

Oman eieee Op) St Reg Ma 75 OM 13 Type.

SH eee tate 3 (ii yy ba Sg SE pace 78 36 13)

OB, bg 3 May Chim sdain’ a 35 ib

OG Ra ee O: £5). oe leave 75 35 12

157, female, juv., 3 Aug.1912 ... 75 38 12

158, 99 op. 5) 9 oF NEKO) 34 12:5

161, cf Ee es Dante reams AS) 36 12

165, i ee Ot eamn Bae x Neen) 38 12

Average of 18 adults of both sexes:— 77°3 36 12°6

For cranial measurements see Table at p. 828.

This is a very remarkable new Shrew, distinguishable at a

glance from the common British S. araneus castaneus by its dusky

upper side in strong contrast to the light flanks. The dentition is

peculiar in the fact that more than half of the individuals have

lost the posterior upper unicuspid, sometimes on one side only,

but more frequently on both sides, the absence of this tooth in

these cases being compensated for by enlargement of the four

remaining unicuspids

MR. M. A. GC. HINTON ON HEBRIDEAN MAMMALS. 825

Description.—The size and proportions are about as in

S. araneus castaneus, but average slightly larger than in north

British specimens of the latter.

Colowr.—Adults have the upper side deep blackish brown,

perhaps darker than ‘‘clove-brown”*; slightly grizzled on the

upper side of the head and neck with brown; the under side is

silvery or smoky grey rather than yellowish or brownish as in

castaneus, and this colour runs far up the flanks and shows up in

contrast to the dark upper side. Between the colours of upper

and under sides a narrow, inconspicuous, grey-brown flank-band

intervenes; rarely it is ‘‘ wood-brown,” in which case it contrasts

more sharply with the back. Nos. 72 & 76 (23 & 26 April) are

moulting into a scarcely less dusky summer coat, but some speci-

mens taken in May and Nos. 157, 161, & 165 (3, 5, & 8 August)

are ina lighter brown coat, between “seal-brown” and ‘ clove-

brown,” somewhat as in S. a. castaneus, with a flank-band near-

‘““ wood-brown.” These specimens are no doubt immature and the

brown pelage is that of immaturity.

Cranial and dental characters—The skull agrees in form and

size with that of S. a. castaneus, but attains rather larger dimen-

sions than skulls of the latter species from northern Britain and

the Inner Hebrides (vide Table, p. 828). The teeth agree in form

with those of S. araneus; but the posterior unicuspid (p. 3) tends

to disappear as shown by the following statistics :—

p- 3 present on both sides in 7 individuals, 7. e. 43°7 per cent.

. » ononesideonly in 4 ee ss 25 g

.. absent on both sides in 5 % SB 1LEB}" op

Text-fig. 136.

Right upper tooth-row of Sorex grantii. xX 8b.

When p. 3 is absent the four remaining unicuspid teeth appear

to be somewhat enlarged in compensation (text-fig. 136). There

is no trace of pigment on the hypocones of the upper cheek-teeth

nor on protocone of the last upper molar.

By colour and dentition S. grantii is clearly differentiated from

all Huropean members of the genus Sorex. Some of these, such

as araneus araneus and araneus tetragonurus, have the back as

dark, but none shows the conspicuously contrasted sides.

* The names of colours in inverted commas are from Ridgway.

826 MAJOR G. E. H. BARRETT-HAMILTON AND

In our opinion S. grantii is to be regarded as an insular

development of S. a. castaneus stock. It has maintained or

acquired slightly larger dimensions than has the latter in the

more northern parts of its range; it has developed a peculiar

colour pattern, and is well on its way to reduce the number of its

unicuspid teeth, perhaps because of a tendency to enlarge the

anterior members of the series. The Jura Shrew is geo-

graphically and morphologically the connecting link between

S. a. castaneus and S. grantii: it has the small size and dental

characters of the northern castanews; but its colour pattern is

modified so as to approach that of S. grantii.

The species is named in honour of Mr. W. R. Ogilvie-Grant of

the British Museum, to whose enthusiasm the formation of the

collection described in this paper is so largely due.

SOREX MiINuTUS Linneeus.

Nine, Bute. Dimensions :—

Head & Hind

body. Tail. foot.

No. 18, male, adult; 20 March 1912... 52 35 10

ts ay a Qt Bee Gade SOO 35 10

FORA i 20 April Bate 8 OD 34 10

2, female ,, 2s dMiarcluth ia |... 4 32 10

fhe Sartore A 20y aes b Gdk TOD: 35 10

OS eee aaa a 55 i OO) 33 10

Dangers S e be Sua eo) O) 35 10

Dona mes 8 * * seh een et 55) 35 10

GOs gg gp AD eyoraail Pete aire). 35 10

Average of 9 specimens of both sexes:— 53 aye 2 KW)

All in dark adult pelage, between ‘“ seal-brown” and “ hai-

brown.”

Two, ARRAN.

No, 59, male, adult, 11 April 1912 ... 58 35 10

51, female, ,, Oo sf 59 35 leat

Average of 2 specimens :-— 58° 35 10°5

In dark adult pelage as above.

Three, Istay.

No, 75, male, adult, 26 April 1912 ... 60 40 10

Sy 95 POO) betes iNet Oe) 37 10

86, female, ,, “8 As Lee MeN 5574 Bel 10

Average of 3 specimens of both sexes:— 586 38 10

No. 85 is in dark adult pelage as above, Nos. 75 and 86 in

juvenile or summer pelage.

One, CoLu.

No. 56, female, adult, 23 July 1912... 60 Ef 10

MR. M. A. C. HINLON ON HEBRIDEAN MAMMALS. — 827

Five, TrreEe. Dimensions :—

Head & Hind

i body. ail. foot.

No. 152, male, juv., 16 July 1912... 50 34. 10

Genie Se lal eli ia ahaa aan 1)

154, ,, - Wider se Met | e508 TO) 34 10

OO! Js a Whe ane ae le) 34 10

151, female, fully |

mae pe gen He 60! 38! sypi0

The larger specimens from Coll and Tiree are in the light

summer coat of young born in the same season; in this pelage the

hairs are more plentifully grizzled or peppered with tawny than

in adults. The very young specimens show an interesting early

pelage, in which the upper side is lighter than in adults, the under

side very clear and nearly white.

For cranial measurements see Table at p. 828.

Elsewhere in the Hebrides S. minutus is known to inhabit Skye,

Higg, Sanda, Great Cumbrae, Lewis, North and South Uist,

Benbecula, and Barra. (Barrett-Hamilton, ‘ History of British

Mammals,’ 11. pt. 8, pp. 111-115.)

EyoroMYs GLAREOLUS BRITANNICUS Miller.

Two, Butr. Dimensions :—

Head & Hind

bodys. Mails toot. aHars

No. 6, male, adolescent, 13 March 1912. 90 42 17 10

Zils eee echullit: PA soaker 5 Smo Bir. IS, yO

For cranial measurements see Table at p. 830.

With this material we are unable to distinguish the Hvotomys

of Bute from the form inhabiting the mainland. In No. 6 the

cement-spaces of the cheek-teeth are just closed; in No. 27 the

roots of the cheek-teeth are moderately developed. In each

specimen the last upper molar has a third inner fold. The

skull agrees in size and general form with that of britannicus,

but the external measurements indicate a rather smaller animal

with slightly larger hind feet than the latter.

Hyoromys atstont. (Text-fig. 137.)

Abstract P. Z.S. 1913, p. 18 (April 15).

Five, MULL. Dimensions :—

Head and Hind

bodyzu stan toot.) Har:

No. 134, male, aged, 18 Junel1912 108 44 18 11 Type

See, Cr ee pape lel O emme Aan: Oreille

SOs iss aR OW Gis a LOSE aa Ss Ae

128} female) (ABiaw;,, cpp OO eA On) i Bro neleliga

LSKOS Se ie ales eee 55 BMOO BOA i aS (0)

Average of 5 specimens of both sexes: 105°2 43°38 184 11

For cranial measurements see Table at p. 830.

828

MAJOR G. E. H. BARRE'TT-HAMILTO

N AND

Cranial and Dental measurements of Sorex.

Ke Fas a

ni g le

— a 8 o & | | LES I

be =| | g z fe 2 | 2 | * Be a

Aya | S| al elena ee BP Vise 12 Si 2

rai | Sl || oS | oat ose | oe Er @ Ss || ey le Si)

S| = |. & co Sh SU A lic |= |e ey 8

Ee aee| ee | — | Sula Aue ey evel

SUE ES OMS | 5 | EONS 8) ee caloN ais ica Ponte

S.aranus |>|\/2|e\|e)5/2)|4 | 4/3/\/4|2) 2 Bel 2

castaneus. |) 12) 2) P/E |S )8) SB) Sie) = le sies

S|) 2 2 Seles | Sq Sue ie lok | =

URA. oS a ee aes a eae

IN@s dL opocee 18°7 5°06 3°5 9:0 47 | 8°2 | 4:4 | 6:93) 9°3 (5-17 | 3°52) 4°51) 2°8 | 1°32

Prin 2 Olen 18:3: 5°17) 3°5 | 9°2 | 5:0 | 7-7 ize 66 9:3 5:17 | 3°52) 4:29] 2°75) 1°43

JLL. | | | | | |

Now14 ona 18°7/ 4°84) 3:5 |9°0 | ... |8:1 | 451] 6:93) 9-4 [5-28 |3°63) 4-4 | 2°86) 1°32

NAS: s56be 19:0) 5°28) 3°9 |9°6 | 5:1 | 81 | 44 | 7°04 9°8 [5°28 | 3°57) 4°51) 3:08) 1°32

ee 17°7, 484/33 89 4:9 | 7-6 4:18] 6-49 9°0 [4:95 | 3°52) 4-4. | 2°86 1°32

4), | | |

IN@ BB sesso 18°4 5°06 3°5 19-2 | 4:8 | 8-0 | 4:4 | 6°93 9°3 5°17 | 3°63) 4-4 | 2°86) 1°32

MARR ee): 18°4) 4:95 3°5 |9°0 | 4°8 | 7°9 | 44 | 6°82: 9°3 (5°06 | 3°52) 4rd. | 2°97) 1°32

Syria 17°9| 5:28 3°5 | 9:1 | 5:0 | 7°8 | 429] 6°71 9:0 [5°06 | 3°52) 4-45) 2°86) 1°32

Gir “Sosseo 18°5| 5:12 3°5 |9°2 49 7:9 | 4:29) 6°82 9°2 |5°17 | 3°63) 4-4 | 2°86) 1°43

68 118: H 5:06 3:5 | 92 52 | 7-7 | 434/665 91 5:06 | 3°52 4-29] 2°91) 1:32

CUMBRAE, | | | acid mea ie

iNowoO ee | 187) 5:06 3°7 | 91 7°8 | 4:29] 6°71/9:4 5:06 | 352) 44 | 2°91) 1°37

34 18°9) 5°28 3:7 |... | ... | 7:9 | 4°29] 6:9319°7 15°17 | 3°62)4°51) 2:97) 1:32

SON Rees 18° 3) 5:06 3°53 | 89 48 | 8:0 | 4:29) 6°71'9°5 4:95 | 3°46) 4-4 | 2°86) 1°32

ARRAN. | | aes

IN@s BY soo o05 18-2) 506136 |89 ... | 7°8 | 4:29) 6°71 5:06 | 3°46) 4°51) 2°97) 1:43

GIS) sop ave 18°6) 5°22, 3°6 | 9°4 |5°0 | 8:0 | 44 | 6°93 9°3 5:11 | 3'57| 4°62) 2:97) 1:37

S. granti.

ISLAY. |

No. 73 .| 1971 5:28 3°7 |9°3 | ... | 8:1 | 4:4 | 6°93) 9°6 5°17 | 3°52) 4°51) 2°75 1-48

Ty eee 19°83) 5°39 3°8 | 9°6 5:0 | 8°4 | 4°62) 7:15 9°9 5:28 | 3°63 4°55) 2°8 1:43

MN TWO” 7B © sasooe 19°0 5°28 3°8 (91 52 | 83 | 4°62) 6:93 9:9 517 | 3°57) 4°51) 2:97) 1:32

Ml 18°9) 5°39) 3°8 | 9°3 | 5:1 | 8:2 | 44 | 682 9°7 5°17 | 3°63) 4°56) 2°86 1esy7/

80 18:8) 5:28) 3°8 | 9:4 | 5:2 | 81 | 44 | 6°82, 9°6 5°17 | 3°63) 4°62| 2°97 1:43

Sou raat 19:0|5°5 | 3°8 |9°2 51 | 84 | 4°62) 7-2 | 9°9 5:39 | 3°63) 4°62) 2°97 1:43

SA: 191 5:28 3:8 |9'2 52 82 | 4:51) 6:93 98 5:28 | 3°63) 462] 2°97 1:43

95 18°6| 5°17 3°8 |} $9 5:0 | 81 | 4:4 | 6:82 9:4 5-17 | 3°63) 4°51] 2°86 1°32

HOW scsccc 18°2,6:17 3°5 |90 5:0 | 7°8 | 4°29) 6°71) 9:4 817 | 3°57 4:4 | 2°75 1:37

OMe eeeee 18°5 5°33: 3°7 | 9°2 5:0 | 7:9 | 4:29) 6°71 9:6 \5°06| 3°46 4°67|2°5 | 1°43

NOS. cceoce 18°4 5°39 3°8 | 9°4 |50 | 7:9 | 429/66 9°6 4°06 | 3°52) 4-62) 2°86 1°32

NOS soacoe 18°6| 5°17 3°7 | 9:1 |5:1 |.7:9 | 44 | 6°71|9°3 (5°17 | 3:57| 4-4 | 2°86 1°32

S. minutus. LA WHE. I

BUTE. el

Nowr2) gine 15:2) 407) 2:7 | 7°3 | 35 | 65 [3525-4 |7°3 4°07 | 2°97) 2:97 214 088

¢/ 15°5| 4°18) 2°7 | 7:2 | 4:0 | 6:6 | 3°74! 6°61) 7°7 |4°29 | 3°08! 3:08) 2-2 0°88

24 15°2| 3°96 2°7 |72 | 4:0 | 63 | 3°52) 5°39 7:4. 4:07 | 2°86) 2°97| 2°14 0°88

DASA ewe 15:2] 3:96 2°8 | 7:3 | 4:0 | 6:3 | 3°52] 5°39) 7°4 4°07 | 2:97 2:97) 2°14 0°88

26 15:0) 3°96) 2°8 |'7°3 | 4:0 | 6:2 | 3°47| 6°8 | 7°3 4:07 | 2°86) 2:97) 2°2 0°88

CORP ae 15°3] 3°96] 2°7 | 7:1 | 4:0 | 6°83 | 3°52) 5°5 | 7-4 4:07 | 2:97} 3°08) L 98 O'77

ARRAN. | | |

No. 51 159) 418 2°8 | 7-1 4:0 | 68 | 3°85 5°83 7°8 4:18) 3:08) 3°08 2:2 | 0°88

59 .| 15:9] 4°18) 2°8 | 7-2 | 4:0 | 6:6 | 3°74) 6°72) 7-8 4°29 | 3°14) 3:08) 2:2 | 0°88

ISLAY. |

No. 85 | 15:4) ... 127 |7-2 | 4:0 | 65 | 8°74) 6°61) 7°6 4:23 | 3:08) 3:08) 2:25, 0-94

| SGN ieee 15°6) 4:18 2°8 73 (40 | 65 (3°74 561,76 418 | 3°08 3°08) 2°31) 0°88

* These measurements were made with an eyepiece micrometer and 5-inch objective,

vide Hinton, “ British Fossil Shrews,” Geol. Mag. n. 8., dec. 5, vol. vil. pp. 537 & 538.

MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS. 829

General characters.—This mouse is a little larger than LZ.

glareolus, approaching EL’. norvegicus, but the tail is shorter than in

the latter, and about as in glareolus; smaller than H. skomerensis

or cesarius. ‘The ears are about as in #, glareolus. The hind

feet are as in norvegicus and skomerensis.

Cranial and dental characters.—In size the skull is larger than in

H. glareolus and agrees with that of norvegicus. As in the latter

form, the jugals are heavy, but the curvature of the zygomatic

arches agrees with giareolus. The brain-case is very broad and

Text-fig. 137.

Dorsal view and profile of skull of Evotomys alstoni. Natural size.

smoothly convex, the temporal ridges being but faintly indicated

even in aged skulls: the parietal region is convex in dorsal profile.

with the highest point a little behind the middle of the parietals,

instead of flattened ; these features impart an appearance of

relatively greater cranial capacity than is seen in any of the

other European species of Hvotomys. The postorbital (squamosal)

processes are not conspicuous. The interorbital region is broad,

with a wide shallow median sulcus. The nasals are rounded

or slightly and narrowly emarginate behind, ending flush with

or slightly behind the ends of the premaxille; they are slightly

longer than the diastema, expanded in front, their lateral borders

slightly but distinctly concave. The rostrum is shallow, as in

EL. norvegicus, its least depth behind the incisors not exceeding

the anterior width. The bulle are nearly as in norvegicus. The

last upper molar has a third inner fold and fourth inner salient

angle—the latter being usually well developed.

The colowr is much darker than in /. skomerensis, but 1s very

similar to that of #. glareolus, being deep russet above ; the under

side is richly washed with yellowish or buff.

This is an interesting and quite distinct species, recognizable

externally by its generally large size combined with small ears

and short tail, while in addition its skull characters are quite

distinctive. On the whole it makes a nearer approach to H. nor-

vegicus than to any other species of Hvotomys.

830 MAJOR G. E. H. BARRETT-HAMILTON AND

€

Cranial measurements of Hvotomys.

eee | x

ele | are | 2 \2

oD ae | dc) 5 = 4

2 | *S | a a | = | | at : 5

eeu t| SSlch ea te Ome iS aha e ere

ese een ea ol Sess P=surs l= 2

|e ps sees a |>S| 6 a=

eee |e era ene

S| See Se eS he |e Is

6 5 lS (Se EAE h eae Seale

peGlarmoritaniicss ===. 2a |2s ee a a a

| | | | |

Chingford, EssEx.........) 24°2 14:0) 3°9/ 11:2) 60 68 65) 55 145 51 Aged. |

Bure, No. 27, male ...... | 23°5| 12°8) 3°8 11-2) 6:0) 6:7 | 64 5°4/14°6 5:4) -Adult. |

| | | | |

| |

. | | | | | |

E. alstoni. -———|—— — ny S| Site

Mutt, No. 134, male ...| 25°3) 18°9| 8°8)11°5 64) 7°5| 7-1) 56/156 5-4) Aged. |

ss S60 (2573/1473) 3:9)| 12-1, 6:61) 7-81) 7271) bez) e671 4) ee

ee 136, ,, _.../ 249145 41/11°8 63) 7:3) 68) 56 15:3 51)

ye 130, female .| 24"1) 135) 3°9| 11-4) 6:0) 6'8| 63| 52|1571| 54)

lie icone

EH. norvegicus. fe | a ne | ee eee

Hardanger, 5.8.5.2 ...... 24°5, 134) 3°8/ 11-1] 6:0! 6:9) 7:0| 5:°4| 15:3) 51) Aged.

Bergen, max. (Miller) .... 26°2 14°6 4°0| 12:0 6°6 80) 7°8| 58160 54) Adult.

MIcROTUS AGRESTIS EXSUL Miller. (Text-fig. 138 a.)

Four, Muu. Dimensions :—

Head & Hind

bod\zylbarlearetootameeLan:

No. 139, male, ad., HY wine WOU, MAD. (BG UD 12

NL se ie oy OTe aa ike a LOO “SO . Me

132, female, subad., 15 _,, us Iss NasA By ee

145, a UNE 3 July as OEY GO hss)

Hight, ARRAN.

No. 41, male, ad., Pyare We = [PAS aI IN) Le

AOS Tae ae sVULV See % NOG wage UO sl

AG eemesuioaGs. se4 o ey BBM) oclkss

DOL ee 3 Dy, ae : WAS 2 Way AS) LE

S25 ks 3 Ongar = Greta Gs The

G25 Seems peel yee 5 NAGY SG) IG 4s

AS Lemaleweiives | oO)», r NOB BR. WO Ll

AU. Tees ACen. GA.” ee ; Ud Leagan lle WLS): 9 hel

(damaged)

One, GIGHA.

No.127, male, adult, 23 May 1912. 125 33 18 12

MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS. 831

Hight, Jura. Dimensions :—

Head & Hind

body. Tail. foot. Bar.

No l00S male ssubad Lo May 1912) 15s "30 19 1s

TESS . +5 lO >; i AS ee MOR we 2

116, 3 “A if so i TUS BY OAD). cess

ina Stenralesre Ori: ss IPAQ) SVE co ME A

Sea BA aan aS Ss TIL a4 TE) 112

EAD tee lie. Pee . IAG BOQ Wes We

Tite Tenemaene eter ts oak Mi (N95), (TLOMAMiTS

124 oo LPs bed asl Sa ew est Hunts alls Dl aOMeaAS

For cranial measurements see Table at p. 834.

M. agrestis exsul Miller, of which, in addition to the material

in the British Museum, we have examined several specimens

from North and South Uist, kindly lent by Mr. Eagle Clarke,

of the Royal Scottish Museum, Edinburgh, is a thick-coated form

with light upper side and flanks, and under side heavily washed

with buff. The fur of the back in winter reaches a length of

about 12 mm., including the hght tips of about 2-3 mm.; the

longer hairs reach about 15 mm. Weare not able to distinguish

the specimens from Mull, Arran, Gigha, and Jura from this sub-

species. Those from Mull are practically quite typical.

The following Table shows the character of the anterior upper

cheek-tooth in the skulls examined :—

Mutu. ARRAN. GicHa. JURA.

Ath inner angle quite absent... 1

represented by microscopic ‘

; VESEICO aed as eo Porcine 2 yee 12 : ae

5p RCTEMIL LONG GUIS BUOVCE 5 sconecoae 1 4 3 8

Specimens examined ...... 5) 9 1 8 21

Some features of the skull are discussed below in connection

with the form living in Islay.

MIcROTUS AGRESTIS MACGILLIVRAI. (Text-fig. 138 B.)

Abstract P. Z.8. 1913, p. 18 (April 15).

Fourteen, Isuay. Dimensions :—

Head & Hind

body. Tail. foot Ear.

No. 78, male, subad., 26 April 1912. 115 34 19 13

OA siesshs | sae iamles 3 i OB ellis “anal

Hy Ween exohribin’ SON, LAOS A Ride P12

BOC vig cls nel barca te, el PELE SOM ee

OOo Me, 2 May LB SYP IG) 1

vlna 25), 1132) edi yaeertlyspe

832 MAJOR G. E. H. BARRETT-HAMILTON AND

Head & Hind

body. Tail. foot. Har.

No.103, male, juv., @) Jileng TESA IO aah MS) TL

MOST as excholke, IMO) 8 Ff Wy ase) Ns)

159, ,, subad., 3 August ,, WO, aie IS)

NOS Am 5s a3 (ee me . ie 8H IS le

87, female, , 30 April - ay Say UO

I 4,0 adulty 2 May. re IPAQ By IDS

160. |e subbadk., 3 August 4 Itsy ee IG)

G2 es Fan 46 e a WO aa It

For cranial measurements see Table at p. 854.

This mouse resembles J/. agrestis exswl in size and general

proportions, but may be recognized at all seasons by its much

thinner coat and the much lesser amount of yellowish wash on

the under side, resulting in the slaty bases of the hairs being

visible and taking part in the general coloration; the under side

thus resembles that of immature exsul. The upper side is slightly

less brightly coloured than in easwl; the hairs of the back reach

a length of about 8 mm.

Text-fig. 138.

Dorsal view and profile of skulls of (A) Microtus agrestis exsul and

(B) UM. a. macgillivraii. Natural size.

A quite young male (No. 79) is in dusky pelage, with very

slight development of the ight hair-tips. The hair-tips of the

under side are of a very ight tint, increasing in depth through

the flanks to the back and poll, where they are deepest but never

approach the rich buff of adults. The rump of this specimen is

more dusky than the back.

In each of the eight skulls examined the anterior upper cheek-

MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS, 833

tooth has a distinct and often rather large fourth inner angle. As

is usual in microtine subspecies, the subspecific cranial characters

first become obvious in old age, although, if care be taken to

compare only skulls of equal age, close observation will find the

beginnings of such characters in younger stages too. The oldest

skull, the type (No. 102), is fairly comparable with that of No. 139,

a M.a. exsul from Mull, from which it is distinguished by its deeper

rostrum, lighter jugals, and more nearly vertical occiput; because

the occiput is less sloping the interparietal is a little less

reduced,—its posterior border is straight instead of sinuous,

and it is rather longer antero-posteriorly. The most striking

difference is seen, however, in the form of the ‘“ shield” or

flattened dorsal area delimited by the temporal ridges upon the

sides of the braincase. In ewsul (text-fig. 138 a) that part of the:

temporal muscle which arises between the interorbital region

and the glenoid articulation continues apparently throughout

life to extend its area of origin by creeping up the sides of the

braincase and so pushing the temporal ridge backward ; the result

is that in old age the anterior width of the “shield,” taken

between its antero-external angles, is considerably less than its

glenoid width: in the Islay mouse the upward creeping of the.

temporal muscle ceases at a relatively early stage of growth, and

the result is that in old age the ‘ shield” presents a form which

is found only in relatively early stages of development in exswl.

The oldest skull of ewswd from South Uist (Edinburgh, No. 300)

is younger than the skulls from Islay and Mull just described,

the temporal ridges not yet being quite fused in the interorbital

region, but the form of the “shield” is already more modified

than in the aged type of the present subspecies.

MICROTUS AGRESTIS NEGLECTUS Jenyns.

Three, Bute. : Dimensions :—

Head & Hind

body. Tail. foot. Lar.

INOmOomemalemanuve OPA pri onic Ne BO lly NB

Scena, 4, IP Wileigela By soc ICO "2G eh

6 a ea SeAsorilbh” io atey cs. PEL NG alates:

For cranial measurements see Table at p. 834.

The short tails are, no doubt, due to immaturity.

Making allowance for the youth of these specimens, we

ean detect no character by which the skins or skulls can be

differentiated from those of J/. agrestis neglectus of the mainland.

In his recently published ‘Catalogue of the Mammals of

Western Europe’ (1912), Mr. Miller treats the various agrestis-

like forms as subspecies. We think it preferable, somewhat

as proposed by Barrett-Hamilton in 1896 (P.Z.8. 1896, 19th May,

pp. 599-608), to regard these forms as belonging to two species,

834 MAJOR G. BE. H. BARRETL-HAMILTON AND

a larger northern with isolated southern colonies, now known to

be older in western Europe, a smaller southern, now known to be

a recent comer. Such an arrangement shows the relationships

most clearly. The earliest known member of the group in Europe

is a species of which the fossil and fragmentary remains have

been found in the early middle pleistocene brickearth of the

Thames at Grays in Essex; whether this form has any close

affinity with any living now it is impossible to say. In the

late pleistocene deposits of Britain, such as that fillmg the

Ightham Fissures, and in many of the caves, remains of a

large form not certainly distinguishable, with the available

materials, from J/. agrestis neglectus occur. The Skandinavian

agrestis, the Hebridean exsul and macgillivrai, the pleistocene

and recent British neglectus, together probably with the southern

French and Swiss levernedii, are all intimately connected forms

and may be regarded as subspecies of J/. agrestis, and of thera

macgillivrati may be counted as the most primitive. At some

time since the close of the pleistocene period J/. agrestis

neglectus has been succeeded in England and the lowlands of

Scotland by a smaller, brighter form, which being a newer

immigrant, may well be granted specific rank as J/. hirtus

Bellamy, with Jf. hirtus baillona de Sélys of France as a

subspecies.

Cranial measurements of Jicrotus agrestis.

es : = | [3

ey il) cel > | |=

& = a ee es | 3

= |wiBeall & Saplomeialveame \ 2.

heen ie Be et ehh SS | es sf SS

M. a. exsul. Ss fia a So 4 Sys

Murz. No. 139, male. | 286 | 162 | 34 | 195 | 71 | 84 | 81) 69

ARRAN. 43, juv. f. | 26:8 | 15:2 374 | 116 6:2 68 7-4 69

52, 5, m.| 27:0. | 15:8 316 | 125 6°4. 75 TA 70

JURA. 119, female | 27°3 | 16°5 vA | 12°5 6°7 WS || BPS 70

N. Uist. CY op | 26°) 14°6 33 | 116 | 64 Vs NW YB 68

S. Uist. R00, 4 |) Br3 | 168 | 33 || Tp, 68 79 | 78 65

BN sath |

M. a. macgillivran. | |

i | |

Istay. No. 89, male. 264 | 15°3 38 | 12:0 67 79 | 7:2 66

TO eel 26:5 | 16:3. |) 3:6 1 d1s8) |, G2, 16 726. Ives) eee

OS See al e2OrOn oA: 37 12°0 62 | 76 7-4, 68

87, female .| 27°4 | 16:0 37 | 127 1 OB 2) 7 68

102, Type .:.| 287 | 164 | 37 | 196 | 67 | 87 | 78 | 71

M. a. neglectus. | | | |

| |

Bure. No. 67,juv. m. | 243 | 140 36 | 115 | 60 | 68 69 61

Bh SUS paley | Gl Ge Ge) ay

MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS

APODEMUS SYLYATICUS SyLyATICUS Linneus.

Seventeen, Buts.

Head &

body. Tail.

IN@ Il, imeiley @ Wiesel OR Say ahs}

Ae dess gos eee ; VO SO

ead eo Loud ese Ms 90 80

10, 29 16 7) ? 90 (es

11D), dane rg 90 77

Deo Be WE 100"! 86

23, br) bh) 9) 9 87 76

21 Ah eT A aN NSE

3], 2? 23 2? 9? 95 75

5, female, 12 Be 80 78

Sia es ga etek aes 59 S879

9, ”? 14 ” ” 90 75

gg OT HD

A, gg AW Ne ; 90 90

22, ” ” ” ’ 83 67

DORR RUM ae , 80 80

3 tne 4 ett, 22) A) a 807-73

Average of 17 specimens of AEE

both sexes : sCachsterae wach | Ber Oe

Six, GREAT CUMBRAE.

No. 33, male, 27 March 1912. 95 92

SD) el Maer Deke aa OH OO

SOR POM: ae 5 95a Oe

37, ” ” ” ” 90 88

38, female, ,, 3 be 93 90

Ona: vO. os. 3 JO 90

Average of 6 specimens of | 93 90-3

A rather rufous series, perhaps due

worn.

both sexes .

Nine, ARRAN.

No. 44,

Average of 9 spec

towards the large foot and short ear of hebridensis.

male, 3

female,

April 1912.

ye)

both sexes

This series shows a rather large reddish mouse, with a tendency

>)%)

imens of

* Trapped on moss-covered bank.

t Trapped in middle of rough pasture.

Dimensions :—

100 «88

10S es

105-86

100. 85

LOO Wess

SORES

ea) to)

JOM RE Ts:

wa) fen)

98°3 84°59

SH SCN

bol po bh po by pp

WNMDNWPH bd w&

kD bD DO bO BO bo

Gs Oo CO to KD CO

Skull

Kar. Coaldy Ve

TLegeey

115)%15)

15 22D

Loui yy 2289

15 23°2

14 M95)

15 Sal

Dee ae 2)

14:5 21:5

14:7 22-2

S25) Die

16 Hil

15 22°8

14°5 20-4

14°5 21°8

16 21°6

14-76

5 23°4

14:5 22°8

15 Q32,

16 22°4

22°8 15:4

14:5

15:5

14:5 §

wo)

=r)

bo bo bo bo

02 He 02 19

OH Oo

+ Trapped in grass near sea-shore.

§ 'Frapped in heather on side of cliff.

Proc. Zoou. Soc.—1913, No. LV.

to late winter coat being

836 MAJOR G. BE. H. BARRETT-HAMILTON AND

One, GIGHA. Dimensions :—

Head &. Hind Skull

body. Tail. foot. Kar. c.eb: I

No. 126, female,22 May 1912. 100 85 22°5 15

A rather large nursing specimen with dimensions tending

towards those of hebridensis.

Ten, Istay. (8, Sheppard ; 2, Royal Scottish Museum.)

No. 97, male, 3 May 1912. ee 4b 283 13}

99, eee mY Gee Ms Vo ie ses Ns Ba)

93, female, 2 ,, 5 SSS Ome 2 melee 22:1

4 a. Re e LOO. sh 23 115) 24°]

10405, Ome ss 5S {3 Bo 23°7

GAs aes ger AKWe® JOM SO nee 22 onal 3

GBs 55 She ee) Go OO tom a oem

NGA eRUVGS: ye ae Co CLG 8

Royal Scottish Museum :—

INO, BOG HemNAes ocwanbesodsonsces OIL Baie BR. abo

SO Ot emEey PES Li Sia ee Pec 15 OO 2a 1G

Average of 9 adults of |

Inatilh SEEElosesyese: 93°3 81:6 22°5 14

A moderately large mouse, with back heavily washed with

black, probably indicating the new summer coat; stomach clear

white, and shght trace of yellow collar.

Three, JURA.

iINoswl22; male, 18!) May W912] 9884 24) a4. 5

123, 9 - HOOe 5. 722) EL oy 23-9

Les, Resanenlley 16 - a On me 1 en)

Average of 3 specimens aa :

of both sexes......... | HY GSO 2a) Mew

Five, Mutt.

Non 29) male, 13 June 19125 9a 922 2:oe It

1th, 115 Gy Oh toe) foi) 24 tlie ea

TSS ce tlO: Lt) OE NOOw) Ging e22nn gl eeu eS

Sie ” 20 ” 39 92 88 24 14:5

140, Ne Te ak KOO IO Eb NEG) Bdlom

Average oe WINES, so0ose Vi Sie 2322) ASA

Four, TIRE.

No. 146, male, 8 July 1912. 105 88 23°5/13°5

148, PO es, at as, 105 88 24 (13

149, a cla tag ee TOOW S73 ete guage

iLO), Fennel. Layeng t LOOM RS Sire tego

Aver age of + specimens of 102° 84:25 23:1 13:5

both SOMES manatee:

MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS. 837

Six from Tiree, in Royal Scottish Museum :—

Head & Hind

bodys Tail; foot. Har:

No. 344, male, 19 Nov. 1906. 80 85 23:5 14

Be), og, lB) IPGlo SOV, OB SR WS} i eyes

342, female, 19 Nov. 1906. S68 OO BF la

Oe ess |, Maat tse 4s SHA fe BB A

Ato). RR eet oe oA VS (ee Bil 1S

BAG Re ae Wee 5 S277) 2805 ono

Average of 6 specimens of |... : :

both sexes ........ esl Boe BO le

Average of 10 ara 92-9 84:5 21-1 13-8

The specimens from Tiree and Mull are, when adult, apparently

above the average size, and tend to have the large hind feet and

small ears of hebridensis. They also approach hebridensis in a

tendency to have the under side washed with yellow.

Cranial measurements of Apodemus sylvaticus sylvaticus.

We are unable to distinguish the specimens from the islands

from A. sylvaticus sylvaticus of the mainland. The dimensions

recorded above afford an idea of the variability encountered in this

form: even allowing for errors on the part of the collector, and

for the differing ages of individuals, there seems to be an inherent

tendency to vary. On all the islands, except Bute and Great

Cumbrae, there is a dimly seen tendency to acquire greater size,

larger hind feet, and shorter ears, 7. ¢.,a tendency to vary in

the same direction as hebridensis, as further shown by the speci-

mens from Tiree and Mull, in which the belly tends to be

washed with yellowish tints.

DD*

ener aR 7 |

| | os : a Ge rage

| 3 oS og ape 28

= 5 S| “ | rece 38

2 @ ices Py Bey ae | oe | aS

ee | eS eS) iy cS om | oo Be] se SS

Se S 25 aS) S : B Sa | 2a |

Bute, 21, male ...... 23:2 | 13:0 40 | 112 | 80 | 102 66 39 | 142 3°6 Aged. |

POO Napea Theat: 23°1 | 12:8 Lomeli? 80 | 95 67 40 | 143 38 jAduli.|

| Cumbrae, 38,female.| 23°2 | 13°2 38 | 11:8 82 | 92 | 68 39 | 145 36 | Aced. |

| Arran, 48, male......] 24:0 | 136 | 4:0 | 12-0 83 | 68 4°2 | 149 3°7 |Adult.|

| Islay, 94 female ...... 241 | 140 | 41 | 114 HS Mes ew a ZeO 37 | 149 37 | Aged. |

Jura, 123, male......| 23°9 | 13:0 |. 40 | 11:6 | 82 9°4. 7-0 4:0 | 144 3:8 |Adult.|

Wind, WO Gy earn 24°2 | 13:0 | A) | wale, | SL) OA PS 4-1 | 14:7 | 38 | Aged. |

838 MAJOR G. E. H. BARRETY-HAMILYTON AND

Mus muscuuus Linnzeus.

One, TIREE. Dimensions :—

Head & Hind

body. Tail. foot. Ear.

No. 147, male, 10 July 1912... 80 Co 18% 11?

An example in the yellowish outdoor pelage. It was trapped

in sand-hills.

In conclusion we have to sum up the distributional evidence

and offer a few remarks, of a quite tentative character, upon

its meaning as it appeals to us at present. Of the shrews,

S. minutus oceurs throughout the Outer and Inner Hebrides,

and in the Orkneys, Ireland, and other islands; S. araneus,

on the other hand, is confined to the mainland, the Inner

Hebrides and such islands as Anglesea and Wight; it is

represented in Islay by the peculiar S. grantii connected

morphologically and geographically with the parent form by

means of the partially differentiated Jura form. We may

therefore perhaps conclude that S. minutus was the first form to

arrive in this region; but this point is of little importance,

because both species have had representatives in Britain from an

extremely early period, and the problem before them in the past

may have simply resolved itself into a contest of endurance to

decide which of them could survive the rigours of the glacial

period in the outlying districts. Be this as it may, one fact

stands out clearly, that Islay has been separated from the

mainland longer than have the other islands of the Inner

Hebrides, with Jura approaching it in age. The microtine

species lead us to much more definite conclusions. We have in

the Inner Hebrides two genera, Microtus and Hvotomys ; the

former alone, so far as is known, inhabits the Outer Hebrides as

well. As is well known, neither genus occurs in Ireland, and

Microtus alone is met with in the Orkneys. There is a great

difference, however, between the Orcadian and the Hebridean

species of Microtus: the Orcadian forms are members of the

M. arvalis group, while those of the Hebrides are members of

the WM. agrestis group. Now, at first sight, having regard to the

fact that J. oreadensis and WW. sandayensis have a near ally in

the Guernsey J/. sarnius and another in the late pleistocene

English J. corneri, as well as to the fact that this group is no

longer represented on the mainland of Britain, one might feel

inclined to look upon the Orcadian fauna as an older one than

that of the Hebrides. On the other hand, we have to remember

that both the MW. agrestis group and the genus Hvotomys appeared

in Britain long before any of the other modern groups of Voles.

Each is represented in such an early horizon as that marked by

the Grays brickearth by a form much like the living ones; while

MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS. ——— 839

the J. arvalis group in a modern guise does not appear until the

late pleistocene era. Secondly, we have to face the remarkable

fact that both #. alstoni and M/. agrestis exsul, with its ally

macgillivrait from Islay, are most closely related to the Skandi-

navian species. Moreover, MW. agrestis macgillivrati appears to be

a little more primitive in its organization than ewsul; the latter

form in turn a little more primitive than true agrestis. The

interpretation of these peculiar facts which we suggest is that

the Hebrides have formed a refuge for some very old mammals ;

that from the former continuous land area, of which they are

now the disconnected remnants, certain of these old mammals

were dispersed along former land bridges to Scandinavia. The

proximity of the Atlantic would ensure the existence of much

milder conditions in the Hebridean area during the glacial

period, at whatever moment in pleistocene time that remarkable

event transpired, than would be present elsewhere to the east.

It will be seen therefore that we are practically in full accord

with the views put forward by Mr. L. Stejneger in his very able

and suggestive paper “ On the origin of the so-called Atlantic

Animals and Plants of Western Norway ” (Smithsonian Misc.

Coll. vol. xlviii. p. 458, 1907). Just as the presence of S. granti

implies that Islay was separated from the mainland at an earlier

date than were the other islands of the Outer Hebrides, so the

existence upon it of J/. agrestis macgillivrati and of a distinct

subspecies of Stoat, Mustela erminea ricine (found in Jura as

well), may be taken as evidence that it was detached from the

common Hebridean land surface at a relatively early moment.

The early detachment of Islay and its peculiar mammals has

an important bearing on the origin of the Irish Fauna. It

indicates that Islay and Ireland have not been connected since

the genus M/icrotus and Sorex araneus reached the Inner Hebrides.

And if that were so, the existence of the deep North Channel

between Ireland and South Scotland must at the same time have

interrupted communications in that direction also. We are

thus unable to accept E. R. Alston’s suggestion (Fauna of

Scotland, 1880, p. 5) that Ireland received its mammals from

Southern Scotland, and are forced to derive the Irish Fauna from

England and Wales (see Barrett-Hamilton, Clare Island Survey,

Mammalia, Proc. Roy. Irish Acad., March 1912),

EXPLANATION OF PLATE LXXXIV.

Fig. 1. Sorex grantii. Natural size.

2. Sorex araneus castaneus. Natural size.

840 MR. T. H. WITHERS ON

47, Some Miocene Cirripedes of the Genera Hexelasma

and. Scalpellum from New Zealand. By Tnomas H.

Wiruers, F.G.8.*

[Received May 6, 1913: Read June 3, 1913. }

(Plates LXXXV. & LXXXVI.7 & Text-figures 139, 140.)

INDEX.

Page

Introduction ........ AR La cde Ae SOCK. ct OM0 Leet ner nA)

Hexelasma Ron iandicuit (Hector VOM b ardaeeE cadaunaeeeeneradcumely tor all

Structure and Affinities of H. aucklandicum .................. 846

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Distributions (Geological) yeas) eeeeseheaseeaes B41, 848, 851

This paper contains the results of a study of the remains of

the ‘gigantic Cirripede” (= Hexelasma aucklandicum) from

New Zealand, as well as some notes on a smaller species of

Hexelasma, and descriptions of two new species of Scalpellum.

One of the latter is founded on some valves in the Geological

Department of the British Museum, and the remaining species

of Scalpellum and the small Hexelasma were found associated in

the matrix with the remains of Heaelasma aucklandicum.

Remains of a gigantic Cirripede have long been known to

oceur in the Waitemata Beds (Miocene) of Motutapu Island,

Auckland Harbour, New Zealand. These remains have been

considered by Sir James Hector (1887) and Prof. W. Blaxland

Benham (1903) as belonging to a pedunculate Cirripede; but

while the former referred them to the genus Scalpellum, the

latter thought that they approached more closely to the genus

Pollicipes.

On learning of my wish to see some of these remains,

Prof. James Park was good enough to write to Dr. J. Allan

Thomson, Paleontologist to the Geological Survey, Dominion

Museum, Wellington, who most kindly sent me the actual

specimens collected by Prof. Park in 1887. Prof. Park wrote

also to Prof. Benham, who sent me plaster-casts of the specimens

figured by him in 1903; these casts are now in the Geological

Department of the British Museum. My thanks are therefore

due to Professors Benham and Park and Dr. J. Allan Thomson,

and I have also to acknowledge the kindness of Dr. A. Smith

Woodward in allowing me to describe the new species of

Scalpellum in the Geological Department of the British Museum.

* Communicated by Dr. W. T. Catman, F.Z.S.

+ For explanation of the Plates, see p. 854. j

WARS MNS) ENE ILYOCQVi

Huth imp.

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AH. Searle

HEXELASMA AUCKLANDICUM, Hector sp.

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+6.SCALPELLUM SUBPLANUM. sp.n:

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TI8.SARCOSG ALPELLUM) UNGULATUM. sp.n.

Fis

FOSSTL CIRRIPEDES FROM NEW ZEALAND. 84]

BALANIDA.

Genus HEXELASMA.

1913. Hewelasma P, P. C. Hoek, Siboga-Expeditie, Cirripedia

Sessilia, p. 244.

“Compartments six; carina, carino-lateral, and lateral compart-

ments with ale, but without radii, the rostrum having neither

radii nor ale. Parietes not porose and without longitudinal

ribs on their inner surfaces; basis membranous. Opercular

Vvalives sulo-uanoular. eee by ky 0. bloel:

HEXELASMA AUCKLANDICUM Hector sp. (Pl. LXXXYV.)

1888. Scalpellum aucklandicum Uector, Trans. N.Z. Institute,

vol. xx. (1887) p. 440.

1903. Pollicipes ? aucklandicus Hector sp.: W. B. Benham, “ On

some Remains of a Gigantic Fossil Cirripede from the

Tertiary Rocks of New Zealand,” Geol. Mag. London,

dec. 4, vol. x. p. 111, pls. 9, 10 (non figs. 8, 9).

1905. Pollicipes aucklandicus Benham: E. Clarke, ‘‘'The Fossils

of the Waitemata and Papakura Series,” Trans. N.Z.

Institute, vol. xxxvi. (1904) p. 419.

1910. Pollicipes ? aucklandicus Hector sp.: J. Park, ‘ Geology

of New Zealand,” p. 115 (pl. 7), pp. 1138, 134.

Diagnosis. Compartments attaining a length of at least 187 mm.,

carinal, carino-lateral, and lateral compartments with simple ale

(i.e., there is no distinct upturned extension at the margin).

Sheath feebly developed, almost absent, and with no suturai edge

to abut against the longitudinal ridge formed on the inner

surface. Opercular valves unknown (except for probably a single

tergum).

Material. 7 rostral compartments, 9 carinal, 13 lateral (8 right

and 5 left), 5 carino-lateral (3 right and 2 left), together with a

small tergum (Pl. LXXXYV. figs. 13 a, 6), which may or may not

belong to the species; all these are more or less imperfect. The

specimens are in the collection of the Geol. Surv. New Zealand,

and are marked with the locality-number 695. They are pre-

sumably the syntypes of Hector. In addition to these specimens

I have examined plaster-casts of the specimens figured by

Prof. Benham.

Holotype. From among the syntypes of Hector I select as

holotype the rostrum here figured on Pl. LXXXY. fig. 1.

Horizon and Locality. Miocene, Oamaruian, Base of Waitemata

Beds: Motutapu Island, Auckland Harbour, New Zealand.

General Remarks. Sir James Hector (1887) first called at-

tention to this fossil, and at a meeting of the Wellington

Philosophical Institute he remarked on some remains of it there

exhibited :—

“Specimens of a large fossil stalked Cirripede, recently

842, MR. T. H. WITHERS ON

collected by Mr. Park, at Motatapu Island, Auckland. A careful

restoration will have to be made before definitely determining

this fossil, but it will probably be found to belong to the genus.

Scalpellum and is distinguished provisionally under the name

S. aucklandicum. In size, this fossil Cirripede greatly exceeds

any previously known, in 8. magnum the capitulum being only

13 inches in length, while in the Auckland specimen it is at

least 8 inches. These fossils oceur in a breccia, marking the old

shore line of the upper part of the W aitemata Series, similar

to the Cape Rodney beds. The associated fossils are Corals,

Brachiopods, and Hchinoderms. Among the latter are two

specimens having plates of a Cidaris of enormous size.”

Attention was again called to this fossil in 1903, when

Prot. Benham described and figured certain valves. He con-

sidered that they showed closer resemblance to the capitular

valves of certain species of Pollicipes, and doubtfully referred

them to that genus.

After an examination of the present material I am convinced

that the valves belong to a sessile Cirripede allied to Balanus.

There are six compartments—a rostrum, carina, right and left

lateral, and right and left carino-lateral. Prof. Benham figured

only four valves, namely, “a carina, left scutum, ?rostrum, and

upper latus.” The carina figured by him is the same as that

now considered as a carina, and the scutum and ? upper latus.

correspond to the right and left lateral compartments respec-

tively; but the valve figured (Benham, 1903, pl. 10, figs. 8, 9);

as a “?yrostrum” is really a carina of Scalpellum subplanwm,

sp. n. (see p. 848). The valves considered here as rostral and

right and left carino-lateral compartments were not figured

by Benham, and it has now been possible to give figures of the

inner surface of each different compartment.

Description of Valves. Valves with solid walls of variable

thickness, apparently not more than 2°25 mm.; externally

marked with prominent, more or less regular, transverse growth-

ridges, rather more strongly marked on the carina; sometimes

ridged longitudinally, and in one or two cases the valves are

distorted by linear depressions; but all the valves are more

or less irregular in shape, and this is obviously caused by the

surface of attachment; inner surface not longitudinally ribbed

near the base as in Balanws, for the smooth imner surface slopes

gradually to meet the outer surface and forms a definite, more

or less smooth edge.

Rostral compartment (PI. LXXXV. figs. 1-3) without radi,

almost symmetrical, moderately convex transversely and slightly

convex longitudinally, bluntly angular at the apex, and either

rounded or slightly concave at the basal margin; triangular in

shape when young, but in the older and consequently longer

valves the lateral margins for the greater part are almost

parallel to each other. On the inner surface two more or jess

prominent ridges extend from the apex, and die out at a point.

halfway from the apex in the young valve (fig. 2) and at a

POSSIL CIRRIPEDES FROM NEW ZEALAND. 843

Text-fig. 139.

OUTER VIEWS.

Text-fig. 140.

INNER VIEWS.

Hexelasma aucklandicum Hector sp.

Restored compartments.

r., rostrum ; 7., lateral; ¢./., carino-lateral; c., carina.

844 MR. T. H. WITHERS ON

point about one-third the length of the valve from the broken

apical portions in the larger valves (figs. 1, 3). Obviously these

ridges serve for the reception of the angle of the ale on the

adjacent lateral valves, and the space between the ridges is

marked, in the older specimens, with transverse lines. The

portion of the valve enclosed by the ridges is half as wide as the

adjacent parts of the valve in the specimen figured (fig. 3 6), but

in the valves figured (figs. 1 6, 26) it is wider. At the base of

the two ridges extending from the apex, and only in the largest

valve (fig. 16), a slight transverse ridge is formed by the

thickening of that part of the valve enclosed by the ridges.

From a point about one-third from the base of the valve, a small

extent of the inner surface, parallel to the outer margins, is

marked with lines which extend upward, and bend abruptly

inwards towards the base of the ridge on either side ; these lines

are made by the ale of the lateral compartments which are

overlapped by this part of the valve.

Lateral compartment (figs. 4-6) with an ala on the rostral

side, moderately convex transversely, and longitudinally almost

flat, irregularly convex, and in one case bent in an elongately

S-shaped curve; the whole valve is usually strongly bent

towards the rostrum, but one valve is strongly bent away from

the rostrum. Parietal portion very much wider than in the

carino-lateral compartment, as much as three times as wide as

the widest part of the ala in one valve, and in others from two

to under one and a half times as wide. The two margins of the

ala form an obtuse angle, the upper margin is practically straight

and the lower somewhat concave, but their shape is influenced by

the degree of curvature of the valve; the growth-lines on the ala

are closely set and extend obliquely upwards from the base, and

on reaching almost to the upper margin curve downwards to the

angle of the ala; on the lower margin, near the angle, a small

smooth portion is left just below where the growth-lines bend

downwards. On the inner surface, almost at the middle of the

parietal portion, a more or less prominent ridge extends from

the apex and dies out at a point opposite the angle of the ala;

near the parietal margin the inner surface is marked with lines

which extend upwards, and on reaching a point just above the

base of the longitudinal ridge bend sharply inwards and down-

wards to meet its lower extremity; these lines are obviously

made by the ala of the carino-laterai compartment, the angle of

which abuts against the longitudinal ridge. Between the longi-

tudinal ridge and the upper margin of the ala the inner surface

is marked with indistinct and irregular transverse lines.

Carino-lateral compartment (figs. 7-9) obtusely triangular in

general outline, with an ala on the rostral side. The whole valve

bent, especially in its apical half, towards the rostrum, is almost

flat transversely, and the parietal portion is much narrower than

that of the lateral compartment; the two margins of the ala

FOSSIL CIRRIPEDES FROM NEW ZEALAND. 845

form an obtuse angle, the upper margin of which is straight and

obliquely inclined towards the apex, and the lower margin, which

is concave, emerges from just above the base of the valve, and

curves gently upwards and then sweeps sharply outwards to meet

the upper margin. Parietal portion of valve extremely narrow,

the widest part being about one-fourth the width of the widest

part of the ala. On the ala the growth-lines are closely set, and

follow a similar course to those on the lateral compartment.

Beneath the upper half of the ala, a portion of the valve on the

inner surface is somewhat thickened for about one-third the

width of the valve; its inner margin is steep-sided, and forms a

ridge, against which abuts the angle of the ale of the carinal

department; the thickened portion of the valve widens gradually

from the apex, dies out at a point opposite the angle of the ala,

and is marked with fine, regular, closely-set, transverse lines.

Near and parallel to the lower half of the parietal margin, and

for about one-third the width of the valve, the inner surface is

marked with lines which extend upwards, and, on reaching about

half the length of the valve, ave angularly bent downwards and

inwards, and the lowest of them meet the base of the steep-sided

ridge near the upper margin of the ala; these lines correspond

to, and are obviously made by, the ale of the carina, which are

overlapped by the portion of the valve thus marked.

Carinal compartment (figs. 10-12) gently curved longitudinally,

with an ala on each side, and these are bent at a sharp angle

from the parietal portion. Parietal portion narrow, transversely

convex, especially near the apex, much narrower than the parietal

portion of the lateral compartment, but wider than that of the

carino-lateral compartment ; ale about one and a half times as

wide as the widest part of the parietal portion, and in one young

valve about the same width as the parietal portion. The ale

emerge from near the base of the valve, widen gradually upward

until about two-thirds the distance from the base, and here they

bend further outwards and then sharply upwards to the apex ;

the two margins, therefore, roughly form an obtuse angle, the

basal margin of which is somewhat concave, and the upper

margin, which is the shorter, is straight. The growth-lines on

the alee extend obliquely upwards from the base and, on reaching

a point more than halfway across the ale, bend sharply and

angularly downwards to the margin; a smooth triangular portion

of the valve is left beneath the angularly bent growth-lines.

The inner surface is quite smooth except for some transverse

lines, which mark the surface above the angle of the ale and

which are more.prominent at this point.

Measurements. The largest valve in the present series (a carina,

fig. 10) would measure, if complete, circa 90 mm., and the smallest

valve (a lateral compartment, fig. 6) cirea 12mm. One of the

compartments (a lateral) figured by Prof. Benham as a scutum

measures 187 mm.

MR. T. H. WITHERS ON

Structure and Affinities. It is evident from the structure of

the inner surface of the compartments of this Cirripede, and

from the modification of the side wall of the compartments to

form ale, that the elements combined to form a shell something

like the compartments in the genus Salanus. This is proved by

the fact that the lines on the inner lateral portions of the rostrum

correspond to, and are obviously made by, the growth-lines of

the ala of each lateral compartment, which was overlapped by the

lateral portions of the rostrum; similarly shaped lines on the

lateral compartment correspond to those on the ala of each

carino-lateral compartment which was overlapped by the lateral

compartment; the more angularly bent lines on the carimo-

lateral compartments correspond to those on the ale of the

carinal compartment which were overlapped by the carino-lateral

compartment. In addition, more or less prominent longitudinal

ridges are developed in the upper portions of the rostral, lateral,

and carino-lateral compartments against which the angle of the

ala of the adjacent valves abutted. The transverse lines on

the portions of the compartments not covered by the ale are

similar to those in Balanws, in which they are caused by the

successive exuviation of the opercular membrane. To a similar

cause may be attributed those in the present Cirripede.

That this form is a sessile Cirripede is, | think, beyond doubt,

and the irregularity in shape of the several compartments, as

well as the fact that some are externally and irregularly ridged

longitudinally and others distorted with linear depressions, 1s.

further evidence in support of this conclusion, for these features

could be caused only by the irregularity in the surface of

attachment. In accordance with the above interpretation, six

compartments would complete the wall of the shell, as in

Balanus; and it is important to note that every fragment in

the present collection can be allocated to one of the six valves

figured. (Pl. LXXXV. & Text-figs. 139, 140.)

The shell of this form could not have been very strong, for, in

comparison with the great length attained by the compartments,

the walls aré relatively quite thin. The compartments could

have been only weakly attached, and are in consequence always

found quite apart from one another. Moreover, the absence of

radii, together with the absence of sutural edges to the alee and

the comparatively feeble ridges or shoulders developed on the

inner surface, and against which only the angles of the ale could

have abutted, show « quite clearly the great ‘structural frailty of

the shell. It could hardly have been a littoral barnacle.

Although this form agrees with the typical species of the genus

Balanus in the number of compartments forming the walls of the

shell, it differs markedly in the structure of these compartments.

These differences are (1) the absence of radii, (2) the simpler

structure of the ale, (3) the absence of longitudinal ribs on the

inner surface, (4) the feebly developed sheath, as well as the

FOSSIL CIRRIPEDES FROM NEW ZEALAND. 847

absence of a sutural edge to abut against the longitudinal ridge

on the inner surface. All these are primitive characters, and

show that in this Cirripede we have a Balanid more primitive

than Balanus.

In my preliminary consideration of this form I regarded it as

being related, mainly in the absence of radii, to the recent species

Bataan Honaunms and BL. corolliformis, winich were included by

Dr. Hoek* in a new section (G) of Balanus, and B. hoekianus

and B. callistoderma, which were referred by Dr. Pilsbry f to the

same section, and I intended to found a new genus to include

these species. On seeing a proof of Dr. Hoek’s work (1913,

‘ Siboga-Expeditie, Cirripedia-Sessilia,’ pp. 244-246), however,

I found that he had included these recent species, together with

two new species (/7. velutinwm and H. arafurae), in a new genus

Hexelasma.

Dr. Hoek kindly sent me drawings of the type-species HZ. velu-

tinwm, and from these it could be seen that while Hewxelasma

differs from Balanws in the absence of radii and the absence of

longitudinal ribs on the mmner surface, it agrees in having a well-

developed sheath, and in the carino-lateral compartments having

an upturned extension of the ale as well as a well-developed

sutural edge.

Since ‘ Pollicipes (¢) aucklandicus” agrees with Hexelasma in

the absence of radii and of longitudinal ribs on the inner surface,

it seems advisable to refer it to that genus ; but in some respects

it appears to be somewhat more pr imitive than the typical species

of Hexelasma, especially in the feeble development of the sheath,

in the absence of a distinct upward extension to the ale of the

earino-lateral compartments, and of a sutural edge to the ale of

the carino-lateral compartments.

The species included in Hewelasma are all deep-sea forms, and

occur at depths varying from about 100 to 900m. In length

the shell of the largest species, HY. corolliforme Hoek, measures

nearly 45 mm., and since the largest-known compartment of the

fossil H. aucklandicum measures about 190 mm., the great

difference in size is apparent.

Except for Balanus psittacus Molina sp., which has been known

to attain a length of 9 inches (circa 225 mm.), Heaxelasma auck-

landicum 1s the largest-known Cirripede. Salanus evermanni

Pilsbry, another large barnacle, is recorded as measuring 150 mm.

HEXELASMA sp.

A large number of the disconnected compartments of a small

Balanid are to be seen scattered about in the matrix containing

* 1883. P. P. C. Hoek, ‘Challenger’ Report, Zoology, vol. viii. pp. 155-160.

a Ugh, Jets Jeiilsjony, Barnacles of Japan and “Bering Sea,’ Bull. Bureau

Fisheries, Washington, vol. xxix. 1909, pp. 76-80.

3848 MR. '. H. WITHERS ON

the compartments of Heaxelasma aucklandicum and the valves of

Scalpellum subplanwm, sp. nu. Those that I have been able to

extract and clean appear to be somewhat worn, and the largest:

of them does not measure more than 5 ov 6 mm. in length; their

outer walls are thrown into comparatively wide longitudinal

folds. Owing to their worn appearance the finer characters are

not well shown, but since they do not appear to possess radit

they must be referred to the genus Hewelasma. They differ,

however, from the typical species of Hewelasma, as well as from

the fossil H. wucklandicum, in having a well-developed sheath,

and in the presence of strong ribs on their inner surface. Balanus :

hoekianus Pilsbry (1911, ‘ Barnacles of Japan and Bering Sea,

Bull. Bureau TDi Werate, Washington, vol. xxix. 1909, p. 2

text-fig. 8), which has now been referred by Dr. Hoek to his

genus Hexelasma, agrees with the present compartments in the

presence of ribs on the inner surface, but it would be rash to

say that they are related specifically. In view of the unsatis-

factory preservation of these compartments, and in the absence

of the opercular valves, I do not think it advisable to institute a.

new species.

Horizon and Locality. Miocene, Oamaruian, Base of Waitemata

Beds; Motutapu Island, Auckland Harbour, New Zealand.

Collection. Geol. Surv. New Zealand.

POLLICIPEDID4.

Genus SCALPELLUM.

1817. Scalpellum Leach, Journ. de Physique, &e., Ixxxv. p. 68.

SCALPELLUM SUBPLANUM, sp.n. (PI. LXXXVI. figs. 1-6.)

1903. Pollicipes (?) aucklandieus Hector sp.: W. B. Benham,

Geol. Mag. dec. 4, vol. x. p. 114, pl. 10, figs. 8-9.

Diagnosis. Carina not separated into tectum, parietes, or

intraparietes, flatly arched transversely, basal margin bluntly

angular; tergum with the upper carinal margin unusually short,

and making with the occludent margin an obtuse angle ; rostrum

with a wide, flat median keel extending from the apex to the

basal margin.

Material. 2 carine, 2 scuta, 4 terga, | rostrum, and 1 sub-

carina; most of these are incomplete, and they were all extracted

from the matrix containing the valves of Hexelasma aucklandicum.

Holotype. The carina figured on Pl, LXXXVI. fig. 2.

Collection. Geol. Surv. New Zealand.

Horizon and Locality. Miocene, Oamaruian, Base of Waitemata

Beds: Motutapu Island, Auckland Harbour, New Zealand.

Carina not separated into tectum, parietes, and intraparietes,

bowed moderately either inwards or outwards, widening gradually

FOSSIL CIRRIPFEDES FROM NEW ZEALAND. 849:

from the apex to the basal margin, which is more or less bluntly

angular; flatly arched transversely and indistinctly carinate.

The apical portion of the smaller specimen is thickened to quite

a third of its extent, the inner portion forming almost a flat

surface extending from each side of the carina.

Length (fig. 1, basal half of valve) 16-5 mm. ; breadth 10 mm.

Length (fig. 2, basal 2 of valve) 16°6 mm.; breadth 6-4 mm.

Scutum trapezoidal, nearly twice as long as wide, divided

almost equally by an indistinct, flat, wide ridge extending from

the apex to the basi-lateral angle. Oceludent margin convex ;

basal margin almost straight. “Tere ral margin slightly coneave,

of about the same length as the convex lateral margin, with

which it forms an obtuse angle. Along the tergal margin the

valve is rounded towards the inner surface. The inner occludent

edge is broad, flat, of about the same width throughout, marked

with growth-lines, and overhangs the depression for the adductor

Eoutor um opposite the tergo-laterai angle; the inwardly rounded

tergal edge is marked with growth- Hines, but the extent thus

marked narrows rapidly towar -ds the ter go-lateral angle.

Length (fig. 3, incomplete valve) 28: 7 mm.; breadth 17 mm.

Tergum subrhomboidal, with an obscure ridge extending in a

straight line from the apex to the basal angle, and dividing the

valve unequally, the occludent portion being in its widest part

almost twice as wide as that of the carinal portion ; lower carinal

margin weakly convex, somewhat longer than the scutal margin,

and forming with it an acute angle; upper carinal margin

slightly convex, unusually short, and forming an obtuse angle

with the slightly convex occludent margin, which is almost twice

its length. The occludent margin forms a somewhat raised

border, and this is followed by a wide, shallow depression

bounded by an obscure ridge extending from the apex to a point

on the scutal margin, which is slightly produced about one-third

the distance from the basal angle; from this ridge the valve

slopes upwards towards the apico-basal ridge, and slopes rapidly

down to the carinal margin. On the inner surface a narrow

portion of the valve along the occludent and upper carinal

margins is marked with growth-lines, and the extent thus marked

is wider beneath the apex.

Length (fig. 4) 20 mm. ; breadth 12:4 mm.

Rostrum subtrianglar, strongly convex transversely, bowed

inwards, basal margin convex ; lateral margins slightly concave ;

a flat submedian keel extends from the apex to the basal mar Sn,

and this on the right side is followed by a further longitudinal

ridge near the lateral margin, but on the wider left side this

ridge, if present, is extremely obseure.

Length (fig. 5) 10°5 mm. ; breadth (when complete) circa 8 mm.

Subearina triangular, moderately convex transversely, bowed

inwards; apex rounded ; basal margin concave ; lateral margins.

slightly concave. On the inner surface a slight, but well-defined

850 MR. T. H. WITHERS ON

ridge extends from the apex to about the middle of the valve,

and is there met by two further ridges extending from each basal

angle; the valve is thus divided into three almost equal portions,

the basal one being smooth and doubtless covered at one time by

the corium, the two upper portions being marked with growth-

lines and most probably overlapping the adjoining carino-lateral

valve on either side.

Length (fig. 6) 5 mm.; breadth 5°2 mm.

Comparison with other Species. This species is referred to the

genus Scalpellum (sensu lato), a course that is advisable until we

can find the remaining valves of the capitulum, which may enable

us to refer the species to one of the subgenera into which

Scalpellum has been divided. Judging from the known valves

(carina, scutum, tergum, rostrum, and subcarina), this species is

related to S. zancleanwm Seguenza* from the Pliocene of Messina,

Sicily. The carina of S. zancleanum differs in having a strong

median keel, from which the sides of the valve slope steeply, and

in the less angular growth-lines; the scuta differs in the rounded

basal. margin and in the usually less acute tergo-lateral angle ;

the terga differ in being much narrower in proportion to width,

in the much more acute apical portion, and in the carinal margin

not being divided into an upper and a lower portion; the rostrum

differs in the absence of a wide, flat, median keel.

Subgenus ARCOSCALPELLUM.

1907 (Oct.). Arcoscalpellum P. P. C. Hoek, Siboga-Expeditie,

Cirripedia Pedunculata, p. 59.

1907 (Nov.). Holoscalpellum H. A. Pilsbry, Bull. No. 60, U.S.

Nat. Mus. p. 25.

1908. Arcoscalpellum Hoek; H. A. Pilsbry, Proc. Acad. Nat.

Sci. Philadelphia, p. 109.

1912. Arcoscalpellum Hoek; T. H. Withers, Proc. Zool. Soe.

London, p. 538.

SCALPELLUM (ARCOSCALPELLUM) UNGULATUM, Sp. n. (Pl, LXXXVI.

figs. 7-13.)

Diagnosis. Carina with its tectum almost flat, parietes more

than half the width of the tectum, intraparietes narrow and bent

abruptly inwards, basal margin rounded; upper latera sub-

triangular, with rounded basal margin and wide lateral portions

obliquely inclined towards the umbo; rostral latus with sub-

parallel scutal and basal margins, and about one-third of its

apical end, which is much thickened, projecting freely beyond the

scuta,

* 1876. G. Seguenza, Atti Accad. Pontaniana, vol. x, p. 386, pl. vii, figs. 1-13.

FOSSIL CIRRIPEDES FROM NEW ZEALAND. B51

Material. 6 carine, 4 scuta, 2 terga, 1 upper latus, and 1 rostral

latus.

Holotype. The carina, I. 15409 (fig. 8).

Collection. Brit. Mus. (Nat. Hist.).

Horizon and Locality. Miocene, Upper Oamaruian. Blue

Clay: Pareora, South Canterbury, New Zealand. Jimestone :

Takiroa, Oamaru District, N. Otago, New Zealand.

In the Geological Department of the British Museum there is

a series of 14 valves of a species of Sealpellwm (sensu lato),

registered I. 15409-I. 15422. Although these come from different

localities, they are of interest, since they appear to represent a

single new species and enable us to gain some idea of the form

of the capitulum (see restoration, fig. 13).

Three of the valves (a carina, scutum, and rostral latus) are

labelled “ Scalpellam | (Blue Clay) | Parimoa, New Zealand | Mr.

C. Mantell.” The remaining eleven valves (5 carine, 3 scuta,

2 terga, and 1 upper latus) are labelled “Scalpellwm | Takiroa, New

Zealand | Mr. C. Mantell.” With the latter series is a MS. label

(probably written by the collector) bearing the words “‘ Lime-

stone | Takiroa—Crinoline Qu. 1852.”

It is fairly obvious that the “ Mr. C. Mantell” was intended

for the late Hon. Walter Mantell, from whom a collection of

New Zealand fossils was obtained- by the British Museum in

1856. On failing to trace the above-mentioned localities, I wrote

to Prof. James Park, who kindly informed me that ‘‘ Parimoa ”

should be Pareora, South Canterbury, and Takiroa is in the

Oamaru District, N. Otago; he was of the opinion that the

Cirripede valves came from beds of Miocene (Upper Oamaruian)

age. ‘Crinoline Qu.” may mean Crinoline Quarry.

Description of Valves. Valves thick, ornamented exteriorly

with numerous exceedingly fine ridges radiating from their

apices.

Carina narrow, widening very gradually from the apex,

moderately bowed inwards, basal margin rounded. ‘Tectum

almost flat transversely, not carinate. Parietes more than half

the width of the tectum (in one old specimen nearly as wide),

bent nearly at right angles to the tectum. Intraparietes very

narrow, bent abruptly inwards and meeting just below the apex,

so that a small portion of the valve projected freely. Umbo

apical.

Length (fig. 7, incomplete valve) 19:2 mm. ; breadth 6°5 mm.

Length (fig. 8, small complete valve) 16 mm.; breadth 3°7 mm.

Scutum trapezoidal, almost twice as long as wide; basal and

lateral margins slightly concave, nearly at right angles to each

other; tergo-lateral angle reaching a point a little above the

middle of the valve; tergal margin straight and its outer edge

flat ; tergo-lateral portion of valve slightly concave; occludent

margin moderately convex and very obliquely inclined towards

the terga. Umbo apical.

Length (fig. 11) 27-4 mm.; breadth 15 mm.

Proc. Zoou. Soc.—1913, No. LVI. 56

852 MR. IT. H. WITHERS ON

Tergum subtriangular, convex transversely, thick, elongated,

with a prominent, nearly straight ridge, extending from the apex

to the basal angle. On both sides of this ridge the valve slopes

rapidly to the outer margins. Carinal margin gently convex ;

occludent margin gently convex, with a raised border, followed

by a wide depr ession, extending ‘from the apex to about half the

width of the scutal margin, whichis “ncleniter! to that extent.

Length (fig. 9, incomplete valve) 21°4 mm.; breadth 10°8 mm.

Upper latus subtri angular, thick, convex transversely and

longitudinally ; umbo a little below the apex, owing to a thick

ledge formed beneath it, which extends from the lateral angles to

just beyond the umbo. Scutal margin slightly convex, its upper

surface forming a prominent ridge, followed by a wide depression,

which is bounded by an indistinct ridge, and between these two

ridges the growth-lines are abr uptly upturned; a second in-

distinct ridge runs almost parallel to the slightly convex tergal

margin, and between these the growth-lines are also abruptly

upturned.

Length (fig. 10) 15-4 mm.; breadth 13°5 mm.

Rostral latus about four times as wide as long, with a promi-

nent, rounded, wide ridge extending its whole length, from which

the surface of the valve slopes rapidly ; outer (rostral) extremity

bluntly angular, much thickened, and this must have projected

freely more than one-third the extent of the valve; inner

(lateral) extremity obliquely truncated. Scutal margin concave,

sloping upwards to the inner margin of the thickened portion,

and then descending rapidly in a curve to the umbo; basal

margin straight and almost parallel to the inner two-thirds of

the scutal margin,

Length (fig. 12) 4-4 mm.; breadth 15:1 mm.

Remarks and Comparison with other Species. In a paper by

Dr. G. A. Mantell* (1850, p. 329) a Cirripede from the Ototara

Limestone (Miocene, Upper Oamaruian) is recorded as ‘“ Pollicipes,

resembling a Cretaceous species.” There is nothing to indicate

whether this is the present species or not. ‘The two scuta

figured by Zittel T as “ Cirrhipedenschalen,” and recorded from

Whaingora and Aotea, North Island, New Zealand, are certainly

distinct from the present species. Zittel’s figures were sub-

sequently reproduced and recorded as Scalpellum sp. by Prof.

James Park, who considered the specimens to be of Miocene

(Upper Oamaruian) age.

* 1850. G. A. Mantell, ‘* Notice of the Remains of the Dinornis and other Birds

and of Fossils and Rock-Specimens, recently collected by Mr. Walter Mantell in

the Middle Island of New Zealand; with Additional Notes on the Northern

Island. WithsNote on Fossiliferous Deposits in the Middle Island of New Zealand,

by Prof. E. Forbes.” Quart. Journ. Geol. Soc. London, vol. vi. pp. 319-343,

pls. xxvill., xxix.

+ 1865. K. A. Zittel, “ Fossile Mollusken und Echinodermen aus Weuscelandig

Palaontolozgie von Neuseeland (Novara-Exped.), Geol. Teil., Bd. 1. Abth. 2, pl. ix.

figs. 12 a, b.

£1910. J. Park, ‘ Geology of New Zealand, p. 141 (pl. xu1.), figs. 9a, 6.

FOSSI“L CIRRIPEDES FROM NEW ZEALAND. 853

From a study of the isolated valves here described, it seems

reasonable to suppose that they formed a capitulum just as is

shown in the restoration (fig. 13). No examples of the carinal

and inframedian latera have yet been found, but doubtless these

will prove to be somewhat like the valves diagrammatically

indicated in the restoration. The capitulum, as_ restored,

possesses 13 valves, but this number may have been increased

to either 14 or 15 by the addition of a rostrum or subcarina, or

both. In any case, it is apparent from the structure and dis-

position of the valves, that the species falls into the group now

included in the subgenus Arcoscalpellum Hoek.

Scalpellum (A.) ungulatwm may be compared with S. michelot-

tianum Seguenza*, from the Pliocene of Messina, Sicily, and

S. quadratum Dixon t sp., from the Eocene of Bognor, Sussex,

but both these species differ in the valves being appreciably

thinner.

Although size is often of little account, none of the numerous

valves of S. quadratum that I have seen are more than half the

size of the largest valves of S.(A.) wnguwlatum. The carina of

S. quadratum difters in having inwardly bent intraparietes, the

scutum in having a raised border to the tergal margin, the tergum

in the carinal margin being elongately §-shaped and the apical

half bent towards the carina, the upper latus in being more

symmetrical and proportionally wider, and the rostral latus

differs in the absence of the prominent, transverse, rounded

ridge or fold and in the outer extremity of the valve being

thickened only to a small extent.

In S. michelottianum the carina differs in having broad,

rounded, longitudinal ridges on the outer margins of the tectum,

in the tectum being proportionally much wider and the basal

margin less convex, the scutum in having the tergo-lateral

portion of the valve markedly convex and rounded inwards at

the tergal margin, the tergum in being much thinner and very

much flatter transversely and the median longitudinal ridge

hardly perceptible, the upper latus in being more triangular in

shape and in the absence of a thick ledge formed beneath and

beyond the umbo, the rostral latus in having a much smaller

extent of the valve thickened at the outer extremity and in the

inner extremity not being obliquely truncated.

S. nuchelottianwm var. gassinensis Alessandyvi ~ (Pliocene to

Miocene) agrees more closely: with S. (A.) wngulatum in having

the basal margin of the carina more strongly convex, but the

other characters of this and the remaining valves differ quite as

much as the valves of S. michelottianum Seguenza.

* 1876. G. Seguenza, Atti Accad. Pontaniana, vol. x, p. 381, pl. vi. figs. 15-25 ;

p. 464; pl. x. fig. 26.

+ 1846. F. Dixon, in J. de C. Sowerby, Min. Conch. vol. vii. pl. 648; 1851.

C. Darwin, Pal. Soc. Monogr. Foss. Lepadide, p. 22, pl. i. fig. 3.

£ 1906. G. de Alessandri, Palwontogr. Ital. vol. xii. p. 252, pl. xiii. figs. 10-15,

854 ON FOSSIL CIRRIPEDES FROM NEW ZEALAND.

EXPLANATION OF THE PLATES.

Prare LXXXV.

Hewxelasma aucklandicum Hector sp.

Miocene, Oamaruian, Base of Waitemata Beds: Motutapu Island,

Auckland Harbour, New Zealand.

Fig. 1. Rostrum. a, outer view ; 6, inner view of apical portion of same.

2. Id. a, outer view of small example; 6, inner view of same.

3. Id. a, outer view; 0, inner view of same. :

4. Lateral compartment (right). a@, outer view; 6, inner view of same.

5. Id. (left). Outer view.

3. Id. (right). Outer view of small example.

7. Carino-lateral compartment (right). Outer view.

8. Id. (right). Inner view.

9. Id. (left). Outer view of incomplete example.

10. Carina. a, outer view; 4, side view of same.

11. Id. Outer view of small complete example.

12. Id. Inner view of incomplete example.

13. Tergum. Probably of the same species. a, outer view; 6, inner view of

same.

All figures reduced to } nat. size, except figs. 2, 6, and 9, which are nat. size, and

figs. 13 a, 6, which are X 4 diam.

Prate LXXXVI.

Sealpellum subplanum, sp. n.

Miocene, Oamaruian, Base of Waitemata Beds: Motutapu Island,

Auckland Harbour, New Zealand.

Fig. 1. Carina. a, outer view of basal half of valve; 6, side view; c, transverse

section.

2. Id. a, outer view of basal two-thirds of a smaller valve; 6, side view ;

c, transverse section.

3. Scutum. a, outer view of incomplete valve; 4, inner view of sane.

4. Tereum. a, outer view; 6, inner view of same.

5. Rostrum. Outer view.

6. Subearina. a, outer view; 6, inner view of same.

Scalpellum (Arcoscalpellum) ungulatum, sp. n.

Miocene, Upper Oamaruian, “ Limestone”: Takiroa, Oamaru District,

; N. Otago, New Zealand.

Fig. 7. Carma. a, outer view of basal half of valve; 0, inner view; c, transverse

section.

8. Id. a, outer view of small complete valve; 6, side view.

9. Tergum. a, outer view; 6, inner view of same.

10. Upper latus. a, outer view ; 0, inner view of same.

Miocene, Upper Oamaruian, “ Blue Clay”: Pareora, South Canterbury,

New Zealand.

11. Scutum. a, outer view; 5, inner view of same.

12. Rostral latus. a, outer view; 6, inner view of same.

13. Restoration of capitulum, based on the disconnect®d valves here figured.

The inframedian and carinal latera are not known, but these valves

: have been diagrammatically indicated in the restoration.

All figures of nat. size, except figs. 5, 6, 8, and 12, which are twice nat. size.

PZ. 8. MS, Fil, L2Oovir

OSTEOMALACIA IN A BABOON.

LXCeWVviit.

INS, Il.

lige

3ABOON.

IN A

OSTEOMALACIA

eo 4e Se IIS, Pi, ILSOOMX,

OSTEOMALACIA IN A BABOON,

PROF. T. WINGATE TODD ON OSTEOMALACTA. 855

Observations on Osteomalacia in the Zoological Collec-

tions of Manchester and Cleveland. By T. WinGate

Topp, M.B., F.R.C.S., Professor of ent orie Western

Reserve University, Cleveland, Ohio *

[Received May 5, 1913: Read June 3, 1913. ]

(Plates LXXXVII.-LXXXIX.7)

It is the object of this communication to call attention to the

widespread appearance of osteomalacia among animals in captivity,

and to point out the opportunity afforded by the disease for the

study of changes in formed bone-tissue.

Through the kindness of Messrs. Jennison of Manchester and

Director Springborn of Cleveland, I have been enabled to study

the disease as it exists in the Zoological Gardens of both these

cities.

In Manchester, while the cercopitheques, macaques and small

baboons were kept in a large cage in a moderately heated

monkey-house, the animals sickened and died of miliary tubercle

within a few months after their arrival. When, however, the

glass was removed from the windows of the monkey-house and a

free current of air permitted to pass through the building, the

animals remained healthy for as longas three years. Eventually,

however, most of them begin to show a disinclination to move

about, crouching and walking with the aid of the fore-limbs—the

hind ones being curled up beneath the body in squatting attitude.

When attacked by healthy animals they seem unable to defend

themselves, or, if they try to run away, they do so groaning with

apparent pain. ‘This is more marked if they are compelled to use

their hind legs, as, for instance, in climbing, Left to themselves

they are very much addicted to masturbation. Indeed, in Cleve-

land, my attention was first drawn to the condition by my being

asked to inspect some macaques of both sexes which had developed

this objectionable habit. As the disease progresses the joints

become stiff and enlarged, although post-mortem joint-changes

have not been present in the cases which I have examined. The

disease is not of itself fatal and may last two years, although it

is usually found necessary to destroy the animals after twelve

months. If the animals die, there are frequently indications of

broncho-pneumonia found at the autopsy. The food on which all

the apes, healthy and diseased, are fed consists of rice-pudding,

carrots, onions, potatoes, greens, apples, and bananas, with a little

meat occasionally.

While suffering from the disease, the hair may become dry,

ruffled, or fall out. This is more frequently the case in rabbits

* Communicated by the SecRETARY.

+ For explanation of the Plates see pp. 859-860.

856 PROF. T. WINGATE TODD ON OSTEOMALACTA.

and rats than in monkeys. The disease seems to localize itself

in different sites in different animals. In the macaques and

baboons it usually affects the hind limbs, which are in con-

sequence badly bent and twisted, but that the whole skeleton is

affected is shown by the brittle character of all the bones. The

ability of the bones to unite after fracture does not seem to be

impaired. Fig, 4 (Pl. LX XXVIII.) represents the tibia of a

young Anubis Baboon (Cynocephalus anubis) after healing of

a fracture near the upper extremity. This bone was broken

by a fall from a height of some three feet while the animal was

attempting to escape from a pursuer. The animals may become

affected by the disease while still young and before the epiphyses

have joined, so that the disease is readily confounded with rickets.

The lesions, however, are the same at whatever age the disease

occurs. In some instances the bone-changes are found most

marked in the fore limbs. This was the case in a peceary from the

Manchester Collection. The animal struggled about with its fore

limbs bent backwards beneath the body. In other animals the

bones of the face are affected. The maxille are much enlarged

and give a puffed-out appearance to the cheeks. This condition

occurred in a horse, a leopard, a rabbit, and a chimpanzee. In

some cases the necks of the teeth are decayed and the teeth may

fall out. This was observed in the leopard and rabbit just

referred to, and also in a Californian sea-lion from Cleveland.

Uleeration and falling of the teeth has not occurred among

the apes in Manchester since the glass was taken out of the

windows of the monkey-house. The bones of the chest are

frequently softened and deformed, which would seem to pre-

dispose the animal to respiratory disorders. The lungs of the

sea-lion showed emphysema and patches of broncho-pneumonia,

the latter being confirmed by histological examination.

Post mortem, the viscera show no lesion whatever in animals

which have been intentionally killed. Nor are there any atrophic

changes in the muscles of the disabled limbs.

The ductless glands are apparently normal. Pathological

appearances are to be found only in the nervous system and

the bones. As regards the former opinion is very varied. Gayet

and Bonnet describe an increase in volume of the nerves with

an overgrowth of the fibrous tissue between the nerve-bundles

and disappearance of myelin in certain cases. The blood-vessels

of the nerves exhibit endarteritis (1). They found no cellular

lesion nor any alteration in nerve-fibres in the spinal cord.

Morpurgo, on the other hand, describes diffuse chromatolysis

in the cells of the anterior horns of the cord, but denies any

change in the cells of the spinal ganglia (2). These changes

occur very early in the course of the disease, but there would

appear to be no definite evidence to show that they initiated the

disease (8).

It may be that these different histological pictures were

PROF. T. WINGATE TODD ON OSTEOMALACIA. __ 857

produced by differences in the etiology of the disease. For the

bone-changes can only be a symptom, caused, perhaps, in many

different ways.

The bone-marrow in osteomalacia is much modified. The

marrow-cavities of the long bones are enlarged and filled with a

gelatinous fatty tissue and by bright red marrow. Marrow is

divided microscopically by Ziegler into splenoid and fine-fibred

constituent factors (4). Of these the latter form is much in-

creased, partially filling up the marrow-cavity and penetrating

the bony tissue.

In early cases the compact bone is invaded by this vascular

fibrous tissue, which has already replaced to a large extent the

spongy bone and which is not preceded by osteoclasts. The same

observation has been recorded by Morpurgo (8).

In later stages the whole of the compact tissue of the shaft has

disappeared and is replaced by fibrous tissue, in which are to be

found discreet areas or islets of osseous material (see Pl. LX X XIX.

figs. 5, 6). There is meanwhile no subperiosteal deposit of new

bone. The histological picture of the bone closely resembles that

seen in ostitis fibrosa. The remaining islets of bone lose their

ossein and become transformed into a tissue which, from its

staining properties, appears to be hyaline in character. But, at

the same time, in other situations the regressing bony substance

exhibits a fibrillar change of the ground-substance similar to that

described by Retterer as the basis of normal bone (5).

Cells with similar staining reactions to osteoblasts may still be

seen bordering the islets of bony tissue (see Pl. LX X XIX. fig. 6).

In many places so-called ‘“ osteoclasts” are observed. But, on

examining several slides one is struck by the comparative infre-

quency of these cell-masses. As Gayet and Bonnet remark (1)

it is difficult to believe that osteoclasts can play more than a very

subsidiary part in the destruction of bone, because of their scarcity

when compared with the extent of the process. It would appear

that osteoclasts are not at all necessary for the production of

change in bone-substance.

As the bony tissue becomes transformed or replaced by the

fibrous material, the Haversian systems disappear. The peri-

osteum becomes intimately united to the mass of fibrous tissue

which remains in place of the true bone. I have not observed

hemorrhages in subperiosteal or other localities. The joints were

unaffected. At the diaphyso-epipyseal junction, changes similar

to those in rickets were observed.

The theories of causation of osteomalacia have been well

reviewed by Morpurgo (2), who, among others, has succeeded in

producing the disease by inoculation of a micro-organism (6).

The detailed histological changes have been discussed by

Basset (7), whose description is amply borne out by my own

shdes.

The disease is one which may be secondary to some other

858 PROF. T, WINGATE TODD ON OSTEOMALACIA.

lesion, and therefore localised, as in the variety appearing after

trauma.

It may develop in connection with giant-celled sarcoma and be

more generalised in type (Schonenberger, 8).

It may occur in the so-called spontaneous form, such as is seen

in animals in captivity.

The last-mentioned variety is certainly not a simple decalcifi-

cation. There is absorption of osseous substance with rarefaction

of the tissue. The process starts from the marrow-cavity and

involves first spongy and later compact bone. The Haversian

systems disappear and the bone becomes fibrillar in character and

later is transformed into fibrous tissue.

An intermediate hyaline change is shown in places. All these

changes point to a revolution in the constitution of bone as a

whole, which is accompanied by changes in the marrow and

periosteum, as already described. Similar changes in bony

tissue are to be observed in ostitis fibrosa and leprosy. For it

has been my good fortune to be able to investigate all three

diseases at the same time. The clinical symptoms in these cases

of generalised bone-softening, which I have described as osteo-

malacia, suggest a nervous origin. The obvious inference to be

drawn from the histological picture in leprosy is that in the last-

named disease the bone-changes are certainly trophic in character.

I would emphasise the fact that we are as yet ill-acquainted with

the symptoms consequent on lesions to the sympathetic nervous

system. But there would seem to be ample confirmation of nerve~

lesion in the histological changes found in the nerve-bundles by

Gayet and Bonnet (1). Moreover, the intimal proliferation de-

scribed by these authors in the vessels of the nerve-trunks may

be produced by a lesion in the sympathetic nerves, as I have

recently been enabled to show (9).

If the disease is infectious, the incubation-period must be con-

siderable, for it seems to appear spontaneously in animals which

have been isolated for a long while. After inoculation, Morpurgo

found the animal became ill in a week or two. Such evidence as

we have points to the nervous system as the seat of primary

disease, whether it be infectious or not, and suggests that the

bone-changes are consequent on nervous lesion. Treatment is

unsatisfactory. Dr. Fox, of Philadelphia, has recently adminis-

tered calcium lacto-phosphate and adrenalin, separately and in

combination, to animals suffering from the disorder in the

Philadelphia collection, but without success (10).

In making observations on the living animals Messrs. Antliff

and Readinger, keepers of the monkey-houses at Manchester and

Cleveland respectively, have rendered generous assistance. The

histological sections are the work of Mr. Gooding, of the Ana-

tomical Department of Manchester. Mr. J. C. Miller, of the

Laboratory here in Cleveland, has assisted me in gathering and

abstracting the literature. To all these gentlemen, I would

therefore express my obligation.

PROF, T. WINGATE TODD ON OSTEOMALACIA.. - 859

Summary.

1. Osteomalacia, or so-called spontaneous generalised bone-

softening, is not a simple decalcification of bone, but a re-

organisation of bone as an organ, in which the loss of osseous

tissue is not brought about by osteoclasts.

2. From this it is evident that the cell-masses known by the

name of “ osteoclasts” are not indispensable in the transformation

of bone to less specialised tissue.

3. The disease may be infectious in origin. If so, the evidence

at our disposal points to the nervous system as the site of infec-

tion. The bone-changes appear to be the symptoms consequent

on the nervous lesion.

REFERENCES.

(1) Gaver et Bonner.—“‘ Contribution 4 l'étude des ostéo-

malacies.” Revue de Chir. vol. xxiii. pp. 44, 228, 1901.

(2) Morpurco.—‘ Ueber eine infectidse Form der Knochen-

briichigkeit bei weissen Ratten.” Verhandl. d. Deutschen

Path. Ges. vol. iii. p. 40, 1901.

(3) Morpurco.—‘ Durch Infection hervorgerufene malacische

und rachitische Skeletveranderungen an jungen weissen

Ratten.” Cent. f. Allg. Path. u. Path. Anat. Bd. xiii.

p. 113, 1902.

(4) Zrecter.— Ueber Osteotabes infantum u. Rachitis.” Cent.

f. Allg. Path. u. Path. Anat. Bd. xii. p. 865, 1902.

(5) Rerrerer.—‘ Structure et histogénése de Vos.” Journ. de

VAnat. et de la Physiol. p. 561 eé seg., 1905.

(6) Morpurco.—‘ Ueber die infectidse Osteomalacie und

Rachitis der weissen Ratten.” Verhandl. d. Deutschen

Path. Ges. vol. xi. p. 282, 1907.

(7) Basser,—“ Anatomie pathologique de ’Ostéomalacie spon-

tanée et expérimentale.” Arch. de Med. Expérimentale,

Meant, joy (sy IQOG.

(8) SCHONENBERGER.—‘ Ueber Osteomalacie mit multiplen

Riesensarkomen und multiplen Fracturen.” Virchow’s

Archiv, Bd. clxv. p. 189, 1901.

(9) Topp.—“ The Arterial Lesion in Cases of Cervical Rib.”

Journ. Anat. & Physiol. vol. xlvii., 1913.

(10) Fox.—The 40th Annual Report of the Board of Directors

of the Zoological Society of Philadelphia, p. 55, 1912.

EXPLANATION OF THE PLATES.

Prats LXXXVII.

Fig. 1. Right Radius from young Anubis Baboon (Cynocephalus anubis). Advanced

stage of osteomalacia.

Note large size and dark colour of marrow cavity. ‘The dark appearance is

due to the large quantity of red marrow. The rarefaction of bone and the

progression of the osteoporosis from marrow-cavity outwards is well shown

in this photograph. The compact tissue can be seen to have been replaced

by spongy bone.

Proc. Zoou. Soc.—1913, No. LVII. 57

Fig. 3.

Fig. 5.

PROF. T. WINGATE TODD ON OSTEOMALACTA.

. Right Ulna from same animal. Here and there are districts of gelatinous

fatty marrow giving rise to lighter areas between the dark-coloured sites

of red marrow. The thickened periosteum and its intimate union with

the tissue composing the bone can be seen well, especially near the centre

of the shaft. In this situation the bone was accidentally broken while

stripping off the muscles.

Prats LXXXVIII.

Right Humerus of same animal.

Note the irregular diaphyso-epiphyseal junction at the upper end of the

shaft. The extreme brittleness of the bones resulted in the fracture of the

surgical neck of this humerus while the muscles were being removed.

There is no sharp line between the articular cartilage of the condyles and

the underlying spongy bone. This photograph shows clearly the articular

cartilage being destroyed on its bony aspect.

. Right Tibia and Fibula from same case.

Note extreme recurved upper portion of tibia. The bone had been fractured

some time previously and had united in this position. The union is com-

plete. The fibula was likewise bent, its lower extremity is cut obliquely

and is shown behind the lower portion of the tibia.

Prate LXXXIX.

Longitudinal section of shaft of Left Ulna from the same animal. X40.

Note the entire disappearance of Haversian systems from the bone and the

replacement of osseous tissue by fibrous material. The bony substance is

here shown broken up into islets. The section represents what once was

the compact bone layer of the shaft.

. Longitudinal section of shaft of Left Ulna from the same animal. 206.

A bone-islet is shown surrounded by non-osseous tissue. Note the gradual

transition of the bone into fibrous tissue. There is no sharp line of

demarcation, and no osteoclasts are to be observed. The decalcification

progresses from the centre of the illustration to the periphery. Inter-

mediate between bone and fibrous tissue is a zone where the nuclei are

surrounded by considerable non-calcitied protoplasm. This corresponds

in appearance to the pre-osseous stage in Retterer’s description of

bone (5).

No. 122.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

May 20th, 1913.

Prof. EH. A, Mincuin, M.A., F.R.S., Vice-President,

in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

The Srcrerary read a Report on the Additions that had been

made to the Society's Menagerie during the month of April

1913.

The Lrprarian submitted a list of dates of publication of the

early parts of the Society’s ‘Transactions,’ drawn up from the

records kept by Messrs. Taylor & Francis, printers to the Society.

Mr. R. E. Hotpine exhibited a large number of specimens and

photographs illustrating variations in the growth of the Antlers

of Deer. .

The Rev. T. R. R. Srespine, M.A., F.R.S., F.Z.S., drew

attention to Prof. F, E. Schulze’s important work on zoological

nomenclature, the ‘Nomenclator Animalium,’ giving a brief

description of its objects and scope, and urging its claims for

assistance from British naturalists.

Dr. R. Broom, C.M.Z.S., read a paper “On the South African

Pseudosuchian Reptile Huparkeria and allied Genera.” Besides

* This Abstraet is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Sic Shillings per annum, payable im adyance.

giving an account of the very completely known South African

form, he also discussed the structure of the Elgin allied forms,

Ornithosuchus and others. The group of Pseudosuchians he

regarded as an extremely important primitive reptilian order,

as there is good reason to believe that not only does it eontain the

ancestor of the Dinosaurs, but also the aneestors of the Ptero-

dactyles and Birds. Huparkeria and Ornithosuchus are, in

structure, almost Dinosaurs, and it is held that when the bipedal

habit was more fully acquired the few characters not quite

Dinosaurian would become Dinosaurian. Birds are held to

have originated from a Pseudosuchian which, by a bipedal habit,

had acquired a Dinosaur-like hind limb, and had then become

arboreal in habit and acquired the peculiar power of flight. :

Mr. E. G. Bouneneer, F.Z.S., Curator cf Reptiles, read a paper

giving an account of the experiments which he had, for the past

year, been conducting on the Metamorphosis of the Mexican

Axolotl (Amblystoma tigrinum), and gave a detailed description

of the changes that take place in the course of transformation.

He also exhibited a number of specimens in the perfect or

Amblystome condition. The conclusions arrived at by the

author, as a result of his experiments, were that, in accordance

with Mlle. de Chauvin’s experiments, and contrary to those of

Dr. Powers, the Axolotl will, with a few exceptions, transform

if placed under special conditions which force it to breathe air

more frequently than usual; that starvation, irregular feeding,

and temperature have no influence on the metamorphosis; that

the elimination of oxygen from the water has likewise no bearing

on the point, as the animal will not, under the circumstances,

rise to the surface and make use of its lungs at more frequent

intervals than animals placed under normal conditions.

Mr. G. E. BuLien contributed a short paper, communicated by

Mr. John Hopkinson, F.Z.8., “On some Cases of Blindness in

Marine Fishes.” Work hitherto performed, e.g. that of Hofer,

de Drouin de Bouville, and others, upon the pathology of fishes

has been directed largely upon species of freshwater habitat.

The present author has found, in certain specific cases of blindness

in marine fishes, pathological conditions similar to those described,

and others with slight modifications, in several freshwater species.

The examples dealt with in detail are traumatic corneal opacitis

in a Conger-Hel, corneal opacitis, etc., in a Greater Weaver,

and corneal opacitis and cataract in a Pollack.

Dr. R. W. Suurevpt, C.M.Z.8., sent a paper dealing with the

Patella in the Phalacrocoracidee. From a study of the patella in

a number of species of this family, he had found that in adult

individuals that bone was composed of the true patella solidly

fused with the proximal portion of the cnemial process of the

tibio-tarsus, which became dissociated from the latter early in

the life of the bird. Late in life this fusion obliterated the

tendon of the ambiens muscle, which heretofore had been

described as passing through the patella and persisting through

life,

The next Meeting of the Society for Scientific Business (closing

the Session 1912-1913) will be held on Tuesday, June 3rd, 1913,

at half-past Hight o’clock p.m., when the following communications

will be made :-—

EXHIBITIONS AND NOTICES.

Sir ArrHor H, Cuurcu, K.C.V.O., M.A., D.Sc., F.R.S., F.S.A.

Notes on Turaecin and Turacin-bearers.

-P. Coatmers Mitcuent, M.A., D.Sc., LL.D., F.R.S., F.Z.8.

Observations on the Anatomy of the Shoe-bill (Baleniceps

Tex).

T. H. WitHers, F.G.S,

Some Miocene Cirripedes of the Genera Hexelasma and

Scalpellum from New Zealand.

Prof. ArtHuR Denpy, D.Sc., F.R.S., F.Z.8., and R. W. H. Row,

B. Se.

The Classification and Phylogeny of the Calcareous Sponges,

with a Reference List of all the described Species, systematically

arranged.

Surgeon JosepH C. THomeson, U.S.N.

Contributions to the Anatomy of the Ophidia.

Prof. T. WineatE Topp, M.B., F.R.C.S.

Observations on Gsteomalacia in the Zoological Collections

of Manchester and Cleveland.

The following paper has been received :—

§ pap

D. M.S. Watson, M.Se.

Batrachiderpeton lineatum Hancock & Atthey, a Coal Measure

Stegocephalian. :

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpon,

Re@ent’s Park, Lonpon, N.W.

May 27th, 1913,

No. 123.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

June 3rd, 1918.

Prof. EK. W. MacBripz, M.A., D.S8c., F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. D. Sersa-Smitu, F.Z.S., Curator of Birds, exhibited the

ege and young of the Mikado Pheasant (Calophasis mikado), a

rare species, described first in 1906, from the mountains of

Formosa. Some living specimens had been imported in 1912 by

Mr. Walter Goodfellow, and the owners of these birds had

entrusted the eggs to the Zoological Society, where they were

being hatched.

The egg was cream coloured and very large compared with

those of allied species of pheasants, being 57 x 41 mm.

The incubation period proved to be twenty-eight days, instead

of twenty-four as in the majority of pheasants, and the young

when newly hatched were very large, and had the quill-feathers

better developed than was the case in allied species,

Sir Artuur H. Caurcu, K.C.V.O., F.R.S., read a paper, com-

municated by the Secretary, entitled ‘‘ Notes on Turacin and the

'Turacin-bearers.”

This paper contains a summary of the chief facts as to the

composition, properties, and occurrence of turacin, the soluble

crimson pigment of the Musophagide. Special stress is laid upon

its constancy of composition, the limitation of its occurrence to

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

‘it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in adyance,

certain plantain-eaters, and the relation of its spectrum to the

spectra of hemoglobin and chlorophyll. Some current errors

concerning turacin are corrected.

The Secretary, Dr. P. Coatmers Mrircuet, F.R.S., gave an

account of his observations on the Anatomy of the Shoe-bill

(Baleniceps rex), illustrating his remarks with lantern-slides.

He showed that Baleniceps and Scopus shared so many anatom-

ical characters, and of these so many occurred also in Storks, that

if the reasoning generally followed by anatomical ornithologists

were adopted, Baleniceps and Scopus must be placed with Storks

rather than with Herons. He submitted, however, that such a

method was irrational, unless it were accompanied by a much

closer scrutiny of the value of the characters than had hitherto

been made or was yet possible, and that for the present Baleniceps

must be regarded as the representative of a division equivalent to

Storks and Herons. He thought also that the relation of the

Steganopods to these three groups required reconsideration.

A paper on ‘‘Some Miocene Cirripedes of the genera Hexelasma

and Scalpellum from New Zealand,” communicated by Dr. W.

T. Carman, F.Z.8., was read by Mr. T. H. Wrrumrs, F.G.S.

An account is therein given of the ‘gigantic Cirripede’ of New

Zealand, originally deseribed as Scalpellum aucklandicum, of

which remains have long been known to occur in the Waitemata

Beds (Miocene) of Motutapu Island, Auckland Harbour. The

valves of this Cirripede attain a length of 8 inches, and have

been previously supposed to belong to a pedunculate form, but

while Sir James Hector (1887) referred them to the genus

Scalpellum, Prof. W. Blaxland Benham (1903) thought that they

approached more closely to the genus Pollicipes. From a study

of the criginal material collected by Prof. James Park (1887),

it is now shown that this Cirripede is a sessile form allied to

Balanus, and it is referred to Dr. P. P. C. Hoek’s recently

instituted genus MHexelasma (1913). A smaller undetermined

species of Hexelasma, and a new species of Scalpellum (sensu lato),

are also described. These are in the collection of the Geological

Survey, New Zealand, and occur in the same beds as the

‘ vigantic Cirripede.’

A second new species of Scalpellum is founded on some valves

from New Zealand, and a restoration is given, the remains

being sufficient to justify their reference to the sub-genus

Arcoscalpellum Hoek.

A paper on “ The Classification and Phylogeny of the Calcareous

Sponges, with a Reference List of all the known Species, sys-

tematically arranged,” was received from Prof. Artaur Denby,

D.Sc., F.R.S., F.Z.8., and Mr. R. W. Haroip Row, B.Sc., F.L.S.

This memoir aims at a complete revision of the genera of

Caleareous Sponges. Fifty recent genera are recognised and

diaynosed, and all the described species, amounting to 433, are

arranged under these genera. The rejected generic names, which

are listed separately, amount to 97. The fifty accepted genera

are grouped in 10 families, and Poléjaeff’s subdivision into

Homocaita and Hursrocata is abandoned. The systematic part

of the werk is prefaced by a discussion on the principles of

classification, and followed by a discussion on the phylogeny of

the group, accompanied by a genealogical tree. The suggestions

of Minchin and Bidder as to the systematic value of the position

of the nucleus of the collared cell is followed up, and this position

has been determined in 75 species. It appears from the evidence

thus secured, taken, of course, in connection with other characters,

that in the primitive family Homoceelide the position of the

nucleus is basal in some species and apical in others, but that it

is not correlated with other characters so as to justify a sub-

division of the family accordingly. From the Homoceelide,

however, two lines of descent have sprung, the one basinucleate

and the other apicinucleate; the former including the families

Leucascide, Leucaltide, Minchinellide and Murrayonide, and

the latter the Sycettide, Heteropiide, Grantiide, Amphoriscide

and Lelapiidee.

Surgeon J. C. Tuompson, U.S.N., sent a paper, communicated

by Dr. F. E. Bepparp, M.A., F.R.S., F.Z.S., containing eon-

tributions to the Anatomy of the Ophidia,

The SzcreraRyY communicated a paper by Prof. T. WincATE

Topp, M.B., F.R.C.S., entitled ‘“ Observations on Osteomalacia in

the Zoological Collections of Manchester and Cleveland.”

——*

This Meeting closes the Session 1912-1913. The next Meeting

of the Society for Scientific Business will be held on Tuesday,

October 28th, 1913, at half-past Hight o’clock P.M.

The following papers have been received :—

1. D. M.S. Warson, M.Sc.

Batrachiderpeton lineatum Hancock & Atthey, a Coal-Measure

Stegocephalian,

2. R. W. Parmer, B.Sc.

The Brain and Brain-case of a fossil Ungulate of the Genus

Anoplotherium.

3. KREDERIC WILMET.

Notice sur l!’Okapia johnstoni, dont Vespece s'est retrouvée

intacte dans une forét commune a |’Ituri et 4 1 Uelle.

_ Communications intended for the Scientific Meetings should

be addressed to ;

P, CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF LonpDoN,

Recurnt’s Park, Lonpon, N.W.

June 10th, 1913.

30,

3l,

32,

33.

34.

35.

36.

37.

38,

39.

40.

“ill

42,

43.

44,

45:

46,

Papers (continued).

Page.

On the Patella in the Phalacrocoracide. By Dr. R. W. Suursupt, C.M.Z.S.

(Pl, LXI.) Bese eceeecseeeeer set HOF se se sett SPFFso®F es ee ee re eee FH s7 Hse 289899 OD 393

Experiments on the Metamorphosis of the Mexican Axolotl (Amblystoma tigrinum),

conducted in the Society's Gardens. By EH. G. EOUUENS ¥.Z.S., Ourator of

Reptiles. (Text-figs. 75 & 76.) ...... eel ateraie stares sfoie wists eres Bes wegicarets msttaliece's 9 400

Oontributions to the Anatomy of the Ophidia. By Josrru C. Tuompson, Surgeon,

United States Navy. (Text-figs. 77 & 78.) ........++0+.00. mis cletets, Sioielo reise Se faite, 414

The Polyzoa of Waterworks. By Sipney Bo Harmer, S8c.D., F.RS., F.ZS.

(ELS SEGRE aks IDG NTE Siem iichno anion ne on dace cocCU moan AMC ne OOO ED secre emia 4

The Marine Fauna of British East Africa and Zanzibar, fom Collections made by Cyril

Crossland, M.A., B.Sc., F.Z.S., in the Years 1901-1902. Bryozoa—Cheilostomata.

By Artnur Wu. Waters, F.L.S., F.G.8. (Pls. LXIV.-LXXIII. and Text-figs, 79-82.) 458

Notes on Albinism in the Common Reedbuck (Cervicapra arundinum), and on the

Habits and Geographical Distribution of Sharpe’s Steenbuck (Raphiceros sharpet).

By Major J. Srnvenson-Hamitton, C.M.Z.S., Game-Warden of the Transvaal ........ . 537

Contributions to the Anatomy and Systematic Arrangement of the Cestoidea,—X. On

Two Species of Tapeworms from Genetta dongolana, By Frank H, Bepparp, M.A,,

D.Sc., F.R.S., F.Z.S., Prosector to the Society. (Text-figs. 85-94.) -.........s2e000 549

Pacific Salmon: An Attempt to evolve something of their History from an Examin-

ation of their Scales. By Jonn Apam Mitnu, of Ardmiddle, Turriff, Aberdeenshire.

Uber trap O STONE som gee scars moe caress oh ierotai oisr ae ctecaisinGr Stee uae win slaice @ ease e accra srure tie 572

Notes on Peripatoides woodwardi Bouvier. By Karnuuun Happon ......00+-eeeees 611

Field-Observations on the Enemies of Butterflies in Ceylon. By J. C. F. Fryur, M.A,

F.E.S., Fellow of Gonville and Caius College, Balfour Student in the University of

Cambridge ..... Nstvecionshiortu veins ioha ist eaeepart By etre orovslentt atatetera orca a oat ora Miibsises eo. 613

On the South African Pseudosuchian Huparkeria and Allied Genera, By R. Broom,

On some Cases of Blindness in Marine Fishes. By G. E. Butusn, the Hertfordshire

AVES SUITING SEMAN DATS ce Gisece cenclretat nS mesic oadeaeee a aienrs oe Gc opeUN rear Shae alawte Slee orate e ciinTowacn olateTece . 634

Notes on Turacin and the Turacin-Bearers. By Sir ArtHur H. Onurcn, K.C.V.O., F.R.S. 639

Observations on the Anatomy of the Shoe-bill (Baleniceps rex) and allied Birds. By

P. Cuaumers Mircnent, M.A., D.Sc., LL.D., F.RS., ey of the Society.

(Pls. LXXX.-LXXXIII, and Text-figs. 119-132.) perder siete stere ores Sa isrelaereles sais ce 644

The Olassification and Phylogeny of the Calcareous Sponges, with a Reference List of

all the described Species, systematically arranged. By Artuur Denpy, D.S8c., F.R.S.,

F.Z.S., Professor of Zoology in the University of London, and R. W. Haroup Row,

B.Sc., F.L.8., Assistant Lecturer and Demonstrator in Zoology at King’s College.

(CRextafe Gos) ~unbes ss etereraitine se sary wey = 5 mray oboe ofer eta ceatshsTOMh tera =ce cyefaietes vast ciel atets 704

The Transvaal Race of the Cape, or Khama, Hartebeest. By R. Lypzxxnr, F.R.S.,

InolAds bes (USS ¢Failanel BIR Boca. ire rin Capi St. iad Gn ano eid OE ene Bint ceo oe 818

On a Collection of Mammals from the Inner Hebrides. By G. EH. H. Barrent-

- Haminton, F.Z.8., and Martin A.C, Hinron. (Pl. LXXXIY. and Text-figs. 186-138.) 821

47.

48.

Some Miocene Cirripedes of the Genera Hexelasma and Scalpellum from New Zealand.

By Tuomas H. Wirunrs, F.G.8, (Pls. LXXXV. & LXXXVI. and Text-figs. 189 & 140.) 840

Observations on Osteomalacia in the Zoological Collections of Manchester and Cleve-

land. By T. Wincatx Topp, M.B., F.R.C.S., Professor of Anatomy, Western Reserve

University, Cleveland, Ohio. (Pls, LXXXVII.-LXXXIX.) ...... SGonUOhHOnoe acre Leow

wee Page

L. \ :

LI. |

LIT. } Costa Rican Butterflies ............¢ Bae APIA RAS _ 839

LIII. |

LIV. )

LV. ° Acanthistius fuscus ...cccesccree Que moomoO Ne ooodDS )

NaS Kahiiavmiutabunda 6.) eres seen cin inicio ts

LVIL. ~Girellops nebulosus 2. ce ce oes s cco uc uceusecuceses

LVIIL. 1. Pomacentrus inornatus. 2: Labr ichthys big + 368

3. Anampses pulcher....c+..ccrecess AE OTONU Dog Ho

LX. Pseudomonacanthus paschalis .....0..eccersces-vecees )

LXI. Patella of Phalacrocoracidz .........+2+cccceseees -. 898.

PRU Pa idicella Greiculacd, ame mate vies ce a cn eleake s+ | 496

LXIII. Statoblasts of Phylactolemata ........c..cseeeoecees \

LXIV.)

LXV. |

LXVI.

LXVII.

ea (eago7es from Zanzibar .....+++e06 NAIA D RSIS ernie e 458

LXx.

LXXI. |

LXXxIi.

LXXIIt. )

LXXIV. Zonurus giganteus. Rone as oar quaamoo as “sities eke 548°

ene \ Huparkeria capensis ....-- Sher inatietavate eters canine aRecCtene Te i.

LXXVII. 9,10. Ornithosuchus woodwardi. 11. O, taylori.. ee

LXXVILI. Mesosuchus browns and others.....0+..cessecccescees

LXXIX. Browniella africana and others .....-....-- sie eee ae ae

POO TAMIUGHSOS REO ebo co ogo pon pO OU Oo CUO ONO OUbndOGaCdr:

LXXXI, Skulls of 1. Baleniceps. 2. Scopus. 3. enoriynchis

LXXXII. Skulls of 1. Dissura. 2. Tantalus. 3. Cancroma....

UXXXIIT. Skulls of 1. Ardea. 2. Pelecanus. 3. Plotus......2-..

LXXXIV. 1. Sorex grantii. 2. S. araneus castaneus..

LXXXV. Hexelasma auchlandicum Hector sp. ..++-...-ses.seeee

LXXXVI. 1-6. Scalpellum subplanum. 7-13. 8S. (Arcoscalpellum)

UMGULACUTU : ons si vinin® = onto) dere eis = nidietelale reheat an

LXXXVII.

LXXXVIII. | Osteomalacia ina Baboon ......-- eS elaec ere eaeeareicia eee

LXXXIx.

LIST OF PLATES.

1918, Part III. (pp. 339-860).

NOTICE.

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,

so that the complete reference is now P. Z. 8.1913, p.... The Distribution

is as follows:—

Part I. issued in March.

SLs a June.

Poise [El Beeler, September.

Peseta AZ December.

‘ Proceedings,’ 1913, Part II. (pp. 153-337), were published on

May 30th, 1913.

The Abstracts of the ‘ Proceedings,’ Nos. 122 & 123, are

contained in this Part.

SE DS DEP

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.

(1913.

PART IV.

- CONTAINING Paces 861 to 1104, witH 24 PLaTEs

xf eS ae ae ee at 0". — —

J . eae es eee our

3 ’ : 5 ay

; : . 4 Ste

* °

as .

AND 55 TEXT-FIGURES.

DECEMBER 1913.

PRINTED FOR THE SOCIETY,

SOLD AT ITS HOUSE IN REGENT’S PARK.

LONDON:

MESSRS. LONGMANS, GREEN, AND CO,,

PATERNOSTER ROW.

ee a

[Price Twelve Shillings.)

LISET SOF CONT ENS:

1913, Part IV. (pp. 861-1104).

EXHIBITIONS AND NOTICKS.

Page

The Szcrerary. Report on Additions to the Society’s ee during the months of

“Tue Secrnrary, Exhibition of the Alimentary Tract of a young Elliot's Pheasant

G@Phastanusielliote) ae caccies sae oles bs Sule See Ee ee toe LEE Gee 1094

Mr. RB. I. Pococr, F.R.S., F.Z8. Exhibition of young Woolly Opossums (Philander

laniger) and young Degus (Octodon degus) .. 0... .cccer sere oe ce nc ress cdeeeeoers 1094

Mr. D. Srtu-Suiru, F.Z.8. Heel-Pads in a young Toucanet (Selentdera macuitrostris).

(Moxt Roe mlOd\cctek suis as inctseis ser pi\ p weciee yummie en gana ior noe aeT: Sona ods: +. 1095

Mr. CO. Tarn Recan, M.A., F.Z.S. A Deep-Sea Angler-Fish, Melanocetus johnsonii .... 1096

Mr. E. G. Bouncer, F.Z.S. Exhibition of a young specimen of the Matamata Terrapin

(QSL HED EOE HRT OC pe oc ies oro iitiow ee hee nee cone ene 1097

Mr. D. Setu-Ssuru, F.Z.8. Exhibition of Hybrid Birds. (Text-fig. 192.) ............ 1098

Mr. R. I. Pocock, F.R.S., F.Z.S. Dorsal Glands in Armadillos. (Text-figs. 193-195.) .. 1099

The SecreTary. Report on Additions to the Society's Menagerie during the month of

October NOlSr 2. .5.< a. SAO NGG Ga ote Oa aN OU SO ‘Gaielletabeis acetate eee 1103

Prof. H. Maxweti Lerroy, M.A., F.Z.8. Remarks upon the New Insect House..... o.- 1104

Mr. R. H. Borns, M.A., F.Z.8. Exhibition of Foetal Skeletons prepared by the Beale-

Selulizepme tod yejr-couetey civ enelasereve's. ors ote 5 ots vs oteseupinlnte oy tecwions cite eet ao hole ees ene eee 1104

PAPERS.

49. Contributions to the Anatomy and Systematic Arrangement of the Cestoidea,—XI. On

a new Tapeworm from (dicnemus. By Frank HK. Bupparp, M.A., D.Sc., F.BS.,

F.Z.S., Prosector to the Society. (Text=figs. 141149.) oe os en's ot niin sm sips ea &61

50. The Brain and Brain-Case of a Fossil Ungulate of the Genus Anoplotherium. By

R. W. Paumer, M.Sc. (Manchester), Research Fellow in Zoology, University College,

Reading; (Dext-feeS0= Laie): wih csistls w nlas ae oma Setar ae Maric eo bb eae 878

Contents continued on page 3 of Wrapper.

OF LONDON.

ZOOLOGICAL SOCIETY

Tars Society was founded in 1826 by Sir Sramrorp Rarries,

Mr. J. Sasrne, Mr. N. A. Vigors, and other eminent Naturalists,

for the advancement of Zoology and Animal Physiology, and for the

introduction of new and curious subjects of the Auimal Kingdom,

and was incorporated by Royal Charter in 1829,

Patron.

HIS MAJESTY THE KING,

COUNCIL.

HIS GRACE THE DUKE OF BEDFORD, K.G., F.RS, President.

Str Jonn Rost BrapForp, Str Watters Roper Lawrence,

K.C.M.G., M.D., D.Sc., F.RB.S.,

Vice-President.

Ricuarp H. Burns, Ese., M.A.

Sure.-Genu. Srr R. Havenock

Cuargtes, G.C.V.O., M.D.

Tue Rr. Hon. Toe Eart or

Cromer, P.C., G.C.B.,

G.C.M.G., K.C.8.1., F.R.S.,

Vice-President.

F, G. Dawrrey Drewitt, Ese.,

M.A., M.D.

Caartes DrumMonp,

Treasurer.

Kse.,

Str Epwarp Duranp, Br., C.B.

Freperick Gittett, Ese.

Tue Lorp GLENCONNER,

F, Du Canz Gopman, Ese.,D.C.L.,

F.B.S.

Br., G.C.LE.

Str Epwunp G. Loper, Br., Vice-

President.

Ernest W. MacBripr, Ksa.,

M.A., D.Sc., F.R.S., Vice-

President.

Proressor Epwarp A. Mrycurn,

M.A., F.R.S., Vice-President.

P. Caatmers Mrrcnert, Ese.,

Wises IDES, IA aID) 5 JB oIey sie.

Secretary.

W. R. Ogitviz-Grant, Ese.

Aprian D. W. Pottocg, Ese.

OtprieLtp Tuomas, Kse., F.R.S.

AntHony H. WineFrrezp, Hse.

Henry Woopwarp, Ksa., LL.D.,

E.R.S., Vice-President.

The Society consists of Fellows, and Honorary, Foreign, and

Corresponding Members, elected according to the By-Laws. It

carries out the objects of its foundation by means of the collection

of living animals, by its Library, and by its Scientific Publications.

The Office of the Society, Regent’s Park, N.W., where all com-

munications should be sent, addressed to “The Secretary,” is open

from Ten till Five, except on Saturdays, when it closes at One p.m.

The Library, under the superintendence of Mr. Henry G. J. Peavot,

is open daily (except Sunday) from Ten till Five, except on

Saturdays, when it closes at Two p.m. The Library is closed from

Good Friday to Easter Monday, and upon all other Bank Holidays.

It is also closed annually for cleaning purposes during the whole

month of September.

The Meetings of the Society for General Business are held in the

Meeting Room at the Society’s Office on the third Wednesday in

every month of the year, except in September and Octeber, at half-

past Four o’clock p.m.

The Meetings for Scientific Business are held in the Meeting

Room at the Society’s Office fortnightly on Tuesdays, except in

July, August, September, and December and January, at half-past

Hight o’clock p.m.

The Anniversary Meeting is held on the 29th. of April, or the

nearest convenient day, at Four p.m.

The Society's Gardens are open daily from Nine o’clock until

Sunset. Mr. R. I. Pocock, F.R.S., F.L.S., is the resident Super-

intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator

ot Birds and Inspector of Works, Mr. E. G. Boulenger is Curator

ot Reptiles, and Prof. H. M. Lefroy is Honorary Curator of Insects.

The Prosectorium for Anatomical and Pathological work is under

the charge of Mr. Frank E. Beddard, M.A., D.Sc., F.R.S., Prosector,

assisted by Mr. H. G. Plimmer, F.R.S., M.R.C.S., Pathologist to

the Society.

‘TERMS FOR THE ADMISSION OF FELLOWS.

Frrrows pay an Admission Fee of £5, and an Annual Contri-

bution of £3, due on the Ist. of January, and payable in advance,

or a Composition of £45 in lieu thereof; the whole payment,

including the Admission Fee, being £50.

No person can become a Frtxow until the Admission Fee and

first Annual Subscription have been paid, or the annual payments

have been compounded for.

Frtiows elected in November and December are not liable for

the Subseription for the vear in which they are elected.

PRIVILEGES OF FELLOWS.

Fettows have Personal Admission to the Gardens upon signing

their names in the book at the entrance gate, and may introduce

''wo Companions daily.

The Wire or Hussanp of a Fattow can exercise these privileges

in the absence of the Fellow.

Until further notice, Frtnows will receive 40 undated Green

Cards, available on any Sunday or week-day up to the end of

February of the year following the year of issue, and 20 White

Cards available on any week-day up to the same date. Twenty

of the Green Cards may be exchanged for a book containing two

Orders for each Sunday in the year. ‘Twenty White Cards may

be exchanged for a book of dated Saturday Orders, each Order

available for any day except Sunday in the week ending with the

Saturday for which it is dated. Special children’s tickets will no

longer be issued, but the Green and White Cards will be perforated,

and each half will be valid for a Child under twelve years of age.

It is particularly requested that Fellows will sign every ticket before

it goes out of their possession. Unsigned tickets are not available.

Frettows are not allowed to pass in friends on their written

order or on presentation of their visiting cards.

Frertows may obtain a TransreraBLe Ivory Ticker admitting

two persons, available throughout the whole period of Fellowship,

on payment of Ten Pounds in one sum. A second similar ticket

may be obtained on payment of a further sum of Twenty Pounds.

Frttows have the privilege of receiving the Society’s ordinary

Publications issued during the year upon payment of an additional

Subscription of One Guinea. This Subscription is due upon the

1st. of January, and must be paid before the day of the Auniversary

Meeting, after which the privilege lapses. FxEttows are likewise

entitled to purchase these Publications at 25 per cent. less than

the price charged to the public. A further reduction of 25 per

cent. is also made upon all purchases of Publications issued prior

to 1881, if above the value of Five Pounds.

Frtiows also have the privilege of subscribing to the Annual

Volume of ‘The Zoological Record,’ which gives a list of the Works

and Publications relating to Zoology in each year, for the sum of

One Pound Ten Shillings. Separate divisions of volumes 39 to

42 can also be supplied. Full particulars of these publications can

be had on application to the Secretary.

Any Fetrow who intends to be absent from the United Kingdom

during the space of at least one year, may, upon giving to the

Seeretary notice in writing, have his or her name placed upon the

“dormant list,” and will then be called upon to pay an annual

subscription of £1 only during such absence, but after three years

must make a further application to be retained on that list.

Any Fertxow, having paid all fees due to the Society, is at liberty

to withdraw his or her name upon giving notice in writing to the

Secretary.

Ladies or Gentlemen wishing to become Fellows of the Society

are requested to communicate with “The Secretary.”

P., CHALMERS MITCHELL,

Secretary.

Regent’s Park, London, N.W.

December, 1913.

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

FOR

SCIENTIFIC BUSINESS.

1914.

TuErspay, FEBRUARY .... 3rd and 17th.

i WOMROTEE! Bo 5.00 ¢ 3rd and 17th.

a END a OWNS ea 7th and 21st.

a IN IAgY.S eta moa 5th and 19th.

AOS GD ag BeatGlo.c 9th.

The Chair will be taken at half-past Light o'clock in the Evening

precisely.

ZOOLOGICAL SOCIETY OF LONDON.

THE ZOOLOGICAL RECORD.

ee object of the ZooLoeicaL RecorD is to give, by means of an

annual Volume, complete lists of the Works and Publications

relating to Zoology in all its branches that have appeared during

the year preceding the issue of the Volume, together with full

information as to the points they deal with, arranged in such a

manner as to serve as an Index to the literature of Zoology in all

parts of the globe, and thus to form a repertory that will retain its

value for the Student in future years.

The ‘ Zoological Record’ having been amalgamated with the

International Catalogue of Scientific Literature, Zoology, volumes

from 43 onwards can now be obtained only from Messrs. Harrison

& Sons, except when purchasing complete sets from the Zoological

Society.

Under the scheme of amalgamation, Fellows of the Society, and

Institutions already on the subscription-list, have the privilege of

subscribing at the old rate of 30s. per annum, which covers the

cost of carriage of the volume. The subscription becomes due on

July lst. in each year, and lapses if not paid by the Ist. of December

following.

The Society is able to supply complete sets of the Record on the

following terms:—

Vols. 1 to 42, price £16 10s. net. Vol. 43 and onwards at 40s. each.

The prices for separate volumes are as follows :—

Vols. 1 to 42 (except Vols. 4 and 6 which are sold with sets only) 10s.

The price of the ‘Zoological Record, Vol. 43 and subsequent volumes,

obtainable separately only from Messrs. Harrison and Co., is 40s.

each.

Index Zoologicus. An alphabetical list of names of genera

and subgenera proposed for use in Zoology, as recorded in the

‘Zoological Record,’ 1880-1900; together with other names not

included in the ‘ Nomenclator Zoologicus’ of 8. H. Scudder. Com-

piled (for the Zoological Society of London) by Cuartes Owen

WarEruHouse and edited by Davin Suarp, Editor of the ‘ Zoological

Record.’ London, 1902. Price to Fellows, 18s.; price to the

public, 20s., or if sold with a set of the ‘ Zoological Record,’ 10s.

Index Zoologicus, No II. An alphabetical lst of names ofj

genera and subgenera proposed for use in Zoology, as recorded inf

the ‘ Zoological Record,’ Vols. 38-47 inclusive (1901-1910), and

the Zoology volumes of the ‘ International Catalogue of Scientific]

Literature, Annual Issues 1-10. Compiled (for the Zoological}

Society of London) by Cuartes OwEn Waveruovuse, I.8.0., and

edited by Davin Suarp, M.A., F.R.S., Editor of the ‘ Zoological}

Record.’ London, 1912. Price to Fellows, 12s. 6d. net; price

to the public, 15s. net., or if sold with a set of the ‘ Zoological}

Record,’ 10s. :

Divisions of Vols. 39 to 42 of the ‘ Zoological Record’ can be

supplied by the Society, but those of Vol. 43 onwards can be had only

of Messrs. Harrison & Sons, 46 St. Martin’s Lane, W.C.

[e. 2. 0.

SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.

Divisions of the ‘Zoological Record,’ Vols. 39-42, containing

the literature of the years 1902-1905, may be obtained separately

as follows :—

B Ge

List of abbreviations of journals, ete. 2 O net

Special Records, viz. :—

I. General Subjects . 2 Ones

II. Mammalia Nel One

III. Aves AS ia... G00 Ns

IV. Reptilia and Batrachia. . BD) pp

V. Pisces ZO,

VI. Tunicata WO ea

VIL. Mollusca AiO! ois

VIII. Brachiopoda . . ep Owes

IX. Bryozoa Re wO tr

X. Crustacea QeiGiatta.

XI. Arachnida OI ae

XII. Myriopoda Le Gate

XIII. Insecta We Oo

XIV. Echinoderma a Og

XV. Vermes .. 5 BO 5

XVI. Coelenterata .. iby AG, 35

XVII. Spongie PO

XVIII. Protozoa QU Cre

Index of new names of genera and subgenera. 2 O

Divisions from Vol. 43 onwards are now supplied

Harrison & Sons, 46 St. Martin’s Lane, London, W.C.

- Messrs.

P. CHALMERS MITCHELL,

Secretary.

Recents Park, Lonpon, N.W.

December, 1913.

LOOLOGICAE SO@ETY OF LONDON.

LIST OF PUBLICATIONS.

Tue scientific publications of the Zoological Society of London

are of two kinds—“ Proceedings,” published in an octavo

form, and “ Transactions,” in quarto.

According to the present arrangements, the “ Proceedings

contain not only notices of all business transacted at the scien-

tific meetings, but also all the papers read at such meetings

and recommended to be published in the “‘ Proceedings ” by

the Committee of Publication. A large number of coloured

plates and engravings are issued in the volumes of the

“ Proceedings,” to illustrate the new or otherwise remark-

able species of animals described therein. Amongst such

illustrations, figures of the new or rare species acquired in a

living state for the Society’s Gardens are often given.

The “ Proceedings” for each year are issued in four parts,

paged consecutively, during the months of March, June,

September, and December. From January 1901 they have

been issued as two half-yearly volumes, indexed separately.

An “ Abstract of the Proceedings ”’ is published by the

Society on the Tuesday following the date of the Scientific

Meeting to which it refers. It is issued along with the ‘“* Pro-

ceedings,” free of extra charge, to all Fellows who subscribe te

the Publications, but it may be obtained on the day of publi-

cation at the price of Sixpence, or, if desired, sent post free

for the sum of Six Shillings per annum, payable in advance.

The “ Transactions” contaim such of the communications

made to the Scientific Meetings of the Society as, on account of

the nature of the plates required to illustrate them, are better

adapted for publication in the quarto form. They are issued

at irregular intervals.

Fellows and Corresponding Members, upon payment of

a Subscription of One Guinea before the day of the Anni-

versary Meeting, are entitled to receive the Society’s

Publications for the year. ‘They are likewise entitled to

purchase the Publications of the Society at 25 per cent. less

than the price charged to the Public. A further reduction

of 20 per cent. is “made upon purchases of Publications

issued prior to 1881, if they exceed the value of Five

Pounds.

Fellows also have the privilege of subscribing to the

Zoological Record for a sum of One Pound Ten Shillings

(which includes cost of delivery), payable on the Ist. of J uly

in each year; but this privilege is forfeited unless the

subscription be paid before the Ist. of December following.

The following is a complete list of the publications of the

Society already issued.

TRANSACTIONS* OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 19 vols. and Index. ate Briceto the,

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PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND

CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF

LONDON. 8vo. 2 vols. (Letterpress only). rice to ibis uo ule

Fellows. Public.

Tears Jl, USROSil, Ibwoll, GwOs cbceddcccscec0 AS, Bh sooo GBsr

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PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.

8vo. 15 vols. (Letterpress only) and Index. (First Series.)

Price to Price to the Price to Price to the

Fellows. Publi. Fellows. Public.

Part I. 1883.1 vol. 8vo. 4s. 6d. .. 6s. | Part. 1X. 1841.1 vol. 8vo. 4s. 6d. .. 6s.+

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VII 1840. | 4s, 6d, .. 6st | Index 1880-1847, | 4s. 6a, | 6s,

8vo. 13 vols. an Index. (Second Series.)

% 4s. 6d. .. 6s.

o ASNOdaa esa

Letterpress only. With Plates coloured.

Price to Price to the Price to Price to the

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Part WIL 1B4Q, I soll, Bo, 4 Ot 5650 8s soocccce sl @ 8 see & Gp

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Tndex 1848-1860. i As. 6d. Gs.

t Out of print.

* In consequence of a re-arrangement of the stock of the ‘Transactions,’ the Society is

now able to offer for sale, at the reduced price of £30, sets of Vols. v.—xvi. inclusive, and

separate papers, of which a list can be supplied, at about one-fourth their published price.

PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE

ZOOLOGICAL SOCIETY OF LONDON. 8vo. 40 vols. and 4 Indices.

Letterpress only. With Plates uncoloured. With Plates coloured.

Price to Price to the Price to Price to the Price to Price to the

Fellows. Public. Fellows, Public. Fellows. Public.

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* No perfect copies in stock. t Out of print.

PROCEEDINGS or tue GENERAL MEETINGS ror SCIENTIFIC

BUSINESS or toe ZOOLOGICAL SOCIETY OF LONDON.

Svo. 25 vols. and Index.

Price to Price to the

Fellows. Public.

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ef oy. Juli 3 mSdio GH ORO clos oy SO o.ube 0.40.6 b5.0000 chee abo 4k

LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS.

List of the Vertebrated Animals now or lately Living in the Gardens

of the Zoological Society of London. (Highth Edition.) 8vo.

1883. Cloth, 4s. 6d.

List of the Vertebrated Animals now or lately Living in the Gardens

of the Zoological Society of London. (Ninth Edition.) 8vyo.

1896. Cloth, 6s.

Catalogue of the Library of the Zoological Society of London

(Fifth Edition.) 8vo. 1902. Cloth, 6s.

THE OFFICIAL ILLUSTRATED GARDEN GUIDE—11th Edition

—with (1) a Railway and Street Map, showing a direct

route to the ‘‘ Zoo” from all parts of London and the Suburbs ;

(2) a Plan of the Grounds, showing at a glance the location of

the animals; (3) a short description of some of the principal

animals in the Collection (now containing over 3000 spe-

cimeus), together with 50 Photographic Illustrations and

Index. Price 6d. in Stiff Paper Cover, postage 14d., or in

Green Cloth Cover price 1s. 2d. post free.

P. CHALMERS MITCHELL,

Secretary.

Regent's Park, London, N.W.

December, 1913.

These publications may be obtained at the Socinry’s OFFIce,

or through any bookseller.

ON AVIAN CESTODES. 861

49. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea. By Frank HE. Bepparp, M.A.,

D.Sc., F.R.S., F.Z.S., Prosector to the Society.

[Received August 27, 1913: Read October 28, 1913. }

XI. On A New TAPEWORM FROM GQ/DICNEMUS.

(Text-figures 141-149.)

INDEX.

Page

EHugonodeum edicnemi, gen. eb sp. mn. ............ Saw cette eS Ol

Characters of Hugonodeuwm ...... A is NE as een eae Sal eS) 7(7

[DiREOMVMMES Ce OOCMOPIRIIAE sononucansasetcnteectsboseecacdseuedcanooscs ov/4t

(GSMS OQ OSAAAMMONS oocaccéscnceasopsccocss sos ssbaenoaseesmssbonssercoovecca. ot

In Prof. Fuhrmann’s exhaustive list of Tapeworms * (of the

Cyclophyllidea only) which occur in birds, only one parasite

of this order is recorded from an Mdicnemus. The species in

question is Choanotenia coronata, and it occurs in the Kuropean

“dicnemus edicnemus. 1 was, therefore, particularly interested

to find in a New World species of the genus, viz. Gdicnemus

bistriatus, a number of examples of a tapeworm which does not

belong to the genus Choanotewnia, though it should probably be

referred to the same family and even subfamily.

I shall, however, defer the consideration of the systematic

position of this Cestode until an account has been given of its

anatomical characters.

The species dealt with in the present communication was

obtained from a Thick-knee of the species mentioned above, which

died in April of the present year, and had lived in the Society’s

Gardens fora year. It is not unlikely, therefore, that the tape-

worms had infected the bird hefore its arrival. But no fact

of importance can be at present deduced as to locality, since the

very parasite of Gdicnemus owdicnemus mentioned above has

been also discovered in Wgialitis nivosa, which is an American

bird.

The @dicnemus bistriatus contained no other tapeworms in its

intestine, except that which forms the subject of the present

communication to the Society. JI am not quite certain how

many individuals there were; but there were at any rate six or

seven. The worms are somewhat slender and measure at least

120 mm. I infer this from the fact that one of the largest

fragments, consisting of both ripe and unripe proglottids, measured

110 mm. ; but, as it had no scolex nor trace of the thin neck, at

least 10 mm. may be added safely. The greatest breadth of the

* “ Die Cestoden der Vogel,” Zool. Jahrb. Suppl.-Bd. x., 1908.

Proc. Zoou. Soc.—1913, No. LVIII. 58

862 DR. F. E. BEDDARD ON

ripe protoglottids is 2 mm. The accompanying figure (text-

fig. 141) shows the scolex and anterior end of an example of this

worm, as well as a portion of the posterior region of the same or

of another example. The anterior part of the body shows the

Text-fig. 141.

Kugonodeum cedicnemi.

Portions of two examples about twice nat. size.

The left-hand figure shows the rather large scolex and whiplash-like anterior

part of strobila. he right-hand figure consists of mature segment».

rather long and oval scolex followed by a slender neck which

widens out rather suddenly. Only one specimen out of the six

or seven which I have examined showed a gradual increase in

AVIAN CESTODES. 863

thickness from the neck onwards. This characteristic of the

species is not unfrequently met with in tapeworms; it is,

however, perhaps unusually strongly marked in this worm from

“dicnemus bistriatus.

The scolew is long and oval, and provided with four large

suckers which possess no armature of hooks.

When the scolex and the rostellum are extended the suckers are

quite in contact laterally. They are then oval in form. The

rostellum is long and retractile; when fully extended it is as long

as the rest of the scolex. The extremity is entirely muscular

with several layers of stout fibres running both transversely and

longitudinally. But in spite of its great muscularity there are

no hooks. It has been recorded, and I myself have had the

opportunity of observing a case, that among tapeworms which

possess an armed rostellum, the hooks are occasionally absent.

But in the present species I have never found them in a single

individual. J am thus disposed to think that they are really

absent, and to rely upon this as one of the distinguishing

characters of the genus, which will be duly set forth later *.

Behind the scolex there is quite a distinct neck, in which there

is no trace of segmentation to be observed. The segments of

the body overlap in the usual fashion and are never greatly

elongated. The most posterior are only slightly longer than broad.

The last external character to which I draw attention is the

position of the genital pores, which are unilateral. In the ripe

and more elongated segments they lie towards the hinder margin

of each proglottid. These orifices are strictly lateral. In the

narrow anterior segments the genital pores have a peculiar dis-

position. In horizontal sections through a consecutive series of

such proglottids the edges of the proglottids stand out for a con-

siderable distance laterally. The genital pores open on these

extensions, but on to the anterior margin of each where it runs

parallel with the posterior margin of the lateral extension of the

proglottid in front. The orifices are thus completely concealed on

a lateral view. In more mature segments these lateral extensions

do not stand out in the same straight line with the rest of the

segment, but become bent backwards; so that the anterior margin

becomes lateral.

The structure of a proglottid is illustrated in text-fig. 142

(p. 864). The cortical layer is about as thick as the medullary

layer. The former contains two rows of bundles of longitudinal

muscular fibres; the number of fibres constituting each of the inner-

most bundles is greater than that of those of the outer. There

appear to be a larger number of bundles also in the inner row.

These longitudinal muscle-bundles do not extend outside of the

nerve-cord ; they are not at all closely arranged. The water-

vascular system consists of two lateral tubes on each side, of which

the outermost, or ventral, is several times larger in cross-section

* Vide infra, }. 877.

864 DR. F. E. BEDDARD ON

than the innermost, which I take to be the dorsal vessel. The

tubes run nearly parallel to each other; but the dorsal vessel is

slightly to the dorsal side of the proglottid. The ventral vessels

are connected in each proglottid by a transverse vessel which is

shown (in part) in text-fig. 143.

Text-fig. 142.

Part of a transverse section through a proglottid of Hugonodeum cedicnemi, to show

the arrangement of the water-vascular tubes and cf the longitudinal muscles.

d. Dorsal vessel. m. Longitudinal muscles. #. Nerve-cord. v. Ventral vessel.

The ovary of this tapeworm is single and not divisible into two

lobes, as is so often the case. It lies distinctly to the pore side of

each proglottid median of the large (ventral) water-vascular tube

and partly ventral, as well as extending to both sides of the

dorsal water-vascular tube. It les in each case anteriorly in the

proglottid, and not very far behind the transverse water-vessel

of the proglottid in front. In front of the ovary, however, lie

some of the coils of the sperm-duct; it is not, therefore, quite at

the anterior margin of the segment. The vitelline gland lies

behind the ovary and towards its inner (median) side. The

relationship is shown in the accompanying drawing (text-fig. 143).

AVIAN CESTODES. 865

Tt will also be seen, on an inspection of that figure, that the

fully grown ova ave largely grouped together in cavities, and that

the ovary is more or less hollow. J am not, however, disposed

to think that this fact has any morphological significance. Two

interpretations are, of course, conceivable. Firstly, that the

Space is celomic, the eggs having been freed from its walls into

its cavity. The occurrence of a similar cavity in other tape-

worms may perhaps be an argument in favour of this view, which

is by no means impossible (and, indeed, has been theoretically

demanded by Bergh and others) ; yet I am disposed to regard the

Text-fig. 143.

\ es Vv.

Part of a horizontal section through a proglottid of Hugonodeum cedicnemi.

e.s. Egg-sacs. J.v. Ventral water-vascular tube. sp.d. A part of ccil of sperm-duct

lying in front of ovary. ¢. Testes. ¢.v. Transverse water-vascular tube.

v. Vitelline gland ; in front of this is the ovary, of which the darkly stained

mature ova are partly received within cavities of the parenchyma.

cavities as merely due to shrinkage. These hollows might also

be considered as the commencement of the uterus. The position,

however, would be rather abnormal; and, moreover, as again

is insisted upon later in this paper, ripe ova are already scattered

through the parenchyma, the cavities surrounding which can

therefore have nothing to do with the cavity in the ovary, what-

ever may be its nature. In younger stages than that which has

just been described the ovary forms a quite solid mass. Later

on it seems to disappear as a definite structure.

866 DR. F. E. BEDDARD ON

S Genital Ducts.

This worm is characterized by the great depth of the genital

atrium, which is much more developed than in most tapeworms

and quite as much so asin any. ‘The orifices of these follow in

successive proglottids upon the same side of the body. In the

ripe proglottids they are situated quite posteriorly and, therefore,

the genital ducts which open into them run forwards towards the

anterior region of the proglottid. The genital atrium, or genital

Text-fig. 144.

Part of a horizontal section through a proglottid of Eugonodeum cedicnemi,

showing a generative aperture.

c. Cirrus extruded from cirrus-sac (shown lying in front of it) and received within

vagina (v;). e.g. Very deep cloaca genitalis. sp.d. A coil of sperm-duct.

cloaca, is lined by a prolongation of the outer cuticle; outside of

this is a very thick layer of rather stout muscle-fibres which,

when the genital cloaca is cut through transversely to its axis,

are seen to have a radial arrangement. The cavity is circular in

transverse and oblong (more or less) in longitudinal section. At

the base, where the genital ducts join it, it widens out and forms

AVIAN CESTODES. 867

a circular cavity projecting beyond the rest of the genital cloaca.

The general shape is suggestive of a silk hat, if the brim of

the hat were hollow. From the centre of the lumen of the

genital cloaca a slender tube, which from its structure appears to

be morphologicaliy a part of the genital cloaca, leads to the

junction of the male and female ducts. This is shown in text-

fig. 144. It should be noted that the muscular layer of the

cloaca genitalis 1s covered externally by a layer of subcuticular

cells continuous with those which underlie the body cuticle. It

is, therefore, perhaps to be presumed that the muscular layer in

question is a local thickening of the delicate layer of fibres which

underlies the body cuticle. Occasionally I have observed the

cloaca genitalis to be slightly protruded ; in such cases the cavity

was more cup-like than cylindrical owing, of course, to the gaping

of the external orifice.

The sperm-duct and cirrus-sac, as already mentioned, pass

obliquely forwards in the mature segments and are parallel to the

vagina, which follows an identical course. ‘The cirrus-sac in this

worm is large and directed in a straight line towards its opening

into a chamber in common with the vagina. This latter chamber

has nothing to do with the terminal cloaca genitalis from which

it is sharply marked off by the muscular walls of the cloaca,

which have a narrow. tube of intercommunication, doubtless

capable of being widened. In all the individuals which I ex-

amined the cirrus of the fully mature proglottids was largely

protruded from the cirrus-sac, but not through the cloaca

genitalis to the exterior of the body. I found the cirrus, in fact,

to be invariably inserted into the neighbouring vagina, which

latter was as invariably filled with spermatozoa. here is, of

course, nothing new in this record of self-fertilization, which is

well known to occur among Cestodes, and has been recorded by

van Beneden, Leuckart, and others. A large number of instances

are given in Bronn’s ‘ Klassen und Ordnungen des Thierreichs ’

by Prof. Max Braun*. I have not, however, noticed it myself

in the considerable number of species which I have examined,

excepting in the present species. This auto-copulation is there-

fore far from being universal in occurrence. It appears to me,

furthermore, that the structure of the efferent apparatus in the

Cestode, which forms the subject of the present communication,

may at times necessitate this auto-copulation. A closure of the

cloaca genitalis coinciding with the eversion and protrusion of the

cirrus would force the latter into the vagina, which is widened at

its extremity, and does not project into the common chamber

into which both efferent ducts open t. The passage for the cirrus

is therefore not in any way hampered.

The cirrus-sac has thickened walls, as in so many species, and

* Bd. iv. Abth. 1, p. 1462.

+ v. Janicki in describing Schizotenia haymanni (Zeitschr. wiss. Zool. 1xxxi.

1906, p. 585), where there is also a very deep cloaca genitalis, comments on the

mechanical necessity for auto-copulation on rather different grounds.

868 DR. F. E. BEDDARD ON

is somewhat bottle-shaped. ‘The narrow region, however,—the

neck of the bottle—is not anterior where the cirrus-sac opens into

the common genital atrium, but posteriorly where the sperm-

duet opens into it. I have not always seen this narrow backward

prolongation of the cirrus-sac ; but I have seen it in more than one

case. There is no difference in its structure from that of the

main part of the cirrus-sac*. The cirrus, as usual, lies coiled

Text-fig. 145.

e. m. ue

Part of a horizontal section through a proglottid of Eugonodeum cedienemi,

to show further course of vagina.

e. Egg-sacs. m. Longitudinal muscle-fibres. 2. Nerve-cord. v. Vagina, full of

sperm and widening above to form a receptaculum seminis, thence bending

hack again to divide into the two usual branches o. & V7it., which are arranged

in the same straight line with each other.

within the sac and when protruded, as shown in text-fig. 144, is

broader at the protruded end. This is inserted into the vagina,

the proximal dilatation of which it completely fills, as is shown in

the same figure. The sperm-duct, after leaving the cirrus-sac,

runs in a straight or slightly sinuous course, at first quite parallel

* Tt is possibly temporary, and due to nnequal contraction.

AVIAN GESTODES. 869

to the vagina on the anterior side of it. It does not expand any-

where into a vesicula seminalis, but forms a large coil. This coil

lies in the neighbourhood of the two water- vascular vessels.

The vagina opens behind the cirrus-sae and is wider, and with

more muscular walls in that section which lies close to the ex-

ternal pore than elsewhere. It is this region which receives the

cirrus during the auto-copulation referred to above. The tube

then becomes finer and passes parallel to the sperm-duct, as shown

in text-fig. 144. At about the level of the larger, ventral, water-

vascular tube it expands into a not very large receptaculum seminis

(text-fig. 145), which in fully mature proglottids is gorged with

sperm. From this point in fully mature segments the vagina

then bends back again and divides into two tubes, one running

anteriorly and the other posteriorly. In less fully mature pro-

glottids the course of the whole vagina is straight and but slightly

oblique, being nearly parallel to the transverse axis of the body.

The shell-gland in such segments is very plain.

§ Egg Sacs.

The egg-containing cavities (which do not, as I think, collec-

tively represent a uterus) occur in segments which have already

begun to lengthen slightly, although they are still much broader

than long. In such segments the ovary and testes are fully

mature , but show no signs of degeneration. Several of these egg-

holding cavities are dis splayed i in text-fig. 143 (p. 865). They are

all small, but not of uniform size; their shape is quite uniformly

spherical. There is no regularity of arrangement among them

that I can detect; they lie everywhere in the proglottids, even

among the bundles of longitudinal muscle-fibres; they thus

axe into the cortex, a position which is, however, not unknown

in other tapeworms, though it is not common. There is no

connection to be observed between adjacent ege-cavities, though

they may he in actual contact: that is to say, there is no question

of a network—the cavities are totally independent in fact and, as

I shall point out, probably so in origin.

These egg- containing cavities, he largest of which are smaller

than the teste es, are definitely marked off from the parenchyma, in

which they lie, by a thickish wall. The cavity which lodges the

ege@* and the vitelline cells is naturally more apparent in the case

of the larger sacs. In the smaller ones the egg and accompanying

eells fill up the available space completely or nearly completely.

The inference appears to me to be that the cavity is formed later

perhaps by the exudation of fluid as well as growth of the peri-

phery in a way similar to that of the mammalian Graaftian follicle.

On the other hand, the structure of the walls of these egg-

containing cavities suggests another interpretation. As already

mentioned, they are rather thick and thus very conspicuous: in

* Occasionally two eggs are found in the same cavity.

870, DR. F. E. BEDDARD ON

them occur numerous nuclei sometimes close together and some-

times more sparsely arranged. These nuclei are more numerous

in the larger egg-holding spheres, and would thus seem to have

multiplied during a growth. But it must be admitted that the

existence of sacs with the lining apparently epithelial is highly

suggestive of a state of affairs like that shown by Dipylidium

caninum.

Text-fig. 146.

A horizontal section through mature proglottid of Hugonodeum ceedicnemi.

A, Advanced embryo in egg-holding cavity. B. Less advanced embryo in sinaller

cavity. v. Ventral water-vascular vessel giving off transverse vessel.

In this worm, as is well known*, the uterus finally breaks up

into numerous quite separate cavities, 1n which he many ripe

ova, and which are lined by a continuous epithelium. The

arguments against this supposition, however, outweigh those in

* Of. e.g. Beddard, P. Z.S. 1918, p. 555 text-fig. 88.

AYIAN CESTODES. 871

its favour. In the first place there is no trace that I can discover

of a pre-existent uterus*, whose subsequent fragmentation might

produce the result described above. Furthermore, the actual

cavities of the egg-holding apparatus are a later development, or

at any rate they increase in extent as the eggs within them grow.

Indeed it may, I think, be safely asserted that in the youngest

Text-fig. 147.

Part of the section illustrated in text-fig. 146 more highly magnified and showing

younger egg-sacs in parenchyma (e.s.) and testes (¢.), which are very much

larger.

groups of egg and surrounding cells there is no free space at all.

Nevertheless it might be said that all this was due to precocious

development, that in fact the rapid protrusion of ripe eggs from

the ovary had outstripped the growth of the uterus, which in

consequence appeared subsequently in point of time, and that

* Of., however, p. 865 under description of ovary.

872 DR. F. E. BEDDARD ON

the uterus for this latter reason appeared in an already divided

condition.

It is to be noted that egg-sacs of the character just described

occur in the younger proglottids, and that they are also to be

found (text-fig. 145, p. 865) in quite young proglottids, in which

the ovary and vitelline gland are at their full development and

have not commenced to degenerate. Without having any positive

proof, | am disposed to think that the cells which encircle the

ovum are ovarian or vitelline gland-cells*, which come to be dis-

posed round the ovum after the fashion of a follicular epithelium

in higher animals. It looks to me, in fact, very much as if a ripe

ovum with a few adherent cells moved out into the adjacent

parenchyma directly from the ovary. There is here clearly a

similarity with the Graattian follicle of Vertebrates, a similarity

which is even increased by the later appearance of a space (perhaps

containing fluid) which surrounds the growing ovum.

It does not always follow that a proglottid which has attained

the dimensions and acquired the appearance of a fully mature

proglottid should contain only embryos. On the contrary, as

be readily seen from an inspection of text-fig. 146 (p. 870),

a large flat proglottid, quite as large as it will grow, may have

egg-sacs 1n many stages of growth. In this figure, which is

dvawn from a section seen under a low power, the size of the

egg-sacs 18 a rough measure of their relative stages of growth.

The largest are of course the oldest, and contain fully developed

embryos surrounded by a thick shell and enclosed in rather a

large space. Intermediate conditions are to be found between

this stage and that in which an undivided ovum occupies all the

space of the cavity of the body-parenchyma in which it lies. I

have also cut sections of apparently fully mature proglottids, in

which there were no advanced embryos.

This state of affairs is not unlike what has been described in

some other Tapeworms by others as well as by myself. We may

exclude Monopylidiwm and Dipylidium, which only show an

apparent likeness to Hugonodeum. In the former genus, and in

certain species belonging to the latter genus, the inclusion of the

ripe ova singly in compartments of the medullary parenchyma

is preceded by a stage in which a functional uterus exists or

(Dipylidium, various spp.) at any rate a cavity which—full of

eggs——is ultimately broken up into: single compartments. On

the other hand, there is a much greater likeness to Hugonodeum

in a genus which I have lately described as new under the name

of Diplopylidium*. Here we have, as it would appear, no trace

of a uterus; but the eggs are found scattered widely through

the medullary parenchyma enclosed in a cavity singly ; the cavity

grows in correspondence with the growth of the contained

* (Of. description of ovary and vitelline gland above.

+ P.Z.S. 1913, p. 565, text-fig. 93, p. 566, text-fig. 94.

AVIAN CESTODES. 873

embryo. There is no great difference in size between the egg-

holding cavities in the two genera. But there is an important

difference in the structure of the wall of these cavities in Diplo-

pylidium, and in the presently described genus Hugonodeum. In

Diplopylidiwm 1 was unable to find any nuclei lining the cavities,

which are simple excavations in the medullary tissue. This was

Text-fig. 148.

Another part of the same section of Hugonodewn cedicnemi showing older embryos

surrounded by a greater egg-holding cavity in the medullary parenchyma.

A, B. Egg surrounded by thick shell and lying in cavity from which nutritive

cells (?), such as are shown at es. in text-figure 147, have disappeared.

C. Older embryo with larger space surrounding it. D. A degenerating (?)

egg-holding cavity and embryo.

the case, not only with the fully mature embryos, but also in com-

paratively newly formed cavities. In Hugonodewm, on the other

hand, the egg-holding spaces possess a lining of cells. It is true

that in fully mature embryo-containing cavities these cells are

874 DR. F. E. BEDDARD ON

often, perhaps generally, indistinguishable. But a granular

detritus often visible seems to me to indicate that their tendency

to disappear is due to the fact that they serve as nutriment for

the growing ovum. It may be noted, furthermore, that the egg-

shell in Lugonodewm is not nearly so broad as in Diplopylidium.

A reseinblance exists between the disposition of the ripe ova

in the present genus and that which characterizes Oochoristica.

I have re-examined examples of a species of Oochoristica which I

described lately to the Society as a near ally of, if not identical

with, Oochoristica wageneri of Janicki*. In the course of my

account of that species I figured the scattered condition of the

ripe eggst, well known from the investigations of others to he

one of the characters of this genus. I did not, however, attempt

any details, merely showing the inclusion of the ripe embryos

eich im a separate chamber, and these scattered through the

general medullary parenchyma. I have now some details to add

to that account. The accompanying figure (text-tig. 149) shows a

part of a section through one of the last two proglottids of

specimen of this species. It will be seen that the eggs are more

closely packed than in Mugonodeum. Vhey are not, however, in

contact or enclosed in one cavity. This much concerning the

ege-sacs of Oochoristica 1s well known from the observations of

several writers previously to myself. e. gy. Zschokke and v. Janick1.

But neither Zschokke{ nor v. Janicki§ give any detailed figures

in their memoirs dealing with Oochoristica of the structure of the

scattered eggs or oncospheres, or of the spaces which they oceupy

in the parenchyma. Zschokke describes the three membranes

which surround the hexacanth embryos, and observes of them

(the embryos) that they ‘“ liegen je einzeln in dichtgedringten,

rundlichen, kapselartigen Follikeln des Parenchyms.” Goin)

figures the ‘crowded embryos with their three surrounding mem-

branes entirely filling the proglottids at the end of the body ;

but he also does not enter into various histological points which

are, as I think, of interest in comparing the scattered egg-capsules

of Oochoristica with those of other tapeworms.

In longitudinal sections of the entire ripe proglottid these egg-

capsules are seen to be circular or more oval in contour, and to

be larger or smaller in size with no regularity of arrangement.

The differences of size correspond as I think partly at least to

differences of age. The more or less circular contour may have

a relation to the plane of the sections. Contrary to the obser-

vations of Zschokke and Cohn upon the species studied by

themselves, I can see but one egg-membrane which occupies all

the available space in the egg-follicle. V. Janicki gives no

Zeitschr. wiss. Zool. lxxxi. 1906, p. 533.

P. Z.S. 1911, p. 633, text-fig. 150.

“Das Genus Oochoristica Lithe,”’ Zeitschy. wiss. Zool. Ixxxiu. 1906, p. 53.

Loe. cit.

Arch. f. Naturg. 1903.

tt &

AVIAN CESTODES. 875

description of the eggs of Oochoristica wagenert, which is nearest

to, if not identical with, my species, except to state that they are

provided with a shell. The sharply defined edge of the shell

differentiates the ovum or the cells of the embryo, if it has deve-

loped so far, which lie within it from a layer of cells which forms

a follicle, and is so far like that which has just been described in

Eugonodeum. There are, however, differences to be noted which

will be apparent from the two figures (text-figs. 148, 149) which

illustrate the two species, and which are drawn practically to the

same scale. The cells of the egg-follicle in Oochoristica have

Text-fig. 149.

A section through a portion of the medullary parenchyma of Oochoristica, showing

five eggs or developing embryos. p. One pole of the usually elliptical

embryo-sac.

nuclei of a different appearance and form a thicker layer in that

the nuclei are more abundant and closer together. Furthermore,

this cellular layer is just as well developed in the largest as

in the smallest follicle. The cells do not disappear during the

growth of the contained embryo, as would appear to be the case

with Hugonodeum. It is easy to believe, therefore, in accordance

with the opinion of the authors quoted above, that the egg-sacs

in Oochoristica are portions of a subdivided uterus. I have,

however, myself no facts to offer in confirmation of this view.

876 DR. F. E. BEDDARD ON

In many of the ege-follicles the form of the entire follicle is, as

already stated, oval. In these cases, which are illustrated in text-

fig. 149, the layer of cells extends at the two poles of the ellipse

for a considerable distance outwards, and thus a heap of cells is

formed. hese masses of cells generally enclose a distinct cavity,

the function of which arrangement may be to protect the egg

from the effect of pressure due to contortions of the worm’s body.

In any case we have here a state of affairs very unlike that of

Kugonodeum. It appears to me to be very possible to make a

comparison between the egg-sacs of Oochoristica and the cor-

responding structures in two species of Davainea studied by

v. Janicki®. Referring, for example, to fig. 9 of his memoir, the

space lying between the two “ Embryonalhiille” at the poles

of the elongated embryo might well correspond to the cavity

which I describe and figure here in Oochoristica. But this com-

parison, which may be carried a good deal further, is not germane

to the object of my present communication, which is to compare

Hugonodeum with other allied genera.

S General Observations.

On the whole the characters of the present species ally it most

nearly to the genus Monopylidium, the alliance being very largely

due to one species only of that genus, viz. Monopylidiwm rostel-

latum. The genus Monopylidium is thus defined by Ransom }:—

(1) Rostellum armed with a double or single crown of hooks.

(2) A single set of reproductive organs in “each segment.

(3) Genital pores irregularly alternate, rarely unilateral:

(4) Genital canals pass between the longitudinal excretory

vessels and dorsal of the longitudinal nerve, or dorsal of

both excretory vessels.

(5) Testicles numerous (20 to 40 or more), behind the femaie

glands or, also, on both sides of the latter.

(6) Vas deferens coiled, seminal vesicle absent.

(7) Uterus breaks down into egg-capsules, each containing one

or several eggs.

The worm which forms the subject of the present paper differs

from Monopylidium in Nos. (1) and (7), and agrees with the

genus in the other characters set forth. These ditlerences appear

to be quite enough for generic separation were it not for the

structure of Monopylidium rostellatum. This species has no

rostellar hooks, and of the uterus or rather the disposition of the

ripe eggs Prof. Fuhrmann writes: “ Les oncosphéres se trouvent

réparties dans tout le parenchyme: chacune delles est entourée

* “Ueber zwei neue Arten des Genus Davainea,” Arch. de Parasitologie, vi. 1902,

p. 257.

+ Bull. U.S. Nat. Mus. No. 69, 1909, p. 76.

{¢ “ Nouveaux taenias @Oiseaux,”’ Rev. Suisse Zool. xvi. 1908, p. 65.

AVIAN CESTODES. 877

par un parenchyme vacuolaire, qui forme autour de Poncospheére

des cellules hexagonales d’un diametre d’environ 0". 17. Les

proglottis murs ressemblent ainsi beaucoup a ceux de certains

Davainea & capsules utérines ne contenant quun ceuf.” The

difficultyin comparing my genus with this particular Wonopylidium

is that Fuhrmann says very little of a pre-existing uterus.

He merely observes that quite early in the body ‘“ Vutérus

commence déja 4 se former,” and that further back ‘“‘ les segments

sont remplis Veeufs.” Iam disposed, however, to think that this

implies a likeness to the more typical species—such as, for example,

M. unicoronata, which Fuhrmann describes immediately after

Healing with WV. rostellatum. Moreover, in his résumé of Avian

lestodes, Furhmann™* defines the genus Monopylidium by, inter

ae the following character :-—“ Uterus stark verzweigt, lost sich

in Parenchymkapseln auf,” etc. As I. rostellatum had been already

described by him it is to be inferred that this species agrees with

the others, and is, therefore, so far not like the Teeniid which

forms the subject of my present communication to the Society.

We may, therefore, I think, exclude my species from the genus

Monopylidium. Nor does it appear to me that we can refer it

to any other known species. I propose, therefore, to name and

characterize a new genus as follows :—

Eugonodeum, gen. nov.

Rostellum very muscular, long and retractile, without hooks.

Suckers unarmed. Ripe proglottids never much longer than broad.

Dorsal excretory tube much smaller than ventral, lying more or

less laterally to it ; ventral vessels connected by transverse vessels in

each proglottid. Longitudinal muscles in two rows of bundles.

Genital pores unilateral. Genital ducts pass between excretory

tubes. Testes chiefly behind ovary, and then not very numerous.

Ovary to pore side, in front of vitelline gland. Genital atriwm

very deep, with radiating muscles. Cirrus-suc large and muscular,

lying in front of vagina. Sperm-duct with coil. Cuwrus unarmed.

Receptaculum seminis present, but not strongly marked. Uterus

not present. Ova imbedded singly in parenchyma accompanied by

other cells; round each ovum a cavity is formed later which is

lined by cells.

I name as the type Hugonodeum edicnemi, sp. n., with the

characters of the genus.

* “Die Cestoden der Vogel,” Zool. Jahrb. Suppl.-Bd. x. 1908, p. 65.

Proc. Zoou. Soc.—1913, No. LIX. 59

878 MR. R. W. PALMER ON THE

50. The Brain and Brain-Case of a Fossil Ungulate of the

Genus Anoplotherium. By R. W. Paumer, M.Sc.

(Manchester), Research Fellow in Zoology, University

College, Reading *.

[Received May 29, 1913: Read October 28, 1913. ]

(Text-figures 150-157.)

INDEX.

MorruoLoey. Page

Anoplotherium : Skull—description of ..................... 878

s Brain— A FY isan aan es auce tetshs)

SYSTEMATIC.

Anoplotherium related to Orycteropus ............... 886, 892

For some time a cranium from the Phosphorites of Quercy,

together with an exceptionally perfect and well-marked brain-

cast obtained from it, has been awaiting description in the British

Museum. The cast was submitted to Prof. G. Elliot Smith,

F.R.S., of Manchester, who intended to describe it, but he was

prevented from so doing by the pressure of other work, and very

generously passed it, together with the notes he had made on it,

to myself. By the courtesy of Drs. Smith Woodward and C. W. :

Andrews, of the British Museum, I have been allowed to describe

both the skull and brain-cast.

The whole of the facial region of the specimen is missing, and

since the dentition cf mammals forms such an excellent guide to

affinities that in original descriptions cranial features are for the

most part entirely overlooked, the identification of a toothless

skull such as this is a matter of difficulty. A study of the

literature of the subject and of such specimens as are available,

however, leaves little doubt as to the genus of the creature.

The skull is broken away at the back of the orbits, and the

paroccipital, zygomatic, and pterygoid processes are incomplete.

In dorsal view (text-fig. 150) it is seen to be narrow, elongate, and

somewhat fusiform. Its greatest width in front of the zygo-

matic processes is 49 mm., but behind the orbits it is greatly

constricted and its width reduced to less than one half, or 23 mm.

At the back of the zygomata, not including the lambdoid ridge,

it measures 40 mm. across. The marked postorbital constriction

is by no means uncommon in early Tertiary mammals, and

indicates at once a poor development of the frontal lobes of the

hemispheres.

The most striking feature of the dorsal surface of the cranium,

however, is the great strength of the sagittal and lambdoid crests.

The former arises in front by the union of the two low postorbital

ridges on the frontals, and traverses the whole length of the brain-

.* Communicated by Dr, C. W. Anprews, F.R.S., F.Z.S.

BRAIN OF A FOSSIL UNGULATE, 879

ease as a knife-like ridge which rises as high as 16 mm. above the

general surface. As it nears the occiput the crest somewhat

thickens and meets the equally highly developed and more massive

lambdoid crest at right angles.

The root of the zygomatic process is very extensive and slightly

convex antero-posteriorly. Its widely concave form in a trans-

verse direction indicates a temporal arcade standing well out from

the skull.

Text-fig. 150.

Dorsal view of skull of Anoplotheriwm.

fr., frontal; par., parietal; sq., squamosal.

In lateral view (text-fig. 151) the great height of the sagittal

and the true form of the lambdoid crests are displayed. This

latter strikes off at right angles to the sagittal for about 2 ems.,

slightly dropping in elevation as it does so, then falling very

sharply downwards, forwards, and slightly inwards, is con-

tinued as a much less pronounced ridge to the posterior root

of the zygoma. Here it bifurcates, the anterior branch being an

extension of the vertical flange of the zygomatic process, while

the posterior runs down, and slightly back, on to the huge par-

occipital process, on which it dies out. The aperture of the

external auditory meatus betweer. the paroccipital and zygomatic

processes is remarkable in being formed above by a groove in the

squamosal, while the tympanic forms its floor alone, there being

no ordinary bony tube.

In occipital view (text-fig. 152) the foramen magnum is seen to

be nearly circular and the condyles massive. Above, the lamb-

doidal crest forms a large shield-shaped area much roughened for

59*

880 MR. R. W. PALMER ON THE

Text-fig. 151.

SS SS ==

PAS

il) eam pg

Lateral view of skull of Anoplotherium.

ais., alisphenoid; e.a.m., external auditory meatus; j:ov., foramen ovale; gl.s.,

glenoid surface of the squamosal; m,f., mastoid foramen; m.per., mastoid

portion of periotic ; 0.s., orbitosphenoid; par., parietal; par.p., paroceipital

process ; p.g,f., post-glenoid foramen; p.g.p-, post-glenoid process; p.orb.p.,

post-orbital process; IT., optic foramen; III.—VL., united sphenoidal fissure

and foramen rotundum ; AIT., condylar foramen.

Text-fig. 152.

( i poy)

Occipital view of skull of Anoplotheriwn.

Lettering as in text-fig. 161,

BRAIN OF A FOSSIL UNGULATE. 881

muscular attachments. The paroccipital processes are seen to

diverge slightly below.

The palate (text-fig. 153) is very long and narrow. The glenoid

surfaces of the squamosals are large and flat with strong post-

glenoil processes. On the left side the tympanic has been

preserved, while on the right this is conveniently missing, leaving

Text-fig. 153.

Palatal view of skull of Anoplotheriwm.

Lettering as in text-fig. 151, with addition of 6o., basioccipital ; f£, venous foramen ? ;

flm., foramen lacerum medium; f/.p., foramen lacerum posterius ; per,

periotic; p.s., presphenoid; sy., squamosal; séin.f., position of stylomastoid

foramen; é.hy., pomt of attachment of tympano-hyal; f.y., tympanic; wf,

canals for vessels of the diploé.

882 MR. R, W. PALMER ON THE

the lower surface of the periotic exposed. The secondary palate

is completely lost.

In a longitudinal section of the skull (text-fig. 154) the great

thickness of the bones and the general development of diploé are

striking features. A skull which supported such massive tem-

poral muscles as this beast possessed had of necessity to be of a

strong build. The extreme shallowness of the pituitary fossa and

the practical absence of clinoid processes are features of interest,

while the mode of attachment of the cribriform mass to a groove

in the pre-sphenoid is curious. The pear-shaped cerebral fossa is

partly separated from the cerebellar by the heavy tentorial ridge,

Text-fig. 154.

ae Sie

eS Sis

Ya mieat

bo ue fim fou bs vi. bs,

Median section of skull of Anoplotherium.

Lettering as in preceding figures, with addition of 6.s., basisphenoid ; er.p., cribri-

form plate; f.fo., floccular fossa; v.d., canal for vessel of diploé.

most of which belongs to the squamosal. The spacious cavity for

the transverse snus opens immediately behind this ridge above

the periotic. It goes straight down to the post-glenoid foramen

and is joined by Bioanal for a vein which ran in a deep groove on

the side-wall of the skull, and was formed by the eonlinenes of

two vessels which ran in the substance of the alisphenoid.

The main features of the individual bones of the skull will now

be outlined.

The bones of the occipital segment are indistinguishably fused

together, but their anterior sutures are for the most part distinct.

The thick supraoccipital is of great extent owing to the extreme

development of the lambdoid crest, of which it forms the entire

BRAIN OF A FOSSIL UNGULATE. 883

upper portion. It articulates with the parietals by a jagged

transverse suture anterior to this crest. The exoccipitals were

also large bones, including, as they no doubt did, the large con-

dyles and practically the whole of the paroccipital processes.

They take no share in the lambdoid ridge, for their suture with

the mastoid and squamosal lies just posterior to this ridge. It is

a Jagged line, starting at the base of the steep drop in the crest,

and travelling behind it, runs down on to the paroccipital process.

The basvoccipital is clearly marked off from the basisphenoid by a

slightly irregular groove. It is very wide behind, between the

paroccipital processes, and notched by the condylar foramina

which must pierce the skull between it and the exoccipitals (text-

fig. 153, p. 881). In front it is narrowed by the lacerate foramina

and by the gap in which the periotic is set. Its lower surface

is here heavily marked for muscular attachments. In section,

the bone is seen to be thin and dense posteriorly, but in front it

is thicker and more cancellous in structure (text-fig. 154).

The parietals are remarkably extensive, and externally they

appear to form the entire cranial roof. Anteriorly, they con-

siderably overlap the frontals, and the extent of this overlap can

be seen in section (text-fig. 154), where the junction of the bones

is indicated by a break in the structure of the roof (at “a” in the

figure). The parietals do not share in the formation of the lamb-

doid crest, but meet the supraoccipital anterior to it. There is no

trace of a distinct interparietal, and the strength of the sagittal

ridge has necessitated a firm union of the parietals. The suture

with the frontals is first seen as a jagged line on the postorbital

ridge. It travels forwards along this for about 12 mm., then

turns suddenly backward and curves down on the side of the

skull till it strikes the alisphenoid. Below, the parietal is

suturally connected with the alisphenoid along a short horizontal

line, and posteriorly it is considerably narrowed by its bow-shaped

suture with the squamosal (text-fig. 150, p. 879). Inside the

skull, the fronto-parietal suture 1s seen as a very irregular line

running vertically on the wall of the cerebral fossa, and the

squamoso-parietal suture as an even more irregular line in front

of the tentorium. Neither suture can be completely traced.

The sguamosal forms a considerable part of the side-wall of the

skull. Posteriorly it overgrows the mastoid, so that only a small

isolated surface of this bone is exposed. The suture with the

mastoid is anterior to the lambdoid crest, and as this bone gets

submerged the suture crosses the crest to form the junction with

the exoccipital. The squamosal sends a small vertical thin flange

on to the root of the paroccipital process. It is partly anchylosed

with the tympanic, and in front of the meatus is bored by the

very large post-glenoid foramen. Almost the whole length of the

bone is concerned in forming the zygomatic process, which is con-

cave from side to side above, has a vertical flange to the exterior,

and a flat glenoid surface below. The flange is roughened on its

upper edge for attachment of the temporal fascia. ‘T'o the inside

884 MR. R. W. PALMER ON THE

of the glenoid fossa, where the squamosal meets the alisphenoid,

there is a narrow ridge which both bones help to form, and which

runs parallel to the middle line to separate the glenoid and

tympanic cavities. Where this ridge meets the post-glenoid

process there is a depression which may represent a venous

foramen (text-fig. 153, /,, p. 881).

Seen from the inside the periotic is quite an extensive bone.

The double openings of the internal auditory meatus and of the

aqueductus Fallopii are seen in the centre of the lower triangular

part. Above this, the floccular fossa (/,fo.) is excavated, and

from this region the large mastoid portion inclines backwards.

The mastoid part, as mentioned, 1s exposed on the outside of the

skull only as a small island between the exoccipital and the

squamosal, forming the lambdoid crest for about 1-5 ems. Through

its suture with the exoccipital runs the small mastoid foramen

(mf.). he inferior surface of the petrosal portion is seen as a

subovate mass inclined steeply outwards and jammed in between

the basioccipital and squamosal. It separates the median and

posterior lacerate foramina. Posteriorly it is pierced by the

fenestre rotunda and ovalis.

The tympanic is preserved on the left side. It is partly

anchylosed to the squamosal, but not to the periotic. It consists

of a small, very thick, uninflated bulla which does not meet the

basioccipital, but projects freely under the petrosal. A cylin-

drical hollow on the bone represents the attachment of the

tympano-hyal (¢.hy.), and on the right side the position of the

stylomastoid foramen (sém.f:) is seen to be immediately behind

this. The rest of the bone forms an irregular mass below the

meatus and presents no points of interest.

The alisphenoids ave firmly fused with the basisphenoid, and

the three bones must be taken together. The basisphenoidal part

is very elongated —behind, it 1s wide and notched by the median

lacerate foramina. In front, it narrows very considerably, and

at the junction with the alisphenoidal part the vertical pterygoid

wings arise. The alisphenoidal part is a hatchet- or L-shaped

bone, having a narrow limb confined to the base of the skull

between the squamosal and the basisphenoid and a square limb

in front of this, forming part of the side wall of the skull and

meeting the parietal above and the frontal in front. ‘The long

limb is deeply grooved for the Kustachian tube, and in its middle

is perforated by the foramen ovale (fiov.). Anteriorly, below

the pterygoid flange, the bone is tunnelled by a canal which

represents the united sphenoidal fissure and foramen rotundum

(III.-VI.). A couple of venous foramina (v,f.) appear externally

on the alisphenoid. These were traversed by the vessels which

united inside the skull to form the large vein already mentioned

as running into the transverse sinus. On its upper surface the

alisphenoid is very deeply grooved to shelter the cranial nerves

lying on it. These grooves are better seen from the cast. They

will be mentioned later. In section, the basisphenoid is seen to

BRAIN OF A FOSSIL UNGULATE. 885

be a thick bone, and it is scarcely thinned by the shallow

pituitary fossa.

On the dorsal surface of the skull the frontals form a flat

arrowhead-shaped area between the post-orbital ridges, Laterally

they are concave. Below, they unite by suture with the ali- and

orbito-sphenoids, and they extend well down on the skull since

the palatines cannot be traced below them. Within the skull

they form the division between the olfactory and cerebral

cavities. In front and above they are hollowed out by the

frontal air-sinuses.

The orbitosphenoid is quite small. Its suture with the ali-

sphenoid is not plain, but it appears to completely surround the

optic foramen. Itis indistinguishably fused with the presphenoid,

which is a very long and thin bone with a marked median palatal

ridge. It is very thick, and its upper surface, as seen in section,

is Shaped like the roof of a house, that is, its surface is very high

in its middle length and slopes down both forwards and backwards

(text-fig. 154, p. 882). The posterior slope shares in the tunnel for

the optic nerve, while the anterior one is excavated by a rounded,

undercut hollow which gives a firm support to the cribriform

plate (cr.p.). This last bone i is a solid mass of perforated tissue

and presents no special points of interest.

Comparison of the specimen with the figures given by

Juvier [1] and Blainville [2], as well as with the casts available at

the British Museum, leaves little doubt but that the skull has been

correctly identified. In Cuvier’s figures the general form of the

skull with the postorbital constriction, the form of the condyles,

the immense paroccipital processes, the peculiar formation of the

meatus, the post-glenoid process, its foramen, and a host of other

details are identical. Fig. 1, pl. xlvi. of Cuvier’s 1822 edition

shows the upper surface of the base of the skull of an Anoplo-

theriwm determined from the teeth, and the arrangement of the

foramina, of the grooves for the nerves, and of the transverse

sinus agree precisely with this specimen. The agreement with

Blainville’s figures is no less conclusive, and though the crests on

the skull are considerably stronger than those in any specimen

figured by these writers, they are of the same form, and their

degree of development can hardly be of greater than specific

value when other features agree so markedly.

Support for the identification also comes from the brain-cast,

for it agrees essentially with the natural one figured by Blainville

on plate u., but not with the one given by Cuvier (plate lv.,

1822 edit.), which is incorrectly identified as an Anoplothere.

Brain-cast.

The narrow, elongate form of the skull is naturally also a

feature of the brain. Its total length from the front of the

olfactory bulbs to the caudal end of the cerebellum is a fraction

under 10 ems. Of this total, the cerebrum accounts for 5°6 cms.,

886 MR. R. W. PALMER ON THE

the cerebellum for 3°35 ems., and the olfactory bulbs for the

remaining | em. The eranial capacity, as measured with sand

from the cast, is 86 cc.

The parts of the cerebral hemispheres chiefly concerned with

the sense of smell, are, as usual in early Tertiary mammals,

highly developed. This is seen in the large size of the olfactory

bulbs and of the rest of the “smell-brain,” and in the small size

of the neopallium. The greatest width of the cerebrum, which

is near its caudal extremity, is 45 mm.

The cerebellum is separated from the hemispheres, but there is

no evidence that the corpora quadrigemina were exposed. Its

width cannot be measured accurately, owing to the casts of the

cavities for the transverse sinuses being continuous with its mass

on each side. Hstimating the extent of these sinuses, its trans-

verse is not much more than its antero-posterior diameter—that

is, about 37 mm.

Text-fig. 155.

/ 7

(

afb lef’

Lateral view of brain-cast of Anoplotherium.

fic f., floccular lobe; f-l.m., cast of foramen lacerum medium ; f.l.p., cast of foramen

lacerum posterius; f-rh., rhinal fissure; o1.b., olfactory bulb; s.coz., coronal

sulcus; s.orb., orbital sulcus; s.ss/., suprasylvian sulcus; v., vessel; IT.

IIT-V1., V3, VIL.-VILIL., casts of foramina of cranial nerves.

Looked at in side view (text-fig. 155), the cerebellum stands as

high above the axis of the brain as does the cerebrum. This is

no doubt a primitive feature, for in recent Ungulates the hemi-

spheres are distinctly more elevated than the cerebellum. The

Middle Kocene Palwosyops agrees with Anoplotherium in this

particular [3].

On the base of the brain the courses of the cranial nerves and

the casts of the lacerate foramina are conspicuous, while below

the floccular lobe on each side the periotic has left a ragged scar.

The brain, in general form, arrangement of parts, and furrows,

is remarkably similar to that of Orycterepus. ‘This is seen in side

view, and especially from below. If the figure of the ventral

BRAIN OF A FOSSIL UNGULATE. 887

surface of the brain of the Aard-Vark given by Elliot Smith

[4] be compared with text-fig. 156, the similarity will be seen to

be very striking. The detailed resemblances will be noticed below,

and they undoubtedly help to strengthen the evidence in favour

of aprimitive Ungulate origin for the Tubulidentata—a view held

by Elliot Smith, Max Weber, and Loinnberg.

Text-fig. 156.

fa!

4 Mi

\\ 4

\ y

AN nf

\\ y Gh

\W Wy? it /

iy \

Ventral view of brain-cast of Anoplotherium.

Lettering as in text-fig. 155, with addition of m.f., mastoid foramen ;

ol.tb., olfactory tubercle.

Ventral Surface of Cerebrum.

The whole of this region is well developed. This can be clearly

seen from text-fig. 156, for, seen from below, the pyriform lobes

almost completely conceal the neopallium—a part of which is

888 MR. R. W. PALMER ON THE

visible only to the outside of the anterior parts of the pyriform

lobes. <Anoplotheriwm, then, was far more highly macrosmatic

than any living Ungulate, and not much less so than the bur-

rowing Aard-Vark,

The olfactory bulbs at their widest part are together 21 mm.

across. The olfactory peduncles are distinct thick stalks, swelling

out behind into the anterior ends of the pyriform lobes.

The olfactory tubercles are well shown as smooth circular

elevations placed rather further back than in Orycteropus, and

sufficiently prominent to be noticeable in a side view of the

cast.

Between these tubercles the cast shows the course of the optic

nerves. These tunnelled the base of the skull very near the

middle line. The position of the chiasma is indicated by the

sinking of these tubular casts on to the general level of the brain-

surface. Diverging outwards from the chiasma, the vallecule

Sylvii are distinct, marking off the globular posterior parts of the

pyriform lobes. Running back along the lower surface of each

of these lobes is the cast of the vein described as running on the

side of the cranium. This vessel does not seem to be cerebral,

but rather to collect blood from the diploé of the bones. The

only other feature on the pyriform lobe worthy of note is a short

straight fissure running along it, above the vessel just mentioned,

and below and parallel to the rhinal fissure. A similar fissure

occurs in the pig and the tapir. In text-fig. 155 (p. 886) it

has been marked with the letter ‘‘ x.”

Lateral Surface of Cerebrum.

The olfactory bulbs having been described from below need no

further mention.

The neopallium is clearly marked off from the rest of the hemi-

spheres by the great horizontal rhinal fisswre. In text-fig. 155

there is a break in this fissure at about one-third of its total length

from the front. This break does not occur on the other side, and

seems to be due to damage of the skull-wall. In side view the

rhinal fissure 1s seen to divide the hemisphere horizontally into

two equal parts, and the primitive character of the brain is

emphasized by the relatively small size of the upper part (neo-

paliium) in comparison with the rest of the cerebrum. Its great

development in the higher mammals, which led to the degenera-

tion of the “smell-brain” and to its enwrapping by the

neopallium, has scarcely begun.

About 17 mm. from the anterior end of the hemisphere, there

is a vertical sulcus running into the rhinal fissure below, and

connected, on the right side only, by a shallow furrow with the

lateral sulcus. This sulcus can be no other than the orbital *. Its

* T am using the same terms as were employed in the descriptions of the brain of

Orycteropus [4], to which I have referred, although Dr. Elliot Smith tells me the

whole question of nomenclature of cerebral sulci urgently needs revising in the light

of recent research on anatomical localization of the cerebral cortex.

BRAIN OF A FOSSIL UNGULATE, 889

position, relation to the rhinal fissure, and its possible connection

on the right side to the lateral suleus ave exactly like the con-

ditions in the brain of Orycteropus.

Behind the orbital, the side of the hemisphere is marked by a

clearly defined sulcus, rather under 2 cms. in length, and running

obliquely from behind forwards and downwards to join the orbital.

This sulcus is nearly certainly the coronal. Its relations on the

two sides are similar, but while on the left it is simple, on the

right it forks slightly at its superior or dorsal end, and in its

middle length it is connected with avery short horizontal sulcus

(4), the identity of which is not clear.

There is no trace of a pseudosylvian sulcus on either side of

the brain.

Dorso-laterally the neopalltum is marked by a suleus which,

starting at the back of the hemisphere, runs straight forward,

parallel to the middle line, for about 2 cms., and then, curving

down towards the rhinal fissure, ends on the right side, about

7 mm. from that fissure, but on the left actually joins it, or very

nearly so. This must be the suprasylvian sulcus. Though small

and not so strongly arcuate as in the higher Ungulates, it is of

the typical Ungulate form in the strength of its posterior branch.

Near its caudal end the posterior limb of the suprasylvian is

joined by a very short sulcus (c), which lies external to it. This

sulcus, though minute, is.constant on both sides.

Midway between the suprasylvian sulcus and the rhinal fissure,

and within the curve of the former, there is a short deep hori-

zontal sulcus about 5 mm. long on the right side, and about twice

that length on the left. This sulcus is exactly analogous in

position to a similar short one in Orycteropus. For purposes of

reference it will be spoken of as sulcus “ d.”

On the dorsal surface of the neopallium (text-fig. 157, p. 890),

running in general parallel to, and on an average about 7 mm. from,

the middle Tine. 4 is a deep sullen, which extends from very near the

olfactory peduncle to the posterior surface of the hemisphere.

This long furrow bears several branches and is really compound.

On the left side of the brain, the most anterior part of it, about

1 em. long, lies further from the middle line than the rest of the

furrow, and was probably not directly continuous with it, though

in the cast there is a connection by means of a shallow depression,

This short suleus must be the prorean. On the right side it is

directly continuous with the rest of the sulcus, which is clearly

the lateral. The relation between the prorean ‘and lateral sulci

is exactly as in Orycteropus.

The large lateral sulcus, though roughly parallel to the middle

line, is by no means straight, and pursues a sinuous course along

the hemisphere. On the right side it has two very short branches

going outwards from it, at distances respectively of 1:5 and

3°3 ems. from its caudal extremity.

At the junction of the prorean and lateral sulci on each side

there is another sulcus, a few millimetres in length and running

890 MR. R. W. PALMER ON THE

straight forward. What exactly this represents is not clear.

It may possibly be a rudiment of the crucial.

Text-fig. 157.

‘i My ZB Via

) WV,

in, 24.

aN WY os

ACAN gr

Dorsal view of brain-cast of Anoplotherium.

Lettering as in text-fig. 155, with addition of a.md., area medullaris ; fsp., fissura

suprapyramis; s.crw.?, crucial sulcus(?); s.la¢., lateral sulcus; sp., supra-

pyramis ; s.p70., prorean sulcus; wv., uvula; v., v’., vessels.

At this point a couple of blood-vessels, marked on the dorsal

surface of the cast, may be mentioned. On the right side, a

small transverse ridge in the middle of the hemisphere and near

the middle line may represent the course of one of the meningeal

vessels or of a branch of the internal carotid artery. Branches

of this latter artery probably account for some of the obscurity of

BRAIN OF A FOSSIL UNGULATE. 891

the brain-surface behind the orbital fissure, where the cast shows

the marks of the fronto-parietal suture.

Posteriorly, between the lateral sulcus and the middle line,

traces of a spacious paired vessel (v’) are seen. ‘This is formed by

two branches, one of which strikes out from the median line, while

the other comes inwards and backwards from the surface os the

brain. The main trunk, from its direction, seems to have gone

round the tentorium and to have joined the transverse sinus. It

was probably a vein collecting blood from the superior surface of

the hemisphere.

Cerebellum.

The general proportions of this part of the brain have already

been given, ‘The cast does not, of course, show the minute folding

of the cortex characteristic of a fresh specimen, but the main

areas and the chief fissures are distinctly marked.

The most striking feature of this organ in Anoplotherium lies

in the small size of the lateral lobes. This is probably a primitive

feature, since, in a comparative series, evolution in the cerebellum

is chiefly marked by the development of these parts [4].

The three fundamental divisions of the cerebellum are indicated

by transverse fissures marked on the vermis.

The fissura prima is not very clearly shown, but there is a

suggestion of a groove in the place where it nearly certainly must

have been situated, on the dorsal part of the anterior face of the

vermis.

The fissura secunda is clearly shown on the posterior surface

of the vermis, marking off from the rest of the cerebellum a

smooth hemispherical area, which faces posteriorly and is un-

doubtedly the wvusa—the anterior division of the lobus posticus.

The whole of the dorsal part of the vermis belongs to the lobus

medius. It is divided into two almost exactly equal parts by a

transverse fissure, which can be nothing but the fissura supra-

pyramis. The vermis of the median lobe shows a twist such as

is seen in a great number of mammals of different groups, and

which gives a marked asymmetry to the organ.

To the sides of the suprapyramis—that part of the vermis

of the median lobe anterior to the suprapyramidal fissure—lie

triangular depressed areas indicating the exposure of white

matter such as is found in many small. mammals including most

Marsupials, Rodents, Bats, etc. [5]. For this region Elliot Smith

has suggested the term ‘ area medullaris.”

To the sides of the pyramis he the indefinite masses of the

lateral lobes, the small size of which has been commented on above.

Their irregular form adds greatly to the asymmetry of the whole

organ.

_ To the outside of the lateral lobes he the probably relatively

large jfloccular lobes. ‘These can be traced as being connected

behind, in the usual manner, to the pyramis. Below, they rest

on the periotics and fill in the floceular fosse. Their exact extent

892 MR. R. W. PALMER ON THE

cannot be determined, owing to the continuity of their casts with

those of the transverse sinuses.

Such a primitive form of cerebellum in an organ of this size 1s

unknown, except in Marsupialia (acropus and Thylacinus) {6}.

Brain-stem.

In cranial casts this region is never represented with any

accuracy, owing to the fact that the nerves and vascular spaces,

which lie below the brain, obscure the outlines of the latter.

Conspicuously shown (text-fig. 156, p. 887) to the outside of the

pituitary region are the courses of the nerves escaping by the

sphenoidal fissure (which, as mentioned above, is confluent with

the foramen rotundum) and by the foramen ovale.

The course of the trigeminal nerve can be followed from the

outside of the region of the Pons Varolii, along a groove in: the

alisphenoid to the foramen ovale, through which it gave off its

mandibular branch, and then, still lying in a groove on the bone,

the second and first branches ran forward to pierce the skull with

the third, fourth, and sixth nerves through the sphenoidal fissure.

Occupying the centre of the figure formed by these two pairs of

foraminal casts, the pituitary body is represented, but the extreme

shallowness of its fossa makes it very inconspicuous. The pos-

terior clinoid processes leave a scarcely noticeable impression on

the plaster.

As to the divisions of the rest of the brain-stem nothing can

be distinguished. It only remains to describe the casts of the

periotic region, of the foramina for the veins of the sinus, and

of the last cranial nerves.

The peviotic bone has left an irregular crater-like scar on the side

of the cast (text-fig. 155, p. 886). This hollow is walled by various

nodose elevations, w hich are chiefly casts of parts of the transverse

sinus. In the Ponte of the “crater” a tiny ridge represents the

passage through the internal auditory meatus and the Aqueductus

Fallopii of the ViIlIth and VIIth nerves. The upper boundary

of the crater is formed by the cast of the floccular fossa. In

front and below is the cast of the foramen lacerum medium,

from which a slight ridge runs forward on the base of the brain

to meet its fellow of the other side below the pituitary fossa.

This ridge may have some connection with the course of the

internal carotid. Below and at the back of the periotic depres-

sion, an elevation represents the foramen lacerum posterius. At

the back of the floccular lobe of the cerebellum, a tiny knob marks

the position of the mastoid foramen (text-fig. 156, m.f.).

On the side of the medulla oblongata, in line with the jugular

foramen, the exit of the XIIth nerve is represented by a tubercle

which is the last feature to be seen on the cast.

At this pot a summary of the resemblances between the

brains of Orycteropus and Anoplotherium may not be out of

place. The Aard-Vark’s brain has been described as differing

from a primitive Ungulate type only in a higher development of

the olfactory parts. The only other important contrast with the

BRAIN OF A FOSSIL UNGULATE. 893

Ungulate brain lies in the insignificance of the suprasylvian

sulcus compared with the lateral, which is the reverse of the

usual condition found in that group [4].

The brain of Anoplotheriwm has the general proportions of

that of Orycteropus, and the “smell-brain” is nearly as well

developed. The horizontal continuous rhinal fissure in the brain

of the first agrees with the usual condition in the second. ‘The

relations of the orbital, prorean, and lateral sulci agree in

both. The suprasylvian suleus of the Anoplothere is much more

developed than in Orycteropus, but its size in relation to the

lateral sulcus is intermediate between the usual Ungulate con-

dition and the state of affairs in Orycteropus. The brain of the

Aard-Vark has long been known to approach the Ungulate type,

and in Anoplotherium we have an Ungulate which somewhat

approaches the Tubulidentate type.

On the ventral surface, the differences between the two brains

ave practically all accounted for by the different degrees of macros-

matism. In the cerebella there are no striking similarities, but

there is nothing to contradict a relationship.

If cerebral anatomy be of any systematic value, the view

that Anoplotherium and Orycteropus arose from a common,

though necessarily remote, ancestry can hardly be doubted.

Tn conclusion, | must emphasize my indebtedness to Prof. Elhot

Smith, whose notes on the brain-cast I have freely used, and who

has given me a great deal of help in many other ways. My

thanks are also due to Dr. Andrews and Mr. D. M.S. Watson

for kindnesses received at the British Museum.

LITERATURE.

1. G. Cuvrer.—‘ Recherches sur les ossemens fossiles”...

Vol. ii. 1822.

2. H. M. D. pe Buarnvitte.—“ Ostéographie.” 1839-1864.

3. C. Harte.—‘‘A Memoir upon the Genus Palewosyops Leidy

and its Allies.” Journ. Acad. Nat. Sci. Philadephia,

2nd ser. vol. ix. pp. 267-388. 1892.

4. G. Exiior Smiru.—‘ The Brain in the Kdentata.” Trans.

Linn. Soc. Lond. (Zool.) vol. vil. pp. 277-394. 1899.

5. G. Huxtior Suira.—‘ Further Observations on the Natural

Mode of Subdivision of the Mammalian Cerebellum.”

Anatomischer Anzeiger, Bd. xxii. p. 369. 1903.

6. Descriptive and Illustrated Catalogue of the Physiological

Series of Comparative Anatomy contained in the

Museum of the Royal College of Surgeons of England.

Vol. 1. 2nd ed. p. 168. 1902.

General.

H. Fitnor.—< Recherches sur les Phosphorites du Quercy.”

1876-77.

G. Extior Smiru.— Notes upon the Natural Subdivision of the

Cerebral Hemisphere.” Journ. Anat, & Phys. vol. xxxv.

p. 431. 1901.

Proc. Zoou. Soc.—1913, No. LX. 60

894 DR. F. A. BATHER ON

D1. The Fossil Crinoids referred to Hypocrinus Beyrich.

By Hea. BATEER, MAC aDESc:aheitas heen

[Received September 8, 1913: Read October 28, 1913. ]

(Plate XC. and Text-figures 158-160.)

ConTENTS.

Page

PrebanGe sh iil ane Mea GR AG SOR STR ti aR veh aria al nes MESS 4

Previous Historyscie sania ccm tenta tase ccahe ees EEE

Suborder Graniocrmonies BRO eC AAPE RO ne aan caahcadnauonccocbes ole)

Themis CEICAROGOWMICEA sco casocesococo sna vvo acoso mngcaccasacocceose SS

TEFANAPDUS, CUMENONS, \ so6:ssc 000429003206 000009 095 295 960 a2ebon000 200800 899

JEANS PUOUD SOP DCOUIP, csc 408 s29008 06406080000 S6H0300G0 can esanoseas GOW)

“ Hypocrinus”’ pir aS Batiste Ue n cay aren aN onMes MocnnGlighecobeg neko)

Gener Cidia on OS18/0\7 sora sass cce eee resem en nce seen cee er Rey een eed

SUMMA YE ose cece shee ese el nea ence eeban tte ohim aac teeleh ae eee eae LS

Be olenacion of Plate Beene tba aionan socnnaniatoedands uonanusacianons woo ue)

INDEX.

Hypocrinus, history of Opimion .....5.....-2....- ences cee eee se) 890

5 rediagOs dil aa ce tecannan Hea e eRe ERASE EE eRe OO,

Hypocrinus schneideri vedescribed saan . 900

‘ Hypocrinus”’ piriformis PidescrinedI and ened Ao the

Taxocrinide as a new genus, diagnosed but not named ... 905

@cenocystist GintyAmotiaa Cystidia eee eee eee ee Een eee ere SON]

Myrtillocrinus Sandb., not a Costexoannidl . SEEN OOD,

Lecythiocrinus adamsi compared with Flapnoeray WNUS) Bostcoree 899

Cydonocrinus compared with “ Hypocrinus”’ piriformis ... 911

Gasterocomide diagnosed and discussed ............2....2.00..--.. 898

PREFACE.

Early in 1908 a crinoid cup from the Carboniferous rocks of

Yorkshire, elsewhere described as C'ydonocrinus parvulus ¥, was

submitted to me by Mr. D. M.S. Watson. Some curious features

in it suggested a comparison with Hypocrinus, which had oceupied

my pen ‘ab intervals duri ing the previous twenty years, but of which

IT had not seen actual specimens, Without delay all the known

fossils that had been referred to the genus were borrowed for

study. ‘This proved a lengthy business, involving patient pre-

paration and microscopic examination. By the end of 1910,

however, their description was drawn up and a series of illustra-

tions prepared, when another specimen was reported as obtained

from Timor by the Elbert-Sunda Expedition. Correspondence

ensued with Dr. J. Elbert and Dr. J. Wanner, but the latter

* Publishea by permission of the Trustees of the British Museum.

+ F. A. Bather, 1913, “ British Fossil Crinoids, [X.,” Ann. Mag. Nat. Hist. ser. 8,

vo}. xll. pp. 588-394,

ID ZSis HON Z), IPL, 2G,

H. G. Herring. Phot. London Stereoscopic Co. imp.

1-6. HYPOCRINUS SCHNEIDERI. 7-10. H. PIRIFORMIS.

THE CRINOID HYPOCRINUS, 895

was away in Timor, and access to the specimen proved for the

time impossible.

My paper, therefore, was laid aside and other work taken up.

Until the publication of that work in July of this year (1913) I

could not return to Hypocrinus ; then, just as [ was about to do

so, I was informed that Professor Wanner wanted to see the

specimens that were in my hands, because he intended to re-

describe the species in his account of the Permian Echinoderms

collected by himself in Timor. Naturally I resumed at once the

communications with Professor Wanner, but, friendly though

our exchange of views and of information has been, I regret that

it has proved impossible for us to come to any arrangement more

convenient to our colleagues and to future workers than this:

that I shall publish my discussion of the original specimens,

and that the description of the new material shall be left to

Dr. Wanner.

This paper, then, is admittedly a fragment. References to one

or two recent papers have been introduced, and my observations

have in places been checked or corroborated by the information

courteously sent me for this purpose by Prof. Wanner. But in

its main lines the paper stands as it was written more than

three years ago. It will, [ hope, be accepted as Prolegomena to

Prof. Wanner’s memoir, relieving him of much needful drudgery,

but not forestalling his more important results.

Previous History.

The genus was founded by Beyrich (1862, Zeitschr. deutsch.

geol. Gesell. Bd. xiv. p. 537) to receive the single species Hypo-

crinus schneidert, based on a unique cup from the bed of a brook,

Kali Mati or Ajer Mati (the dead water), about half a kilometre

south of Kupang in Southern Timor. Beyrich (1865, Phys.

Abhandl. Akad. Wiss. Berlin, Jahrg. 1864, p. 83), when publishing

a fuller description, was doubtful whether his new genus was a

Crinoid or a Cystid, being inclined to the latter view by the

curious infra-radial position of the anal opening.

The first author to notice the new genus was Quenstedt, who,

in his ‘ Handbuch der Petrefaktenkunde’ (ed. 2, 1867, p. 751),

definitely referred it to the Cystidea, placing it with Hchino-

encrinites, but separating it from Cryptocrinus. In ‘ Petre-

faktenkunde Deutschlands’ (1876, Bd. iv. p. 687), Quenstedt

put Mypocrinus, together with Cryptocrinus, near Hchino-

encrinites and its allies, although he considered that the probable

presence of five large arms brought it near to the true Crinoidea.

In the Atlas to that work, pl. 113, fig. 94, Quenstedt gave a view

of the posterior side, and another of the base. These views show

the relation of the basals to the posterior side, and thus indicate

the position of the small basal, a piece of information not given

by Beyrich. Unfortunately, Quenstedt’s interpretation of the

base proves incorrect. In the third edition of the ‘ Handbuch

60*

896 DR. F. A. BATHER ON

der Petrefaktenkunde’ (1885, p. 967) Hypocrinus was still placed

with Hchinoencrinites and its allies.

Zittel (1875, ‘Handbuch der Paleozoologie, Bd. i. p. 413),

who established the Cystidean group Aporitide on the lines

indicated by Miiller (1854), placed Hypocrinus therein next to

Cryptocrinus. KR. Hoernes (1884, Elem. d. Pal. pp. 123, 124),

remarking that both genera possessed pores, especially Hypo-

crinus, transferred them to the Caryocrinide. Hypocrinus was

also associated with Cryptocrinus by M. Neumayr (1889, ‘ Die

Stiimme des Thierreiches, p. 403), who, however, made no

particular remarks on the genus.

Hypocrinus was not mentioned by Wachsmuth and Springer

in their ‘Revision of the Paleocrinoidea,’ although the name

appears within round brackets in their privately issued index.

Before 1890 no one had placed the genus anywhere except

with the Cystidea ; in that year, however, in the second of my

papers on “ British Fossil Crimoids” (Ann. Mag. Nat. Hist.

ser. 6, vol. v. p. 324), [ associated it with Achradocrinus, stating

that it appeared to differ from that genus only in having three

infrabasals instead of five. In the concluding part of the same

paper (tom. cit. p. 885) Achradocrinus and Hypocrinus formed

Series 3, Achradocrinites, in the family Cyathocrinide, Series 2

being Codiacrinites, containing Codiacrinus and Lecythiocrinus.

The view that Hypocrinus was a crinoid was accepted by

P. Herbert Carpenter (1891, ‘On certain Points in the Mor-

phology of the Cystidea,” Journ. Linn. Soc., Zool. vol. xxiv.

pp. 14-16). “ Hypocrinus,” he said, “is certainly a very singular

form and one would like to know more about it.” Carpenter

considered it to be allied to Lecythiocrinus White, of the Coal

Measures of Central North America, but also mentioned its

affinity to Codiacrinus.

Our knowledge of the genus was increased through two cups

collected by Dr. A. Wichmann in 1888-9 at the same locality

of Ajer Mati, and described by Professor A. Rothpletz (1892,

‘ Paleeontographica,’ Bd. xxxix. p. 74) under the names Hypo-

crinus milleri Beyr. (a lapsus calami, as Prof. Rothpletz tells me,

for H. schneidert) and Hypocrinus(?) pyriformis (melius pari-

formis), n. sp. Since Dr. Rothpletz stated that Crinoidea were

represented in the Wichmann Collection by columnars alone, and

since between their description and that of Hypocrinus he placed

the account of an echinoid radiole, he must be supposed to have

regarded Hypocrinus as a Cystid.

Whether the views of Herbert Carpenter and myself were

unknown to foreign writers or whether we were merely thought

unworthy of attention, subsequent authorities continued to refer

Hypocrinus to the Cystidea, associating it as before with Crypto-

crinus. Among text-books one notes Félix Bernard (1893, Elém.

de Paléont. p. 159), Von Zittel (1895, ‘Grundziige der Pala-

ontologie,’ p. 155), and Koken (1896, ‘ Die Leitfossilien,’ p. 290).*

In 1895, Dr. O. Jaekel was describing from the Devonian

THE CRINOID HYPOCRINUS. 897

of Germany several crinoids which I regard as close allies of

Hypocrinus, but he did not mention that genus for comparison.

Haeckel (1896, ‘‘ Die Amphorideen und Cystoideen,” Festschr.

fiir Gegenbaur, Bd. i. p. 147) dealt with the genus between Crypto-

crinus and Lichenoer ystis, but concluded his ‘deser iption by saying :

“wegen ihrer sonstigen nahen Beziehungen zu _ einfachsten

Crinoideen kénnte man sie auch fur reduzirte oder verkummerte

Formen dieser Klasse ansehen.” This statement, while perfectly

true as regards Hypocrinus, is not equally applicable to Crypto-

crinus.

Undismayed by this weight of authority, in a “ Phylogenetic

Classification of the Pelmatozoa ” (1899, Rep. Brit. Assoc. 1898,

p- 923) I placed G'asterocoma, Seaeeninie. Achradocrinus, aad

Hypocrinus in the family Gasterocomide, which came in the

suborder Cyathocrinoidea, among the Dicyclica Inadunata.

This family was maintained, with the addition of Manocrinus,

on pp. 177, 178 of my contribution to Lankester’s ‘ Treatise on

Zoology’ (Part III. Echinoderma, 1900).

In the second edition of the ‘Grundziige der Paliontologie’

(1903, p. 141) Von Zittel included in the family Gasterocomidee

the same genera as I had placed in it. His example was followed

by Potta (1904, ‘ Rukovet Palaeozoologie,’ I.), who, however,

quoted Hypocrinus as “ peemelserbon rusky.”

Delage and Heérouard (1904, ‘Traité de Zoologie concrete,’

Tome iil. pp. 379, 380) reduced my suborder Cyathocrinoidea to

the rank of a family Cyathocrinusine, but retained Hypocrinus

and the other genera in the same relative position.

The latest writer to maintain Hypocrinus in the Cystids under

a family Cr yptocrinids appears to be Dr. G. H. Girty (1908,

“Guadalupian Fauna,” Professional Paper 58, U.S. Geol. Surv.

p. 108). This he has done in connection with ‘the description of

Coenocystis richardsoni, an alleged new genus and species of this

Family, from the Delaware Mountain Formation of uppermost

Carboniferous age in Texas. Whatever Coenocystis may be, I

find no reason for supposing it to be a Cystid. But even if it

were, this would not affect the position of Hypocrinus, which, as

Dr. Girty says, 1s evidently distinct.

The preceding account shows that the position now occupied by

Hypocrinus in our leading text-books depends chiefly upon the

opinions expressed by Herbert Carpenter and myself. It is,

therefore, advisable to point out that, when we wrote, neither

of us had examined any specimens of Hypocrinus. For us it

remained, as Carpenter expressed 1t, a very singular form which

one would like to know more about. At last, the great courtesy

and kindness of Professors Dr. W. Branca of the Museum fiir

Naturkunde, Berlin, and Dr. C. K. A. Wichmann of Utrecht

University, have enabled me to study all the described specimens

of this genus. The following pages contain redescriptions of that

material and fresh diagnoses.

898 DR. F. A, BATHER ON

CRINOIDEA DICYCLICA INADUNATA.

Suborder CY ATHOCRINOIDEA.

Dr. Frank Springer, in his paper on ‘“ Some new American

Fossil Crinoids” (July 1911, Mem. Mus. Comp. Zool. Harvard,

vol. xxv. no. 3), has published a very interesting discussion of

my Suborders Cyathocrinoidea and Dendrocrinoidea. While

differing as to the delimitation of these divisions and the

principles upon which they are based, he readily agrees in

recognizing their existence. He prefers to regard them only as

large Families, to which he applies the names Cyathocrinide and

Poteriocrinidee, but admits that this is ‘“‘a matter of detail,

depending upon the general plan of treatment.” It was necessary

for me to treat them either as Suborders or Superfamilies, because

J split them up into Families, several of which are again divided

into Subfamilies. Iam, therefore, emboldened to retain for the

present the Suborder Cyathocrinoidea.

Family Gasterocomide.

Cyathocrinoidea with anal opening in the side of the dorsal

cup, below the level of the arm-bases. Radial facets of horse-

shoe shape, with distinct axial canal. Infrabasals small, often

fused into three plates or one. Orals, so far as known, for the

most part concealed by the cover-plates of the subvective system ;

posterior oral a large madreporite. Stem frequently with peri-

pheral canals.

Dr. Springer, in the above-quoted memoir (July 1911, p. 122),

has given a full account of the Gasterocomide, although he has not

provided any fresh diagnosis. The characters which he mentions

as distinetive are included in the above diagnosis, which differs

only verbally from that published by me in Lankester’s ‘ Treatise

on Zoology’ (Part 11. 1900, p. 177). Dr. Springer further agrees

with Jaekel and with me in referring to the Family the genera

Gasterocoma, Nanocrinus, Scoliocrinus, and Achradocrinus. To

these headds a new genus, Schultzicrinus, found in the Onondaga

Group of the Middle Devonian in Livingston County, New York.

This genus has the infrabasals coalesced and five simple uniserial

arms. Further, Dr. Springer adds Arachnocrinus Meek and

Worthen, of which he has made a complete study, showing that

our previous views as to the structure, and therefore as to its

systematic position, were erroneous. ‘This genus differs from

Schultzicrimus mainly in having dichotomous arms. I am, of

course, prepared to accept its reference to the Gasterocomide.

Apparently, Dr. Springer would also place in the Gasterocomide

the genus Myrtillocrinus Sandberger, of which he considers 7’77-

pleurocrinus HK. Wood to be a synonym. In this he follows

Wachsmuth and Springer (‘ Revision of the Paleeocrinoidea,’ 1885),

as well as some other writers therein quoted. It may, however,

THE CRINOID HYPOCRINUS. 899

be pointed out that the genotype, Wyrtillocrinus elongatus Sand-

berger, is not very well known, and that Dr. Springer here bases

most of his argument on J/. americanus Hall, which possibly has

not the same structure. If, however, we accept the generic

identity of the two species, then it follows that Myrtillocrinus is

devoid of the most characteristic feature of the Gasterocomide,

namely, the passage of the anus through the dorsal cup. It

seems to me, therefore, as it did to Prof. O. Jaekel (1895, Pal.

Abhandl. Bd. vii. p. 75), that the Family Gasterocomide is far

more homogeneous if JMyrtillocrinus be removed from it.

Whether I was right in placing it with the Cupressocrinide is

another matter, not worth discussion until Sandberger’s type-

specimen has been restudied.

Dr. Springer, on p. 138, somewhat casually mentions Hypo-

crinus, but leaves its systematic position uncertain. On the

grounds of its structure Hypocrinus is naturally referred to this

Family, and there does not appear to be any objection to that

course, except the fact that all the other genera of the Family are

restricted to the Middle Devonian, whereas Hypocrinws is not

known below the top of the Carboniferous. This difficulty is

scarcely lessened if the species Lecythiocrinus adamsi Worthen be

proved to belong to Hypocrinus, since that species is similarly

removed by a large interval of time from the rest of the Gastero-

comidee, being found about the horizon of Coal No. 8 of the Lower

Coal Measures in Peoria County, Illinois.

According to Worthen’s description and figure of Lecythio-

crinus adamsi (1882, Bull. Hlinois State Mus. Nat. Hist. no. 1,

p- 87; and 1883, Geol. Surv. Illinois, vol. vil. p. 317), that species

has five infrabasals, and therefore could not be referred to Hypo-

crinus. This fact, however, would not prevent its being placed

with the Gasterocomide, since Achradocrinus also has five infra-

basals. The reason for placing this species with the Gastero-

comide is that ‘Just above the summit of one of the subradials

basals], and in the lateral angles of two of the radial plates, there

appears to have been a circular opening, rather larger than the

arm-facets, which is now filled with stony matter”(Worthen). This

no doubt is an anal opening, as suggested by Worthen himself.

Although, for reasons of structure, Lecythiocrinus adamsi and

the species hitherto referred to Hypocrinus ave naturally placed

in the Gasterocomide, it should not be forgotten that this peculiar

disposition of the anus may have arisen independently in the late

Carboniferous Cyathocrinoidea. In that case, the forms in

question, wherever they might be referred, would not belong to

the Gasterocomide.

HYPocrinus.

1862. Hypocrinus H. KE. Beyrich, Zeitschr. deutsch. geol.

Gesell. Bd. xiv. p. 537.

For other references, see ‘‘ Previous History.”

Diagnosis.— A Gasterocomid with anal opening at the summit

900 DR. F. A. BATHER ON

of the posterior basal, and at the lower angles of the right and

left posterior radials, which meet above. Two pairs of infrabasals

fused ; the remaining small infrabasal being the anterior. Radial

facets small. (Arms unknown.) Stem-facet from +13 to -2

diameter of cup. (Stem unknown.)

The diagnosis is based on the characters of the genotype, since

H., piriformis must be removed from the genus. 'The somewhat

hexagonal shape of the anal opening suggests that it was closed

by six valvular plates.

Genotype.—ZH. schneidert.

Distribution.—This species, as well as “ Hypocrinus” piri-

formis, has been found only in Timor, in beds which are either

Permian or Carvoniferous. Recent workers have regarded them

as Permian, but the Echinoderms at present described would

rather favour a Carboniferous age. JI have already discussed

this in my account of Schizoblastus from Timor (1908, N. Jahrb.

f. Mineral., Beil.-Bd. xxv. p. 318). The affinities of Hypocrinus

also, as shown above, are closer to the Upper Carboniferous

‘“« Lecythiocrinus” adamsi, 1f not to the Devonian Gasterocomide,

than they are to anything as yet known from undoubted Permian

rocks. It is of no avail to discuss the question further in this

place, for the abundant materials collected by Dr. J. Wanner

and others will probably settle it when they have been described.

I would only suggest that the possibility of some of the fossils

being derived from older beds should be borne in mind. Some

of the few Echinoderms that I have studied had certainly been

much worn before they were included in their present matrix.

HYPocRINUS SCHNEIDERI. (Pl. XC. figs. 1-6 ; text-figs. 158, 159.)

1862. Hypocrinus schneideri H. K. Beyrich.

1865. Hypocrinus schneidert Beyr.; H. E. Beyrich.

1892. Hypocrinus millert [err. pro schneideri| A. Rothpletz.

For references, see ‘“‘ Previous History.”

Diagnosis.—This being the only undoubted species of Hypo-

crinus known to me, no specific diagnosis can be given.

Holotype.—The original of Beyrich’s figures (1565) preserved

in Museum fiir Naturkunde, Berlin (our Pl. XC. figs. 1-4).

The only other specimen described is that of Rothpletz (1892).

Locality.— Both specimens come from Ajer Mati, 8S. of Kupang,

Southern Timor,

Horizon.— Permian 2 (vide supra).

Description of Holotype.—A dorsal eup, with no trace of stem,

anal plates, brachials, or tegmen. Surface of stereom stained

pinkish. Infilling matrix, soft micro-erystalline calcite with

small dark specks.

General shape squat, pyriform. Cross-section subpentagonal,

owing to projection of umbones of basals, which is especially

marked in the posterior basal. Height 20°8 mm.; greatest

sagittal diameter 20°38 mm., greatest transverse diameter

19°9 mm., these being at about 14°5 mm, from basal plane.

THE CRINOID HYPOCRINUS. 901

Stem-facet (Pl. XC. fig. 4) circular, markedly concave, bounded

by a slight rounded rim; external diameter 2°7 mm.; internal

diameter 1:75 mm. Lumen circular, about -3 mm. diameter.

There are exceedingly faint traces of about 15 or 16 radiating

grooves.

Infrabasals (Pl. XC. fig. 4) closely united. The posterior

suture is most distinct, chiefly because a’ crack is partially co-

incident with it. When the specimen reached me the right

anterior suture could scarcely be detected, only being indicated

by a faint band of lighter colour. Various processes of prepa-

ration have now rendered it perfectly visible. The left anterior

suture at first seemed conspicuous, but the appearance was due to

a vather irregular crack continuous with that neticed along the

posterior suture. The left anterior suture itself could not be

traced with certainty; it now appears to be only in part co-

incident with the crack. In the concavity of the base, after

careful preparation, there are dimly seen two dark streaks

running from the lumen in the direction of the posterior and

right anterior sutures respectively. There appears to be a third

such streak a little on the anterior side of the above-mentioned

erack ; and this streak probably indicates the position of the left

anterior suture.

Tt follows from the preceding account that Beyrich’s description

and figures of the base,.as composed of two large plates and one

small plate, though unsupported by valid evidence, were in the

main correct. His figures are little more than inexact diagrams,

and neither they nor the text indicate the position of the small

infrabasal ; it is anterior (text-fig. 158).

Text-fig. 158.

RR GS ce (en) e @

Hypocrinus schneideri.

IBB

Analysis of the cup of the holotype.

The periproctal outline is dotted in. Nat. size.

Heights of IBB :—anterior 8-7 mm. ; right large 1B 7-3 mm.

along median line, 9°3 to posterior angle; length of posterior

suture 7°3 mm.

Basals bounded by fairly distinct sutures, emphasized by the

902, DR. F. A. BATHER ON

bands of lighter colour that accompany them (Pl. XC. fig. 3).

The measurements in millimetres are :—

ey ere ile gy maa.

posta eee ee LOS alee lais 118 13°7

THs DOS 1B coe cee eepecn ae A TOG ee ates | 5G |) Rs

HPs, BUI: 183i ganado eos eeeonsane 8:7 | Tale) 9°3 | 17

Rar eB edhe seen Viieccvel S774 il weelO; Olea ei | | 518hs

Ost Neen 9°2 | 11:7 | 103 | ca. 139

The greatest width of each basal is on a level with its umbo,

at a little more than half its height. The umbo is most

prominent in the posterior basal, which is also the widest of the

basals and would be the highest were it not truncated by the

periproct.

Radials sloping from top of basals to the peristome rather

suddenly. Height of each from the lower angle to the inner

margin of the peristome about 8 mm. Width of each below

about 8°3 mm.; width above about 2 mm.

The upper margins of the radials form a distinctly pentagonal

border to the dorsal cup (Pl. XC. figs. 1, 2). Each side of the

pentagon forms a smooth slightly raised rim, and is about 2°3 mm.

long. At each angle of the pentagon les a brachial facet.

Facets rather worn, so that their precise constitution cannot

be detected. A ventral groove leads downwards into the cavity

of the cup. From this to the outer edge of the facet is about

15mm. Width of facet about 13mm. Neither muscle-fossve

nor an axial nerve-canal can be detected.

Within the pentagonal rim, the edges of the radials slope

steeply downwards into the cavity of the cup (cf Pl. XC. fig. 5).

The downward extension appears to be greater near the facets

than near the interradial sutures, and may have served for the

attachment of muscles. There is no trace of any tegminal or

oral plates, or of any sutural surface on which they might have

abutted. This fact and the contraction of the cup at its upper

end lead one to regard the pentagonal opening as a peristome,

covered in life only by oral folds of a flexible and thinly-plated

or naked integument. Below the left posterior facet, in the

peristomial cavity, 1s a fragment of a plate, but its nature is not

clear.

The Periproct (Pl. XC. fig. 1) truncates the uppermost angle

of the posterior basal with an obtuse re-entrant angle, 2°8 mm.

across, and cuts into the adjacent lower angles of the right and

left posterior radials with a curve of approximately the same

chord (2°8 mm.), also somewhat angular, so that the outline

is approximately an irregular hexagon nearly 3 mm. high. Of

THE CRINOID HYPOCRINUS. | 903

anal plates no trace remains, but the relatively large size of the

opening suggests that some probably existed.

Oimament:—Beyrich says ‘‘Sammtliche Platten sind aussen

glatt und scheinen zum Theil von pordsem Gefiige zu sein.”

Rothpletz remarks that his smaller specimen ‘ die pordse Structur

der Platten, welche Beyrich an seinen stark abgeschliffenen

Exemplare nur vermuthen konnte, sehr deutlich zeigt.” As a

matter of fact, the holotype is only ‘“ strongly worn down” in one

or two places, and it shows the texture of the plates (Pl. XC.

fig. 6) no less clearly than the specimen described by Rothpletz.

The latter author continues :—‘‘ Die Porositiit der Platten wird

evkannt an den abwechselnden kleineren und grésseren rundlichen

Vertiefungen, welche die Oberfliche bedecken. Die kleinen

Poren stehen regelmissig und dicht gedrangt, die grosseren

liegen vereinzelt dazwischen und hiufen sich auf den Radialia

nahe den Anzatzstellen und auf den Parabasalia in der Gegend

der Buckel.” This account applies equally well to the holotype ;

but I must add that in neither specimen can I detect any

regularity of arrangement in the smaller pores.

The diameter of the smaller pores is about ‘075 mm. or more.

The diameter of the larger pores is*3 min. or less. There is

every gradation between these sizes.

This micro-structure of the test is so unusual, if not unique, in

its absence of regularity and in the sporadic occurrence of larger

pores, that Beyrich’s use of the word ‘“ scheinen” may have been

due toa hesitation as to the true nature of the appearance. ‘The

irregular surface, especially in the umbonal region of a basal,

markedly resembles some encrusting hydrozoon with autopores

and dactylopores. Were the appearances really due to some such

encrustation, this would not have been the only occasion on

which an echinoderm plate has been described as porous on

similar evidence. But in the present instance, where the rough

surface has been worn down, the same porous structure is still

seen penetrating the plates. The pores are rendered distinct by

the infiltration of a reddish iron colouring.

Since the matrix itself is of a granular micro-crystalline com-

position, with darker specks, it is very difficult to discriminate

between the matrix and the rough surface of the plates. Beyrich

may have been uncertain whether he was looking at the actual

stereom or at an imperfectly cleaned surface. The porous

substance is of a saccharoidal appearance, and is easily scraped or

broken with a needle, even with a brass pin. It does not cleave

with a smooth fracture, as does the denser stereom near the

sutures.

This apparent porosity of the test doubtless weighed with

those authors who referred Hypocrinus to the Cystidea, but

the structure is really nothing but an exaggeration of that seen

in the cup-plates of the larval Antedon before the connective

strands of stroma have impressed on the stereom a radiate or

fasciate arrangement. The exaggeration consists partly in the

greater size of the pores, which, as figured by W. B. Carpenter

904 DR. F. A. BATHER ON

in his classical memoir (1866, Phil. Trans. B, vol. 156, pl. 41),

have, in the * eribriform films” of Antedon, a maximum diameter

of ‘02 mm., as contrasted with -38 mm. in Hypocrinus ; partly in

the greater relative thickness of the plates. But, so far as I

can make out, in a place where the stereom was already worn,

the inner layer, or hypostereom, was much denser, so that the

pores nowhere pass right through the test.

Description of the Specimen figured by Rothpletz. (Pl. XC.

fig. 5 & text-fig. 159.)—In its main features this agrees with the

holotype, and only the points of difference need here be

mentioned. i

Text-fig. 159.

a Os OL eae

Oe es

es eo ae

Hypocrinus schneideri.

Analysis of the cup in the specimen figured by Rothpletz.

The periproctal outline is dotted in. Nat. size.

Height 13°5 mm. Posterior basal not so prominent, partly,

perhaps, because worn down. Greatest sagittal diameter

13°2mm. Greatest transverse diameter 13°3 mm. These both

at about 8 mm. from basal plane.

Stem-facet obscure, but appears irregularly triangular, the

angles directed towards the sutures between the infrabasals.

Greatest external diameter (in direction of |. ant. suture) circa

27mm. No trace of radiating grooves.

Infrabasals. The left larger one is considerably cracked, and

some of the cracks in part coincide with the sutures ; otherwise

all the sutures are very hard to trace. Heights of IBB:

anterior 6°6 mm.; right large IB 5:1 mm. along median line,

5°8 mm. to posterior angle; length of posterior suture 4°7 mm.

Measurements of Basals in millimetres :—

Nar Re ere eee) ete

POSHWB had. welee ene 66 70 6°8 84

if OS tal} eee ee aes | 57 7-0 61 86

TMT ber kan egeeone eee ee 61 85 74 90

OEE Vii BHM Ge atekce Seccauae 66 71 67 87

[PE OSES FB ascent oc conesnns 61 7 6:1 7-8

THE GRINOID HYPOCRINUS, 905

Tt will be noticed that the width is here on the anterior, rather

than on the posterior, side.

The Radials have a height of circa 5-4 mm.,a width below of

6-6 (1. ant.), 6°3 (ant.), 5-7 (1. ant.), and a width above of circa

1-7 mm.

The Facets (Pl. XC. fig. 5), with their ventral grooves, are

rather clearer than in the holotype, but still no muscle-scars can be

detected. From the cavity of the cup to the outer edge of each

facet is circa 1 mm., and this is also the width of each facet.

The Periproct has a rather more regular shape than in the

holotype, and its upper angle is slightly produced adorally. The

height of the opening is 2°4 mm., the width 2 mm.

‘“‘ Hypocrinus” prrirormis. (Pl. XC. figs. 7-10; text-fig. 160.)

1892. Hypocrinus (?) pyriformis A. Rothpletz, 1892, p. 75,

jOlls 3x6 ths Aa).

As will appear from the description, this species must be placed

in anew genus, So long as the imperfect holotype was the only

available specimen, the curious features of its structure might be

interpreted as abnormalities, and the proposal of a fresh name

was scarcely justified. The material obtained by Dr, Wanner

and the facts which he has very kindly communicated to me

render a more precise interpretation possible and necessitate a

new genus, - That Dr. Wanner may not be deprived of the

fruits of his own discoveries, I leave to him the privilege of giving

it a name and of completing my imperfect descr iption.

Holotype.—The unique specimen figured by Rothpletz and

preserved in the Mineralogisch- Geologisch Instituut te Utrecht.

Locality.—Ajer Mati, S. of Kupang, Southern Timor. Dr. J.

Wanner has recorded this [or possibly an allied] species from

Fatu Bitaoni (or Bitatnu) in Insana district, Timor (Centrlbl. f.

Mineral. 1910, p. 737).

Description of Holotype.—A dorsal cup with no trace of stem,

anal plates, or tegmen. Stained pinkish. Matrix fragmentary

organic limestone with crystalline cement. General shape

elongate- -pyriform ; somewhat irregular owing to peculiarities in

the radial circlet ; with a tendency - to be flattened along the per-

radial meridians, or, conversely, to be broadly ridged ‘along the

interradial anemia, from the top of the basals downwards.

This produces a sub-pentagonal cross-section. Height circa

43 mm. Greatest sagittal diameter 28 mm. Greatest trans-

verse diameter 31°50 mm, ‘These diameters are measured by

callipers with the jaws parallel to the main axis of the cup;

but the diameters are not at right angles to the main axis.

On the left-hand side the greatest projection is 26°7 mm.

above the basal plane, on the right-hand side it is 24°5 mm.

The Stem-facet was broken away, and the proximal end of the

cup was therefore slightly ground down by Dr. Rothpletz (Pl. XC.

fig. 7), who says that it Lisst in der Mitte einen Centralcanal

erkennen, von welchem die drei Niihte der Basalia [7, e. IBB|

906 DR. F. A. BATHER ON

ausstrahlen. Nahe der Peripherie liegen fiinf rundliche Poren

in den letzteren, von denen eine auf das kleinere, je zwei auf

die grdsseren [Infra-|Basalia fallen. Dies diifte fir die

urspriingliche Anlage von fiinf [Infra-|Basalia sprechen, von

denen je zwel nachtriiglich erst verwachsen sind.” The cross-

section of the worn end is not regularly circular: the sagittal

diameter is 7°4 mm., the transverse diameter 6°6 mm. The

diameter of the stem-facet may therefore be taken as about -25

that of the cup. The supposed * Centralcanal” is of irregular

outline and is far from ‘“‘in der Mitte,” since its centre lies at

2-7 mm. from the posterior side and 4:7 from the anterior side.

Proceeding from this irregular area are three cracks, which,

however, do not appear to reach the periphery precisely at the

points where it is cut by the sutures between the three infra-

basals. Under careful lighting one can detect traces of sutures

continuing those on the posterior side, and they seem directed

towards a point more strictly central than the supposed axial

eanal. The “ fiinf rundliche Poren,” which represent the axial

nerve-canals, are not quite so obvious as might be supposed.

They are irregular in shape, size, and position, and that in the

small infrabasal lies at the outer edge of a long depression,

which seems to have been regarded by the draughtsman of

Dr. Rothpletz (Taf. x. f. 236) as formed of two pores. That

figure has the small infrabasal towards the observer.

Infrabasals closely united. By removing a little more of the

firm adherent matrix, I have been able to make the sutures

clearer, so that the proof of their position no longer rests on the

doubtful appearances of the proximal end. The axis of the small

infrabasal is in a line with the periproct. Heights of IBB:—

small IB 15 mm.; right large IB 14°4 mm. along median line,

15-1 to upper angle; length of suture between the large IBB

12°3 mm. The infrabasal cirelet seems to bear indications of

growth-lines parallel with its upper margin and crossing the

vertical sutures (Pl. XC. fig. 10).

Basals (Pl. XC. figs. 8, 10) bounded by distinct sutures.

While all rest symmetrically on the infrabasals, none is quite

symmetrical above. The lack of symmetry corresponds with

abnormalities in the radials, and will be described in connection

therewith. The measurements in millimetres are :——

Width Greatest Width Hecht |

below. width. above. ea

ey OCK\ine Bip peenpaseelsoanoc scans 12:7 171 116 26°8

Ye POstoUBin. daencseeaesecees 13°7 168 iLleil ca. 25°6 |

Pian B ie, ee ceed myet2so8 lull 70 ca. 120 248 |

lant. B i ee 145 | 176 ca. 11-0 22°7 + |

L.gpost Bb vx. yeusveners 12:0 16°6 12°5 25°3

THE CRINOID HYPOGRINUS. 907

The greatest width of each basal is at about two-thirds of its

height, and here also the sides slope equably to a prominent

umbo, above which the plates bend towards the oral pole, the

bend being more sudden on the left than on the right.

The Radials (Pl. XC. fig. 9 ; text-fig. 160) continue the slope of

the upper part of the basals. They present several abnormalities

due to growth, and are further obscured by crushing. Only four

were distinguished by Dr. Rothpletz, but I believe I can see

traces of a fifth, though much atrophied [and this is fully con-

firmed by Dr. Wanner in litt.|. Ant. R and 1. ant. R were

Text-fig. 160.

IBB

“ Hypocrinus” piriformis.

Analysis of the cup of the holotype.

The sutures obscured by wear or fracture are represented in dotted line,

as is also the outline of the periproct.

v.p.R denotes the minute plates, of rather uncertain number and shape, one or all of

which probably represent this radial. On l.p.R the outline of the cleaved

surface is represented. Tbe outline of a.R is modified by a slight shear.

Nat. size.

apparently of less modified outline than the others, though in

this specimen weathering and crushing have rendered that out-

line incomplete. The l. ant. R rests on |. ant. B by a longer

side than that which rests on |. post. B. Similarly, the side of

ant. R, resting on the r. ant. B seems to have been longer than

that on |. ant. B. Ant. R is broken, and part of it, still united

to l. ant. R has been pushed inwards and under the remaining

portion. Thus it is difficult to compare the interradial sutures

bounding these two radials, but there is nothing to show that

they were not of equal length. The upper margin of each of

these radials was excavated by a concave curve, and the edge

908 DR. F. A. BATHER ON’

was slightly bevelled so as to give this the appearance of a very

obtuse angle, with its apex about in the middle line of the radial.

This appears to represent a greatly simplified brachial facet, but

there are no articular markings, unless it be a slight depression

near the middle.

The r. ant. R was similar in its genera] outline to the ant. and

l. ant. RR, having, like them, the basiradial suture longer on the

posterior than on the anterior side, It differs, however, in that

its left or posterior side is divided into two tracts: one a very

short suture connecting the upper angle of the plate with what

seems to be part of the r. post. R; the other an excavate margin

to the periproct. The curve of this margin throws the left upper

portion of the radial away from the median line, so that the sub-

angular concavity of the facet is longer on the left than on the

right.

The r. post. R is probably represented partly by a small

fragment still attached to r. ant. R and partly by a fragment

attached to |. post. R. The right-hand fragment has been pushed

slightly inwards. Whether these fragments all represent the

r. post. R, whether a part or the whole may not represent anal or

perisomic plates, are questions that cannot be settled on the

evidence of this specimen. [Dr. Wanner iw Jitt. confirms and

extends my observation of some small piate or plates in this

position, one of which, at any rate, he identifies with r. post. R.]

The really important fact is that r.post.R has been so far

atrophied that we may safely infer the entire absence of any

r.’ post. arm.

The 1. post. R is, in every direction, larger than the others.

It rests, almost symmetrically, on the shoulders of the 1. post. and

post. BB and abuts normally on |. ant. R. On its right (7. e.

posterior) side, however, the radial in its lower part bounds the

peviproct, and in its upper part has a short sutural union with

the supposed r. post. R. The upper, or peristomial, margin is

regularly concave, with a slightly more marked curvature than

that of the other radials, but shows no bevelling, groove, or

depression, such as might suggest an articular facet. Outside

this margin, however, the highest part of the plate rises into a

mass of calcite (possibly stereom), which has been truncated by a

cleavage-fracture, sloping slightly downwards and outwards from

the oral pole. ‘The evidence of the holotype leaves it uncertain

whether this protuberance formed the base of a mere hump, of an

unjointed spine, or of an arm; but Dr. Wanner tells me that

some of his specimens permit “die sichere Beobachtung, dass

l. p. R eine Gelenkflache fiir einen Arm zeigt,” and his diagrams

show that this joint-face corresponds with the cleavage-surface

just described. It will be interesting to learn the exact consti-

tution of this facet, whether it has a fulcral ridge and axial canal,

and whether there is a ventral groove leading down to the

peristome. Of such structures the holotype presents no trace

whatever.

THE CRINOID HYPOCRINUS. 909

The projection of this facet is due not merely to the protu-

berance on the radial, but also to the greater height of that

radial and of the subjacent posterior basal. The sum of these

heights is again relatively exaggerated by the nearer approach of

' the left posterior side of the theca to the vertical as compared

with the right anterior side, which bulges outward. ‘Thus the

oral surface of the theca slopes downwards from the left posterior

perradius towards the right anterior interradius. If the erect

position of the stem be admitted, then all these departures of the

theca from the normal symmetry result in raising the supposed

large |. post. arm further and further from the sea-floor.

The evidence thus far does not absolutely warrant the con-

clusion that the r. ant., ant.,and ]. ant.arms had become atrophied

out of existence, or even that they were so reduced as to be use-

less as food-collectors ; but it does lead us to infer that the task

of sending a food-stream to the mouth was mainly, if not entirely,

thrown on the I. post. arm.

The following are measurements of the radials in millimetres :—

Left Right

: : |

Height. aedeh ee | lower lower

| : a | margin. | margin.

a, Cialis 1 Na ee 75 9°4 Be |) Gal 45

CNG Gh gene ee ? 2 58) | ? Bb yl

preserved. | |

Wants oh een evenvea| ka geo. TIN azo B4or| 54 |

to edge of | AA

| periproct. | |

| | | | |

Ik, OSS IB, searesdencatwecc| ian CeLeen lip hy wlIEC/ AN Ratssyeal Note 83 |

Peristome (Pl. XC. fig. 9).—The measurement “ width above”

indicates how much of each radial enters into the peristomial

margin. The opening is “ elliptische,” as Rothpletz describes

it, only because the plates are shifted; probably it was of

irregular outline, with an angle about the middle of each of

the four large radials, and a longer side in the posterior region.

One would like to know how this peristome was closed: by

oral plates, by other plates of interradial position, by enlarged

cover-plates of the left posterior arm, or, asseems highly probable,

by the brachials of the reduced arms of the trivium.

Periproct (Pl. XC. fig. 9)—Dr. Rothpletz regarded the

subcircular or subhexagonal opening between the post. and

r. post. BB, and the 1. post., r. post., and r. ant. RR as the

periproct. He did not observe all the difficulties to which this

interpretation led, such as the apparent absence or almost com-

plete atrophy of a radial, or, still more, the unusual position thus

assigned to the small infrabasal, or even the necessary conclusion

Proc., Zoou. Soc.—1913, No. LXI. 61

910 DR. F. A. BATHER ON

that his species could not be a Hypocrinus. So long as there was

only one specimen known, it might have been legitimate to evade

some of those difficulties by a vague hypothesis of abnormality,

or, more precisely, to have supposed that the anus occupied its

usual position opposite the small basal (which region is worn

away in the holotype}, but that it had become choked, and that a

new outlet had been formed by the partial resorption of that

radial which in the present paper is called r. post. R, but which,

on that interpretation, would have been ant. R. When, however,

Dr. Wanner informs me that all his new specimens agree in this

curious position of the opening, and that none of them shows any

trace of an opening on the opposite side of the theca, then J am

bound first to accept the identification by Rothpletz, and after-

wards to face the difficulties involved.

The measurements of the periproct in millimetres are: height 5;

width below, 5:4; left lower margin 3°9; right lower margin 2°7.

There is no trace of any periproctal plates, unless they are to be

sought in the minute plates between the left posterior and right

anterior radials.

Ornament.—In addition to the growth-lines already mentioned,

there are to be seen on the better-preserved infrabasals, 7. e. on

the posterior side, traces of pustules, apparently coinciding with

the growth-lines. There also seem to be similar pustules on a

small tract of the posterior basal. The rest of the surface is too

badly preserved for the ornament to be detected. Dr. Rothpletz

says that “die durchweg abgeriebenen Platten lassen ihre pordse

Beschaffenheit zwar noch erkennen.” ‘They are, however, far

from presenting the curious appearance of H. schneideri, and on

the better-preserved portions no pores can be observed. Most of

the surface is irregularly worn and contains numerous holes.

These latter, however, are either due to some boring organism or

to the impression of hard particles in the matrix. Often the

hard particles, which may be grains of a dark mineral or frag-

ments of other fossil organisms, are still seen closely adherent to

the cup-plates, or even forced into them.

Affinities of “ H.” piriformis.—Comparison of the analyses

shows at once that this species is not a Hypocrinus. The position

of the small infrabasal, the relations of the periproct to the pos-

terior basal and to the superjacent radials, and the modifications

of all the radials, are considerable distinctions.

It is even doubtful whether the species should be referred to

the Gasterocomide, although it agrees with the diagnosis of

that Family, as given above, in all the known essential features

except in the shape of the radial facets. No other Gasterocomid,

however, has the infrabasals arranged in quite the same way.

These two points demand closer scrutiny.

The position of the small infrabasal in the right posterior

radius is characteristic of Flexibilia Impinnata rather than of

Dicyclica Inadunata. When the latter forms have a tripartite

THE CRINOID HYPOCRINUS, 911

base, the small IB is usually anterior, as in Hypocrinus. A few

other positions for it have been recorded, but not, so far as I can

find, the same as it occupies here, unless it be in the little-

known Ampheristocrinus Hall (1881) from the Niagara group of

Indiana.

The radial facets of the Gasterocomide are of the horse-shoe

shape so common in Cyathocrinoidea. In “H.” piriformis the

facets are so greatly modified that it is perhaps unsafe to base con-

clusions onthem. The arm-bearing facet seems to have occupied

almost the full width of the peristomial margin of the left pos-

terior radial in the holotype; but the diagram Dr. Wanner has

sent me shows it as occupying about half the width. ‘There is,

in the holotype at any rate, no trace of the axial canal. The

other facets are atrophied, it is true, but such traces of them as

there are seem to indicate a pre-existing facet as wide as the

radial rather than one of horse-shoe outline. Wide facets are,

of course, almost universal in the Flexibilia Impinnata, though

not confined to that group.

If the reference of “/.” piriformis to the Flexibilia be taken

as a possible hypothesis, we have next to inquire whether in that

Order the periproct ever emerges below the summit of the radials.

I am unable to find that such an arrangement has ever been

deseribed in any known genus, but that is no valid argument

against the possibility. Among Dicyclica Inadunata this peculiar

position first appears suddenly in the Devonian, and I have already

hinted at the possibility of its independent reappearance towards

the close of the Carboniferous. Why, then, should it not have

appeared with equal suddenness in some other group, especially

when the form displaying it is peculiarly specialized in other

respects ¢

Here, moreover, some actual corroboration of the hypothesis -

is afforded by the fossil which impelled me to make this first-

hand examination of Hypocrinus. That is a small cup or patina

from the uppermost bed of the Yoredale series, in Nidderdale,

Yorkshire. On it is based the new genus and species Cydono-

crinus parvulus (Bather, Oct. 1913).

Cydonocrinus belongs to that group of Flexibilia in which, to

use Dr. Springer’s words, ‘“‘the rays and their divisions are

rounded exteriorly, and the interbrachial spaces relatively de-

pressed ” (Journ. Geol. vol. xiv. p. 510, Oct. 1906), This character

affects also the anal plates, which are ‘not united by suture with

adjacent rays, but 1 In arm- -like series, more or less separated from

them by perisome ” (Springer, op. ai. p. 919). Such forms con-

stitute the Taxocrinidz as defined by Dr. Springer, and it is clear

that, of the included genera, Cydonocrinus is most nearly allied

to Taxocrinus. Thus it has the same fundamental plan of the

cup as has “4.” piriformis.

Now the features in which Cydonocrinus differs from Tazxo-

crinus all bring it nearer to “H.” piriformis. They are the

ole

912 DR. F. A. BATHER ON

subglobular shape of the cup, the small diameter of the stem- facet,

the relatively large size of the infrabasals, which in the Flexibilia

are rarely visible, the rise of the radials towards their facets

curiously resembling the hump of the arm-bearing radial in

“H.” piriformis, the crowding of the facets round the peristome

which thus becomes relatively small, and, finally, of greatest

interest in this connection, the outward projection of the posterior

basal, which, combined with the constriction of the upper part

of the cup, almost entirely separates the rectal channel from the

general thecal cavity. In the actual fossil the anal plates, which

once rested on the posterior basal, have been broken away, so

that the appearance is that of a periproct, opening at the top of

the posterior basal and at the lower corners of the adjacent

racials.

Given, then, the existence of such a form in Middle Car-

boniferous times, it is easy to see how the accentuation of the

left posterior radial with its arm would have pushed the periproct

further to the right, and crowded out the right posterior radial.

This and the diminution of the other arms would have produced

a form agreeing with “/7.” piriformis. On this hypothesis some of

the minute plates between the left posterior and right anterior

radials may be relics of the perisomic plates.

Physiologically considered, a modification bringing the food-

intake so close to the vent would be curious in any group of

erinoids; but that such a modification would not be altogether

out of character with the Flexibilia may be gathered from a

perusal of two suggestive paragraphs by Dr. Springer (Journ.

Geol. vol. xiv. pp. 496, 497, 1906). Speaking of “that strange

influence which has modified the bilateral symmetry of almost

every genus in this entire group,” he points out that ‘‘if the arms

have an asymmetrical distortion, it is to the right, never to the

left.” If the anals are shifted, it is likewise to the right, so that

the excavation of the posterior basal for their reception is “fon the

right shoulder of the plate.”

Timor is a long way from Britain, and I am not going to say

that the genus which must be established for “//.” piriformis is

necessarily descended from the genus to which our Yorkshire

fossil belongs; but I do claim to have shown some reason for

thinking that “ H.” piriformis may belong to the Taxocrinide,

and that, peculiarly modified though it is, its very modifications

are exaggerations of a tendency natural to that Family.

Accepting this systematic position, at all events until it is

proved erroneous, we may draw up the following :—

Generic Diagnosis.—A Taxocrinid with no radianal, with large

IBB forming a conspicuous part of the cup, with left post. R and

arm enlarged and all others reduced in size, the right post. R

being almost entirely atrophied, so that the rectum passes between

post. and r. post. BB below and 1. post. and r. ant. RR above.

THE CRINOID HYPOCRINUS. 913

SUMMARY.

The two specimens of Hypocrinus schneidert Beyr., described by

Beyrich and Rothpletz respectively, are redescribed and refigured.

The structure of the genus is shown to agree with that of the

Devonian family Gasterocomide, but it is suggested that in this

case and in that of “ Lecythiocrinus” adamsi the distinctive

features may have been independently acquired.

The holotype of “ Hypocrinus” piriformis Rothpletz is ve-

described and refigured, and proved to be no Hypocrinus. It is

thought to be a highly modified descendant of the Taxocrinide,

by way of such a genus as C'ydonocrinus. The left posterior

radial appears to have borne a large arm, but the other arms were

more or less atrophied, and the right posterior radial has almost

disappeared.

EXPLANATION OF PLATE XC,

Hypocrinus schneideri Beyr.

Figs. 1-4 are photographs of the holotype, enlarged slightly

less than 2 diameters.

Fig. 1. View to show the relations of the periproct, which is the opening nearest

the observer.

2. View trom above the oral pole, showing the peristome and radial facets.

hig. 3. The cup from the right side: o., oral pole ; a., anal opening or periproct.

Note the light colour of the dense stereom at the sutures, contrasted

with the staining of the porous stereom of the general surtace.

ig. 4. The cup trom below, with the posterior imterradius away from the observer.

Note the sutures between the infrabasals.

Fig. 5. A photograph os the peristomial area of the specimen figured by Rothpletz.

The periproct is at the lower corner of the figure. X 5 diam.

Fig. 6. Micro-photograph of the upper part of one of the basals in the holotype.

x 7 diam.

“HH ypocrinus” piriformis Rothpletz.

Fig. 7. The ground-down base of the holotype, photographed under water, so as to

bring out the sutures between the intrabasals and the five axial nerve-

canals in those plates (see detailed description on pp. 905, 906). The

orientation is as in fig. 4, and approximately opposite to that of Roth-

pletz, pl. x. f.236. X 4 diam. <

Figs. 8-10. Photographs of the holotype, nat. size.

Fig. 8. Shows the posterior basal with the protuberant left posterior radial (1. p. R.)

above it and the periproct at its right upper corner.

Fig. 9. View from above the oral pole, showing the distorted peristome with the

large but obscure facet of left post. R. to the left, and the anal opening

towards the observer.

Fig. 10. The basal facing the observer is the right anterior. Above it is the peri-

stome (0.), with the left post. R. rising up behind it, and with the anal

opening (a.) to the left. Note the ornament of growth-lines in the lower

part of the cup.

All the photographs were taken by Mr. H. G. Herring, and represent a

selection from many attempts upon these dithcult subjects.

[| Note. The following corrections were received too late for insertion in previous

sheet :—

Page 896, line 15 from bottom ‘ For Hypocrinus miilleri.

Pee X00, = I) 4 ve Read H. mnlleri.

EpITor. |

914 DR. W. T. CALMAN ON

52. On Freshwater Decapod Crustacea (Families Potamonidee

and Paleemonide) collected in Madagascar by the Hon.

Paul A. Methuen. By W.T. Caiman, D.Sc., F.Z.8.T

[Received October 8, 19138: Read November 11, 1913. }

(Plates XCI. & XCII.¢ and Text-figure 161.)

INDEX. Page

Imtroductioniande lis trot Mocalitvesseemereeeeeeaiteeeree ee eee eee eee em Le

Part I. Family Poramontp™.

1. Description of the Material .. 915

Potamon (Potamon) madagascaric iense A. Milne- ‘Edwards,

and its Varietal Forms ...... cet cee OG

Potamon (Potamon) goudoti H. Milne-Edwards ............... 920

Potamon (Potamon) methueni, sp. n. SSomahr escent aoe OOO)

Hydrothelphusa agilis A. Milne- Edwards. Sioa O22

2. Systematic Afhinities of the Madagascar River! ior abe besa eas an Ce

3. Geographical Relations of the Madagascar River-Crabs ......... 925

Part Il. Family PanmMonip 2.

Palemon (Macrobrachium) lepidactylus Hilgendorf ............... 926

Palemon (Parapalemon) dolichodactylus Hilgendort ............ 926

Palemnon (Hupalemon) ritseme de Man ..... .......2.-0.000ee0-00-.. 927

Palemon hildebrandti Hilgendorf, and its Varietal Forms ...... 928

INTRODUCTION,

The collection described in this paper was obtained in Mada-

gascar in 1911 by the Hon. Paul A. Methuen, to whom I am

indebted for the opportunity of examining it. The specimens

are the property of the Transvaal Museum, Pretoria, but, by

permission of the Acting Director of that institution, a selection

from among them, including the holotype of the new species, has

been retained for the British Museum.

For the purpose of comparison with Mr. Methuen’s material

the specimens of Potamonidze and Palemonide from Madagascar

already in the British Museum collection have been re-examined

and the results have been incorporated in the paper.

The following is a list of the localities whence Mr. Methuen’s

specimens were obtained. Those from the localities marked with

an asterisk were collected for Mr. Methuen by M. Herschell-

Chauvin of Tamatave :—

Hastern Region of Madagascar.

Manambato, streams running into Lake Rasoabé. July,

SNL

Ambilo, streams running into lagoon. July, 1911.

* Rapides de l’Ifotry, a deux joursde Tamatave. September,

*Ivondro. August, 1911.

+ Published by permission of the Trustees of the British Museum.

{ For explanation of the Plates sce p. 932.

1 ZS MONS) IP COI,

G,M, Woodward delet lith. fith amp.

POTAMON METHUENI.

Ie Wigs), WS) WS Ie CIO,

G.M.Woodward ta et hth. Huth imp.

PALA MON HILDEBRANDTI.

DECAPOD CRUSTACEA FROM MADAGASCAR. 915

Mountains leading to the Mangoro Terrace.

Ambatonharanana, near Ampasimpotsy. July, 1911.

Mangoro Terrace.

Imerimandrosa, northern end of Lake Alaotra, altitude

800-900 metres. July 1911.

Mountains leading directly to the Plateau.

Ambohidratrimo. June, 1911.

South-western Region.

Andranolaho and Tongoroby, Onilahy River, Dist. of

Betsioky. October, 1911.

Part I.—Family PoramMonip& (River-Crabs).

1. Description of the Material.

The River-Crabs of Mi. Methuen’s collection are referred to

the following species :—

Potamon (Potamon) madagascariense A. Milne-Edwards.

* i goudoti H. Milne-Edwards.

+ methuent, sp. n.

Hydrothelphusa agilis A. Milne- Edwards.

As will be explained below, the specimens included under the

first of these names differ considerably among themselves and

represent at least three distinct forms which may possibly deserve

to rank as varieties or even as species.

In the specific descriptions which follow, regard has been

given chiefly to the characters of the carapace, to the exclusion of

those of the appendages, since the latter seem to afford no useful

points for separating the species here dealt with. Attention may

perhaps be called to the difficulty of deciding whether any single

specimen is to be regarded as adult. In the case of females the

broadening of the abdomen seems to afford a fairly definite

indication, and it is usually easy to sort the specimens into

“adults” and ‘‘immature.” In the males, however, the genital

appendages of the first and second abdominal somites are often

found apparently well developed in very small individuals, which

differ considerably in the outline of the carapace from larger

males.

In the lists of specimens the numbers preceded by the letters

‘“« BLM.” refer to the British Museum Register of Crustacea. The

measurements given in the tables are expressed in millimetres

and were taken by means of a scale graduated to°5mm. No

very high degree of accuracy can be claimed for them, but I

believe that they are fairly comparable among themselves; it is

not in all cases certain that they can be directly compared with

916 DR. W. T. CALMAN ON

measurements given by other writers, owing to the frequent lack

of precise indication of the points between which measurements

have been taken. In those given here ‘‘length” is measured

from the hinder edge of the carapace to the front in the median

line, 7. e., to the bottom of the notch between the frontal lobes ;

‘“‘ breadth” is the greatest breadth of the carapace wherever it

may be; ‘‘exorbital width” is the distance between the tips of

the exorbital teeth, and is therefore least accurate where these

points are blunt and indefinite; “frontal width” is the most

difficult of all to find fixed points for, owing to the obliquity of

the sides of the front ; the measurements here given have been

taken between the innermost points of the orbits, 7. e., just above

the antennal flagella, when the carapace is viewed from in front.

Complete references to the earlier literature relating to the

species dealt with will be found in Miss Rathbun’s monograph, to

which I am glad to acknowledge my great indebtedness.

PoraMon (POTAMON) MADAGASCARIENSE A. Milne-Edwards.

Potamon (Potamon) madagascariensis A. M.-E., Rathbun, Nouv.

Arch. Mus. Paris, (4) vi. 1904, p. 264, pl. ix. fig. 7.

I have ventured to group under this name a number of speci-

mens which differ considerably among themselves, and of which

some at least would probably be regarded by other students as

representing distinct species. I have e so grouped them, partly as

a confession of failure, for I feel no great confidence that they

really belong to a single species, although I have found no

satisfactory characters by which to distinguish them, and partly

in order to emphasize the fact that the various forms can be

arranged, in a way that seems to me significant, around the:

typical form of the species. They radiate outwards from this

typical form towards the other species of Potamonide recorded

from Madagascar, and while in some cases there remains enough

of a gap to justify the retention of the names given to these

other species, the relations are clear enough to suggest for the

Madagascar River-Crabs an autochthonous origin from some

form resembling the typical P. madagascariense. Miss Rathbun,

in her monograph, attributes a certain range of variation to

this species, and notes that one of the forms shows a transition

towards P. hwmbloti. If the view here put forward be correct,

the range of variation must be considerably extended, and

P. humbloti, P. grandidierit, and P. bombetokense must be

included as varietal forms.

The specimens examined appear to fall naturally into five

series as shown on the accompanying table of measurements. ‘To

avoid prejudging the question of their systematic rank, I designate

these series by letters.

DECAPOD CRUSTACEA FROM MADAGASCAR,

Measurements of Potamon madagascariense.

oii

Locality, &c.

Series A.

Madagascar, B.M.81.9 ..

3 B.M. 85.18 ...

BY BICTOOO ss

Betsileo, SMSO Che aise

39 32

SpErRizs B.

| Ifotry, Methuen, No. 90............

SERIEs C.

| Ambatonharanana, Methuen,

No. 25...

Surtes D.

| Madagascar, B.M. 88.5

29 33

SERIES EH.

Imerimandrosa, Methuen,

TB, COL copsprootessee

No. 35

O4 40404040 Oy OY +0+0 044004 05 +0

O303+0+0

OsQ3 00404 400440054005

Sex.|

4005

Length. | Breadth.

50 66

30°25 39

P55) 21:25

11 14°5

10 12°5

16°75 22

13°5 17-25

13 16°75

12°75 16

25°25 33°5

Zi 27-5

20°26 26

28°5 36

20°35 26

19°75 25°75

37°5 46°5

32°5 385

22°25 26°5

19°5 23°25

165 21°5

16°25 21

16 22

1575 | 205 — |

15°5 20°5

14:5 19

14°5 19

NEP || Tes

13°25 17

11 13°5

10°25 13

34 45°5

35 45°5

AT 575

46°5 58°75

45°5 56

38°5 AT 5

Exorbital

width.

Frontal | Breadth

width. | ratio.

20 1°32

12 1:28

ut 1eBy/

5°25 131

4:75 1:25

7-25 1°31

6 OT

5'd 1:28

515 1°25

105 1:32

9 11583

85 1:28

11 1:26

Sion E26

Sion melee)

14°75 1-24,

13 118

9°25 1:19

8 119

i I

7B || 128

CAS | SI

7 | 133

7 | 16

65 1:31

65 131

6 1:27

6 1:28

4:75 1:22

4-75 1:26

14 1:33

14 1°33

17°25 1°22

ile 1:26

16°5 1:23

14°75 1:13

SERIES A.—These specimens are all, without much doubt,

referable to the typical form of P. madagascariense as described

and figured by A. Milne-Edwards and by Miss Rathbun.

characters in which they differ among themselves are relatively

unimportant, with two exceptions, viz. :—(1) the distance between

the epigastric and the postorbital crests, which, in the largest

The

918 DR. W. T. CALMAN ON

Specimen (81.9), isa trifle less than half the distance between the

former crest and the frontal margin, is about 2/5ths of that

distance in the largest specimen from 85.18, 1/3rd_ in the largest

from 82.6, and only about 1/4th in an adult female from 80.27,

while in the smallest specimen from 85.18 the two crests are all

but continuous ; (2) the roughness of the surface of the carapace,

which, in most of the specimens, is in accordance with the

published descriptions and figures, becomes notably reduced in

the specimens from Betsileo registered under the numbers 82.6

and 80.27 ; in these, the whole carapace is much smoother, with

only a few oblique granules or raised lines near the antero-lateral

margins and on the posterior branchial regions, and one or two

obsolescent granules on the front; it is to be noted that the

eranulation or serration of the antero-lateral margins themselves

is not correlated with the general roughness or smoothness of the

surface, being stronger in the Betsileo specimens than in the much

rougher specimens from 85.18.

In the smaller specimens of this series, especially in the two

smallest from 85.18, the external orbital notch becomes very

shallow or is altogether obliterated, and the outline of the carapace

is more quadrilateral than in larger specimens.

Serres B.—The four specimens of this series differ from those

included in Series A in the relative flatness and narrowness of the

carapace, which has the antero-lateral borders less arched, so that

the lateral projection of the carapace beyond the exorbital tooth

is only about 2/3rds of the greatest diameter of the orbit instead

of at least 5/6ths as in the larger specimens of Series A. These

differences, however, are rather less strongly marked in the

larger than in the smaller specimens of this series. The frontal

lobes are very prominent in one specimen, and in all there is a

tendency for the margin of the front to be less deflexed than in

Series A. The exorbital and epibranchial teeth are very pro-

minent, and, although the carapace as a whole is much less rough

than in the largest specimens of the first series, the epigastric and

postorbital crests are sharply defined and the oblique lines on the

lateral regions are numerous, sharply cut, and. granulated. The

marginal granules on the under surface of the merus of the

chelipeds are very prominent and spiniform. It is worth noting

that even in the larger of the two males the genital appendages

are very short, less than half the length of the abdomen, and

apparently immature.

This form or variety seems to me to lead in the direction of

Hydrothelphusa.

Series C.—The largest female in this series is only 16°25 mm.

in length, but 1s apparently adult ; the largest male is a trifle

larger. The carapace in all is rather convex, especially antero-

posteriorly, and the front is bent downwar ds. The carapace, on

the whole, is rather wide, the ratio of breadth to length being, in

one specimen, as high as 1°37. The surface is rather smooth,

DECAPOD CRUSTACEA FROM MADAGASCAR. 919

although not more so than in some specimens of Series A. The

epigastric and postorbital crests are somewhat indefinite; in some

specimens they are nearly in line, but in others they are separated

by a distance estimated at 2/5ths of that between the epigastric

and the frontal margin. ‘The exorbital tooth is not prominent

and the epibranchial i is small.

The specimens of this series approach the P. humbloti and

P. grandidiert of Miss Rathbun’s monograph, especially perhaps

the latter, although they do not agree exactly with the description

of either. They differ so much among themselves in all the

characters that might be used to distinguish them from the

specimens of Series A, that I cannot regard them as specifically

distinct ; from Series B they are more easily defined.

Series D.—In the two specimens of this series the carapace 1s

very smooth, the rugosities on the anterior part of the branchial

region being hardly perceptible. The general antero- posterior

convexity of the carapace is well marked, the various regions are

also convex, and the grooves between them cor respondingly

distinct. The most characteristic feature is the inflation of the

anterior branchial regions, which, in both, but especially in the

female, is much more marked than in any of the specimens men-

tioned above. The antero-lateral marginal line is not prominent

and its granulations are nearly obsolete. In the female the

distance between epigastric and postorbital crests is nearly half

that between the former and the front, while in the male the

proportion is not more than two-fifths.

These specimens, which I cannot believe to be specifically

distinct from the smoother specimens of Series A, approach

P. goudoti in general aspect. and are separated from it chiefly by

the character of the epigastric crests, which do not project nearly —

so far forwards, are much less oblique, and far more clearly dis-

joined from the postorbital crests than they are in that species.

Series K.—This series consists of four large, dark-coloured,

male specimens which differ considerably from all those discussed

above. The carapace as a whole is moderately convex, with the

regions somewhat inflated and the grooves between shen well

marked. The surface is fairly smooth, with oblique striz on the

anterior part and more distinct raised lines on the posterior part

of the branchial region. The crests are well defined and sharp,

the epigastric well in front of the postorbital. The epibranchial

tooth is large and the exorbital moderately prominent.

According to Miss Rathbun’s analytical key to the species of

the subgenus Potamon these specimens would be referred to

P. bombetokense, described by her from a single male specimen.

From the detailed description and figure of that specimen, how-

ever, they differ in the rather narrower carapace, in having the

margin between exorbital and epibranchial teeth straight or

slightly convex instead of concave, and, apparently, in having the

inter-regional grooves more strongly marked.

920 DR. W. T. CALMAN ON

Poramon (Poramon) Goupori H. Milne-Edwards.

Potamon (Potamon) goudoti H. M.-K., Rathbun, Nouv. Arch.

Mus. Paris, (4) vi. 1904, p. 305, pl. xiii. fig. 10.

Measurements of Potamon goudoti.

‘ | | |

Locality, &e. Sex. Length. | Breadth. eee | ae ieee

‘Wedeeasene (no locality), ty ;

Methuen, No 55.| ¢ 30°25 41 265 12°75 1°35

Betsileo, B.M. 82.6............| 6 | 31% 42°5 28 13°5 1°35

r z, .| 3 | 29 39 25°5 115 1:34

Antananarivo, B.M. 82.13 ...| 3g | 29 39°5 26 12 1:36

% 4 3g | 28 38°5 25725) Uo n | ueesiam|

| 55 Hae 27°25 36°25 24°25 11 | 1°33 |

| ff , (dry)| ¢ | 29°75 | 41 26°5 1275 1°37 |

Rs ss ~ | Oh 2 39°25 25°75 12P3 WK

| i ‘ 5 | & | 286 37-5 PES | aul || kei

The nine specimens agree in general with the published de-

scriptions and figures of this species, and present no noteworthy

differences among themselves.

PoraMon (PoTAMON) METHUENI, sp. n. (Plate XCI.)

Measurements of Potamon methueni.

Locality, &c. Sox.| Length. Breadth. Reo | outa Breese

Teac hone nee epee ie EE eee

Methuen, No. 34. | 3 33 46:25 | 26°75 12°5 | 1:4

9 20 36.| ¢ | 30°75 43°75 25 12 | 1°42

3 i 39.| g | 2825 | 39 24 11 | 1°38

Description.—Carapace: length about five-sevenths of its

breadth, very convex, especially from before backwards, gastric

region sometimes below level of inflated anterior branchial regions ;

surface finely punctate, smooth. Oblique portions of cervical

groove shallow and obscure, if produced forwards they would cut

the line of the postorbital crest rather within the outer angle

of the orbit; H-shaped depression well marked; posterior part

of mesogastric region wider than urogastric, the latter fairly well

defined posteriorly. On the anterior branchial region is a faintly

marked oblique ridge nearly parallel to the cervical groove.

DECAPOD CRUSTACEA FROM MADAGASCAR, 921

HKpigastric crests not very prominent, more or less rugose,

hardly oblique, situated entirely behind a line joining epi-

branchial teeth ; they are continued, without interruption, into

the protogastric portions of postorbital crest, which are very low,

rounded, and ill-defined, except in so far as they are limited

by the transverse depression behind each orbit; in general

direction the postorbital crest is transverse, and it 1s not trace-

able external to its intersection by the cervical groove. Groove

between the epigastric lobes bifurcating widely behind, the limbs

angularly bent so as to become parallel. Front strongly deflexed,

but with anterior margin yisible from above; divided into two

low rounded lobes, most distinct from above, less so from in

front; with a smooth raised marginal line. Upper margin of

orbit with a prominent smooth border, inclined forwards, with

only a faint trace of a lobe in the middle, towards the exorbital

angle which is not at all prominent or dentiform. Lower margin

of orbit nearly straight as seen from below, and smooth or

obscurely crenulated; without any trace of a notch or sinus

below the exorbital angle. Antero-lateral margin strongly arched,

with a distinct marginal line, which is smooth or, in the smallest

specimen, obscurely crenulate, and runs on to dorsal surface

behind. Postero-lateral border concave, with more or less distinct

lines. Pterygostomial region and, iess conspicuously, subhepatic

region beset with tubercles.

Merus of third maxillipeds slightly wider than long; ischial

groove rather nearer the inner than the outer margin.

Chelipeds unequal. Lower surface of merus margined with

tubercles, which may become indistinct ; a small blunt tooth near

distal end of inner edge; upper edge rugose with a low sub-

terminal tubercle. Carpus smooth and punctate externally ;

with two teeth on inner side and a variable number of small

granules. Surface of hand smooth and rather coarsely punctate ;

fingers long, slender, and strongly deflexed.

Sixth somite of male abdomen shorter than its distal width.

Holotype, wale, 33 mm. in length, in British Museum. Para-

types, the two other males mentioned above, in Pretoria Museum.

Remarks.—This species resembles, on the one hand, P. goudoti

and, on the other, the smooth form of P. madagascariense described

under Series Dabove. From the former it differs chiefly in the

wider carapace (the concavity of the postero-lateral borders has

the effect of increasing the apparent difference in this respect) ;

in having the epigastric crests nearly transverse, lying behind

instead of in front of the line joining the epibranchial teeth, and

quite continuous with the obsolescent postorbital crests ; in having

the gastric region less elevated and the transverse convexity of the

carapace therefore less marked; and in having no notch below

the external orbital angle. Compared with P. madagascariense

the same differences hold good, even although in that species the

epigastric crests are less oblique and less advanced than they are

in P. goudoti.

999 DR. W. T. CALMAN ON

HypRorHELPHUSA AGILIS A. Milne-Edwards.

Hydrothelphusa agilis A. M.-E., Rathbun, Nouv. Arch. Mus.

Paris, (4) vil. 1905, p. 266, pl. xvii. (Potamonide, xv.) fig. 7

Measurements of ye aioe aS

| |

Locality, &c. Sex.|} Length. |

| width. | width. | ratio.

| Methuen, No. 10) Q 2 | 36°56 PMS | PRS 3) alate |

is % 11] g | 285 | 32% 255 lal Welelco a la |

: ‘ 12, g | 295 | 335 25°5 1225 | 113 |

5 a 131 9 | 305 3455 | 26:5 112;5 10a pela oi

| i a 14) $ | 31-75 | 3625 | 27 | Be | me

| Ambohidratrimo, |

Methuen, No. 28) 9 | 27 BS || HPSS ii 115

| i i 291 g | 315 | 38 27 135 | 12. |

| : rs 301 2 | 385 455 32°75 15°5 118 |

| i cs 31| 9 | 275 325 | 23-5 IS) aeLS |

a xf 32 g | 26 3025 | 2275 | 1025 | 116 |

The arrangement of the spinules on the lower orbital margin

is less regular and constant than is implied by Miss Rathbun’s

description. In other respects the specimens agree well with the

published accounts of this species. All the female specimens,

with the exception of No. 30, are clearly immature.

2. Systematic Affinities of the Madagascar River-Crabs.

In his monograph of the Indian River-Crabs (Cat. Crust. Indian

Mus. pt. i. fase. 1. Potamonide, 1910), and in a short but very

important later paper on the classification of the family as a

whole (Records Indian Mus. v. pt. iv. 1910, p. 253), Lt.-Col.

Aleock has given an entirely new aspect to the system of the

Potamonide. It is necessary, therefore, to enquire how the

Madagascar species stand with reference to the new divisions of

the family. Relying mainly, but not exclusively, on the character

of the mandibular palp, Alcock has divided the River-Crabs of the

Old World (apart from the aberrant Deckeniine) into the two

subfamilies of Potamonine and Gecarcinucine. In the former

(text-fig. 161, A) the terminal segment of the palp is simple,

although it may be thickened amd plumose at the base, while in

the latter (text-fig. 161, C) it is “deeply cut into two lobes which

embrace the incisor process of the mandible.” Alcock refers the

Madagascar genus Hydrothelphusa to the Potamonine, but he

pout out that ‘the thickening at the base of the ter minal joint

DECAPOD CRUSTACEA FROM MADAGASCAR. 923

of the mandibular palp is more than ordinary prominent.” I

find that this statement applies not only to Hydrothelphusa, but

also, and in an even greater degree, to the other Madagascar

species that I have examined. In Hydrothelphusa the proximal

thickening forms a sharply marked ridge ; in Potamon madagas-

cariense (text-fig. 161, B), P. goudoti, and P. methuent this ridge

overhangs a little as a free lobe. This evidently forms a transi-

tion to the bifid palp of the Gecarcinucine, although it is sufli-

ciently far removed from the typical Gecarcinucine condition as

seen in Parathelphusa tridentata (text-fig. 161, C), for example, to

allow us to refer all the Madagascar species with confidence

to the subfamily Potamonine, with which they also agree in

Text-fig. 161.

Mandibular palp (denuded of sete) of three species of Potamonide. The lower

figures represent the palp as seen from below, the upper figures the terminal

segment as seen from infront. A. Potamon fluviatile. B. P. madagascariense.

C. Parathelphusa tridentata.

the shape of the male abdomen. It is possible that a similar

transitional form of the palp may yet be found in species of

Potamonine from other parts of the world, but it is evident from

Alecock’s remarks that it does not occur in any of the Indian

species ; and, in the absence of evidence as to the species of the

African and Malaysian Regions, it may be provisionally regarded

as suggesting specially close affinity between the species found in

Madagascar.

I do not think that much importance attaches to the fact that

in all the Madagascar species the palp consists of only two

g24 DR. W. T. GALMAN ON

segments. Alcock found three segments distinct in all the

Indian species of Potamon, but in his later paper he states that

in the Potamonine the palp has either two or three segments.

From the examination of a number of African species I suspect

that it will not be possible to distinguish sharply between those

species in which the suture-line between the first and second

segments is well marked and those in which it is almost or quite

obliterated.

As regards the generic affinities of the species here discussed,

there is nothing to forbid the conclusion that they are closely

interrelated. ‘They have, indeed, been placed in two distinct

genera, Potamon and Hydrothelphusa, and the species of the

former might not unreasonably be placed in two subgenera,

Potamon and Geothelphusa, but the groups indicated by these

names are probably more convenient than natural. Both Alcock

and Miss Rathbun comment on the close resemblance of Hydro-

thelphusa to Potamon, and Miss Rathbun specially mentions its

affinity with P. madagascariense ; this affinity becomes even more

striking when comparison is made with some specimens of

Mr. Methuen’s collection which I have described above as repre-

senting a form of P. madagascariense (Series B). Potamon

goudots is considered by Miss Rathbun as forming a link between

the subgenera Potamon and Geothelphusa, and the new species

described above as P. methueni goes even further in the direction

of the latter subgenus. There can, however, be little doubt that

Geothelphusa, even as restricted by Alcock, has still very slender

claims to be regarded as a natural or monophyletic group. <A

general softening of the asperities of the carapace, such as we

frequently see within the limits of undoubted species, has only to

be carried a little further to result in the obliteration of the

postorbital and epigastric crests; and there is no difficulty im

supposing that so short a step may have been taken independently

by more than one evolving species. ‘This independence is, in

fact, implied by Alcock’s transference of certain species of ‘ G’eo-

thelphusa” to genera of his subfamily Gecarcinucine, and I think

there is some slight evidence of the same thing among those that

remain in the Potamonine. At all events there is no necessity

to assume that P. goudoti and P. methueni are more closely

allied to the species of ‘‘Geothelphusa” found in Africa and Asia

than they are to P. madagascariense.

Potamon madagascariense itself is placed by Miss Rathbun in

the group of which P. edule (fluviatile) is the type. While it

certainly resembles the species of this group in the characters of

the carapace, it differs a little, as already indicated, in the form

of the mandibular palp, and also in some other small characters,

such as, for instance, the much broader contact of the basal

antennal segment with the front. What the exact value of these

differences may be it is at present impossible to say.

DECAPOD CRUSTACEA FROM MADAGASCAR. 925

' 3. Geographical Relations of the Madagascar River-Crabs.

In addition to the species mentioned above, three others have

been recorded from Madagascar. The African Potamon (Pota-

monautes) depressum was recorded by Lenz and Richters, but its

occurrence is doubted by Miss Rathbun. Potamon (Potamon)

pittarellu, described by Nobili (Boll. Mus. Zool. Torino, xx.

No. 507, 1905) since the publication of Miss Rathbun’s mono-

graph, appears to be allied to P. madagascariense, but is, no

doubt, a distinct species. Finally, Potamon (Parathelphusa)

antongilense, described by Miss Rathbun, is evidently a remark-

able and isolated species, concerning which further information

is much to be desired ; as far as can be judged from the shape of

the male abdomen, it is not a Parathelphusa in the sense in which

that genus has been restricted by Alcock.

Leaving aside these three species, and considering only those

that are represented in the collections now examined, it is

evident that the Potamonid fauna of Madagascar bears no close

relation to that of Peninsular India, which, as Aleock has shown,

belongs entirely to the subfamily Gecarcinucine. This is im-

portant, since geologists seem to be agreed that the connection

of Madagascar with India existed only at a time (not later than

the early Oligocene) when Peninsular India was separated from

the continent of Asia. Of the possible affinities with African

species it is perhaps unsafe to speak until the rich Potamonid

fauna of Africa shall have been examined as thoroughly as that

of India has been by Aleock; but it is worth noting that no

typical Pofamon seems to be known from East Africa; and if, on

the one hand, P. madagascariense is the most primitive of the

Madagascar species, and, on the other, if it is really allied to the

group of P. flweiatile, then it is hard to guess by what route it

ean have reached the island.. Further than this it does not seem

possible at present to go. As I have indicated above, our con-

ceptions of the phylogenetic relationships of the species rest, as

yet, on a very narrow basis of morphological fact; and, until

that basis is broadened very considerably, the group of River-

Crabs, as a whole, appears to me to be a hazardous subject for

zoogeographical speculation,

Parr IJ.—Family Pau #MoN1ID# (River- Prawns),

The Palemonide of Madagascar have already formed the sub-

ject of an important memoir by Coutiére (Ann. Sci. Nat. Zool.

(8) xii. 1900, pp. 249-342, 5 pls.), on which I have relied mainly

for the determination of the four species in Mr. Methuen’s

collection. These are as follows :—

Palemon lepidactylus Hilgendorf.

dolichodactylus Hilgendorf.

ritseme de Man.

7 hildebrandti Hilgendorf.

Proc. Zoou. Soc.—1913, No. LXIT. 62

926 DR. W. T. CALMAN ON

While all the adult males and a good many of the females in

the collection have been referred. to one or other of these species,

there remain over a number of specimens which either lack the

large chelipeds or are clearly immature. It is possible that these

may include representatives of some other species, but I have not

succeeded in identifying them, and it does not seem that any

good purpose would be served by enumerating them here.

The most interesting species in the collection is that which I

identify with Bithynis hildebrandti of Hilgendorf, which does not

appear to have been re-examined since its description in 1897.

The series of specimens collected by Mr. Methuen, together with

those already in the Museum Collection, show, I think, that this

species is not closely related to the Chihan species which is the

genotype of Bithynis; that the single character which led to its

being referred to the genus Bithynis is a matter of individual

verseision 3 and that, therefore, the species should be removed to

the genus Palemon.

Pat#Mon (MACROBRACHIUM) LEPIDACTYLUS Hilgendorf.

Palemonlepidactylus Milgendort, Monatsber. Akad. Wiss. Berlin,

1878 (1879), p. 838, pl. iv. figs. 14-16.

Palemon (Maer obrachium) lepidactylus Coutiére, Ann. Sci. Nat.

Zool. (8) x1i. 1900, p. 272, pls. x., x1. figs. 1-15 a.

Localities. Ambatonharanana, 7 males (Methuen); Rapides de

VIfotry, | male (Wethuen): Betsileo, 3 males, B.M. 80.27.

In the characters of the rostrum and of the larger cheliped all

the specimens fall within the range of variation determined for

this species by Coutiére. In no case, however, does the form of

the smaller cheliped agree with that which he regards as typical.

‘The fingers of this appendage are in all the specimens straight or

nearly so, meeting completely or with only a slight gap, and

having a scanty provision of bristles on their opposed edges. In

these respects the specimens agree much better with Coutiére’s

P. hilgendorfi, and suggest that that species should not be

regarded as distinct from P. lepidactylus.

- PaL#Mon (PARAPALAZ[MON) DOLICHODACTYLUS Hilgendorf.

Palemon dolichodactylus Hilgendorf, Monatsber. Akad. Wiss.

Berlin, 1878 (1879), p. 840, pl. iv. fig. 18.

Paleemon dolichodactylus Coutiére, Ann. Sci. Nat. Zool. (8) xii.

1900, p: 283, pl. xi. figs. 18, 19.

Localities. Andyanolaho and Tongoroby, Onilahy River, 15

males, 3 females (Methuen).

The larger males in the collection agree very well with the

accounts of this well-marked species given by Hilgendorf and by

Coutiére. With them are associated some smaller specimens in

which the woolly covering of the larger chela is very slightly

developed. The rostral formula is ay wibhit4 Suomi thie upper

2-4

teeth postorbital.

DECAPOD CRUSTACEA FROM MADAGASCAR. 927

PAL&MON (EuPALzZMON) RITSEM& de Man.

Palemon(Lupalemon) ritseme de Man, Zool. Jahrb., Abth. Syst.

Ixe SOs po ci4e op. cits x.) 1698 \plyxxcxvan tie. 70!

Palemon ritseme Coutiére, Ann. Sci. Nat. Zool. (8) xii. 1900,

p. 314, pl. xii. figs. 32-33 a.

Localities. Manambato, streams running into Lake Rasovbe,

3 males, 8 females (Methwen); Ivondro, 6 males (Methuen) ;

Ambilo, 2 males (Zethuen); Tamatave, 1 male, B.M. 82.6.

The specimens collected by Mr. Methuen belong, without

doubt, to the same species as a specimen in the Museum Collec-

tion from Tamatave, determined by Mr. Miers as P. ide Heller.

They differ conspicuously from typical specimens of that species,

however, in the fact that the surface of the carapace and of the

tail-fan is completely smooth or presents only very minute and

inconspicuous traces of spinules, while the chelipeds of the second

pair are very finely scabrous. Since Nobili has described a

smooth or nearly smooth variety of P. ide from New Guinea,

and Coutiére records a similar example from Madagascar, this

character may not be of specific value. The chelipeds of the

second pair are shorter, in relation to the body-length, than in

Coutiére’s specimens of P. ide, the longest being only about

11 of that length, and most of them being actually shorter.

Although the carpus is, except in three instances, longer than

the chela, the difference is very much less than is shown by

Coutiére’s measurements in specimens of P. ide of similar size,

and the fingers are always much more than half the length of the

palm. On the other hand, the rostral formula in nearly all cases

falls within the somewhat narrow limits assigned to the variations

of this character in P. ide, and the armature of the fingers agrees

precisely with Coutiére’s description.

In all the points mentioned in which these specimens differ

from the descriptions of P. ide, they approach Coutiere’s account

of the form which he identifies with P. ritseme de Man. Our

smaller specimens, however, which alone are directly comparable

with Coutiére’s largest, have the second chelipeds more asym-

metrical and apparently a good deal stouter, the chela itself, and

especially the palm, being relatively a little shorter. The larger

specimens have many of the proportions rather different, the

greater relative length of the palm in most of them being

noteworthy. Finally, in the specimens now examined, the pos-

~ terior pereopods are, on the whole, stouter than in those

previously described in this species, the ratio of length to breadth

in the propodus of the fifth pair varying from 25 to 21, while

de Man gives the ratio as from 35 to 25 and Coutiére as 25,

On the whole, the balance of characters seems to be in favour

of regarding our specimens as older individuals of the species to

which Coutiére’s specimens belonged, which he has recorded under

the name of P. ritsema@ ; whether they really belong to de Man’s

species is, perhaps, a little more doubtful; and I am not at all

62*

998 DR. W. T. CALMAN ON

confident that they would be found to differ essentially from the

smooth variety of P. ide. It may be noted that, in having parts

of: the surface of the carapace slightly scabrous, some of our

specimens agree with P. sintangensis de Man, which Coutiére

regards as a varietal form of P. ritseme.

The following table gives (in millimetres) the more important

measurements of all the specimens that have one or both

chelipeds of the second pair preserved. The “total length” of

the body and also the ‘‘ total length ” of the chelipeds of the second

pair are only approximate, on account of the impossibility of

straightening the specimens without injury. In an ovigerous

female the eggs measured *5 x -4 mm.

Measurements of Palemon ritseme.

;

| Chelipeds of second pair.

heen «| Total | Rostral

| Locality, &e. Sex Nenoth. ase Tot Hi | |

hansen | Merus. Carpus. | Chela., Palm. | Fingers.

| | | | |

| | |

| Wiktihnen, NOOB ce ING |) PaO BON Sas Wes boi ho 15

| | |

E Ba CaS 23 20150 ate ABE S38 6rn 2Mcome Malet

” BB saps || Ue ay 12 93)| 18 PA) er wri |) Os

} | |

2+9 | | |

x a7...) 6 | 108 | == 88 | 14 2750426 paal lel aaxo

| Manambato, | 948 | | | |

| Methuen, No. 42 ...| ¢ 98 ere pe Eee 22 e23rom elo om eee)

| lvondro, | Oey | 9 | | ¢

| Methuen, No. 99 ...| g | 95 | —Q> | Ve i | 3 | a | ae aoe

| Manambato, | 348 | | |

| Methuen, No. 46...| @ | 92.) == | 2 || 10 165 | 165 |) 85 | 8

247 | | |

| Tamatave, B.M.82.6 | 3 90 way | 2028 35 | SBS || BO 13°

Manambato, I pate | |

Methuen, No. 47 ...| g | 80 3 AGS, 10 hy) Wy 95) 75

| |

. 45 O75 2 2 9 155 96 15 8°5 | 65

| |

PALEMON HILDEBRANDTI Hilgendorf. (Plate XCII.)

Bithynis (?) hildebrandti Hilgendort, Sitz.-Ber. Ges. Naturf.

Freunde Berlin, 18938, p. 244; Coutiére, Ann. Sci. Nat. Zool. (8)

xii. 1900, pp. 250, 256, 259; Ortmann, Bronn’s Thierreich,

Crustacea, ii. 1901, p. 1292; Ortmann, Proc. Amer. Phil. Soe.

Philadelphia, xli. 1902, p. 274.

DECAPOD CRUSTACEA FROM MADAGASCAR. 929

The material which I have examined consists of eleven speci-

mens,from three localities, in the British Museum (Nat. Hist.) Col-

lection and nineteen specimens from one locality in Mr. Methuen’s

collection. ‘The Museum specimens (PI. XCII. fig. 1) agree with

Hilgendorf’s description of this species, except in one or two small

details to be mentioned below. Mr. Methuen’s specimens were at

first regarded as representing a probably new species of Palemon,

since they possess a well-developed hepatic spine on the carapace

(Pl. XCIL. fig. 6). Closer examination showed, however, that

they differed in no other respect from the Museum specimens,

and their specific identity was finally proved by the discovery,

among Mr. Methuen’s specimens, of one which has no trace of a

hepatic spine on one side of the carapace, while on the other

side its place is indicated by a small and inconspicuons tubercle.

The following description applies to all the specimens examined.

The sides of the carapace and the abdominal somites are

rather coarsely punctate. The rostrum is more or less straight,

with the dorsal edge moderately arched, of varying length but

always shorter than the antennal scale, and often shorter than

the antennular peduncle ; of the dorsal teeth, one is postorbital.

The tip of the telson (Pl. XCII. figs. 4 & 5) is rather broadly

rounded, sometimes, but not always, with a minute median point ;

the inner lateral spines are much longer than the outer. Between

the inner spines is a fringe of sixteen plumose hairs.

The third maxilliped does not reach the tip of the antennal

scale. The chelipeds of the first paix extend beyond the scale by

a little more than the chela.

The chelipeds of the second pair (Pl. XCII. fig. 2) are similar,

smooth, with fine scattered setules, and may extend beyond the

antennal seale by the length of the chela and carpus. The carpus

is equal to the merus, widened distally ; the chela slightly com-

pressed (5:6), a little wider than distal end of carpus; palm

longer than carpus and equal to fingers; fingers with a sharp

cutting-edge for the greater part of their length, with two small

teeth near the base of the dactylus, and one, with an indistinct

second, on the immovable finger (Pl. XCII. fig 3).

Posterior legs moderately stout, propodus of last pair abouttwelve

times as long as wide and about four timesas long as the dactylus.

The females are ovigerous from a length of about 38 mm.

upwards. The eggs measure approximately 2 x 1-4 mm.

It will be observed, from the table of measurements given

below, that the females are much more numerous than the males

in the collections examined and also that they considerably exceed

them in size; both conditions appear to be uncommon among the

species of Palemon, but they are found also in P. moorei of Lake

Tanganyika.

Hilgendorf described the mandibular palpas consisting of only

two segments. In all the specimens which I have dissected the

usual three segments are distinct. Hilgendorf also attributes

three teeth to the dactylus of the second chelipeds and states that

930 DR. W. T. CALMAN ON

the carpus “ ragt nur ein wenig unter die Antennenschuppe nach

vorn,” while in our specimen the dactylus bears only two teeth

and the carpus may extend beyond the tip of the antennal scale.

In all other respects, however, Hilgendort’s description applies so

well to our specimens that their specific identity can hardly be in

doubt.

The existence of an affinity between this species and the

Bithynis gaudichaudii of Chile and Peru, asserted only in a

hesitating manner by Hilgendorf, is fully accepted by Coutiére,

but denied by Ortmann, who considers that the resemblance does

not extend beyond a single character (the lack of a hepatic spine)

which may easily be supposed to have arisen by convergence.

Ortmann’s view is strongly supported by the variability of this

character in the present series of specimens. I have examined

the appendages of both species for other evidence of affinity

between them, and have failed to find it. The branchial system

and the mouth-parts of both are very similar to those of several

species of Palemon with which I have compared them. Only in

one point do the mouth-parts of B. hildebrandti present anything

unusual, and that is the reduced size of the epipodite of the

first maxillipeds, but this constitutes no resemblance to B. gaudi-

chaudii, in which the epipodite is quite as large as in the species

of Palemon examined. On the other hand, the differences in

general aspect between the two species are considerable ; B. hilde-

brandti has the chelipeds hardly differing in the two sexes,

slender, symmetrical, smooth, with the carpus equal to the merus,

and the fingers armed with only a few small teeth near the

base ; B. gaudichaudii is a very much larger species, with the

chelipeds very stout, much more strongly developed in the male

than in the female, very unequal on the two sides of the body,

beset with spiny tubercles, with the carpus shorter than the

merus, and the fingers of the larger chela toothed for half their

length or more. If it be advisable to maintain the genus Bithynis

(which seems to me doubtful) it must be for the South American

species alone, and B. hildebrandti must be transferred to the genus

Palemon.

The variability of the hepatic spine in this species recalls th

cases of “mutation” recently described by Bouvier in certain

Atyide. It resembles these cases in its discontinuity, only one

individual out of those examined being in any way intermediate

between the two forms; and it further resembles some of them

at least in the fact that it is geographically limited, for the

specimens of the two forms come from different localities. Both

of these points, however, require to be tested by further collecting.

Tt differs from Bouvier’s cases in that if concerns only a single

character, and one which, were it not for its constancy in other

Palemonide, might be regarded as of trivial importance.

The large size of the eggs may be taken to indicate an abbre-

viation or suppression of the larval development, and this is

likely to be associated, as jt is in some other Palemonide, with

DECAPOD CRUSTACEA FROM MADAGASCAR.

931

an exclusively fresh-water habitat and a restricted area of distri-

bution ; it may be not without significance therefore, that, of the

species of Paleemonide in the present collection, P. hildebrandti is

the only one that does not occur outside the island of Madagascar.

Measurements of Palemon hildebrandti.

Locality, &c.

Ambatonharanana, |

Methuen pene eee

”

»”

a”

”

4.6

Total Rostral | Hepatic |

‘length. formula.) spine. |

Chelipeds of second pair.

| Merus.

Carpus.

Chela.

Palm. | Fingers.

“Tr

932

ON DECAPOD CRUSTACEA FROM MADAGASCAR.

Measurements of Palemon hildebrandti (continued).

| Fingers.

| Chelipeds of second pair.

Air Total Rostral | Hepatic

| Locality, &. Sex. length. formula.) spine. | l Nl

| | Merus. | Carpus. | Chela.) Palm.

foueh. oe i ae nai aaa ag | |

| | | | |

| | 14+5 1 GO) 1. Bo) |) WSO |

| Betsileo, B.M. 82.6.) g | 50 | ~ 3 0 eo | 6:0 | 145 |

a sp il CRA Sh A Aan, sia [pesos felt

ee ieee

| 147 | 657 57 || 13:0

99 ” sisi 2 46 | “o | 0 | ry 5-7 67 12°2

| | 14+6 | (60 60 | 12°2

aera : ea! Rephieeon BoA OO 55 | 12°0

mi ey alee Page ae! Oo 6:0 60 | 120

| | |

| | | |

: Apes, Puen ies = ear 60 | 60 | 136

| | | |

ig | | 147 | 45 | | 45 8-9 |

” 3 ? | 38 SEG ios | 0 ae 4°5 | 89 |

| | |

Madagascar |

B.M.79.20........... 9 | 52 | on GACT Ohi HOR ONT eS Fe

Movie fot @ | a = 0 6:0 60 | 14:0

| |

East Imerina, | |

| BM. 92.7.4.6-7...| 9 | 37 | a8 LO, algae ies Hie! ak |

| | | |

| : 14+6 |

% + | 2 Be To LO eepne tee elm moc ats Ikeaanes

EXPLANATION OF THE PLATES.

Puate XCI.

Potamon methueni, sp. n.

Fig. 1. Male. Holotype. Natural size.

at ES 3 Hand of larger cheliped. 14.

Bs gy ue Abdomen. XX 13.

Aa oe i Third maxilliped. 23.

Pruate XCII.

Pelemon hildebrandti.

Fig. 1. Typical form, ovigerous female. X 23.

2 Cheliped. x 5.

Bo a 4 base of fingers. X 10.

4., as: Telson. XX 10.

D. f Tip of telson. ™ 1d.

6. Form with hepatic spine. Carapace. 2k.

i i)

fay

iat

can

EZ. S TOS El eC nie

RASC

ors:

Ree

sae

nS <

rey At

Nees

> SS

2 oS

Huth, Lith® London.

ML, del.

LE CHRIORCHIS INERMIS.

ON A NEW REPTILIAN TREMATODE. 933

53. A New Trematode of the Genus Lechriorchis from the

Dark Green Snake (Zamenis gemonensis). By Maris

V. Lesour, M.Sc., Assistant Lecturer and Demonstrator

in Zoology, Leeds University *.

[Received September 20, 1913: Read November 25, 1913. |

(Plate XCIL4)

INDEX.

Page

Ethology: A new species of Trematode in the body-

Sy I

cavity of Zamenis gemonensis .......1.ceeee see eee eee 933

Geographical Zoology: Southern Europe; Zamenis

grap sy 5

gemonensis, a new Trematode from the body-

CAVAD YE. cr eecc es dence ae nein ste ence retee CHER Gece te Meebect 933

Systematic: Lechriorchis inermis, sp. n., from the

body-cavity of Zamenis gemonensis .................. 933-936

On June 10th, 1913, a specimen of a Dark Green Snake

(Zamenis gemonensis), from Southern HKurope, died in the Zoolo-

gical Society’s Gardens, and loose in the body-cavity were a large

number of Trematodes which were sent to me for identification.

These prove to be a new species of Lechriorchis Stafford closely

related to Z. validus Nicoll (1911). The habitat is interesting as

all the Trematodes of the group which includes Lechriorchis

whose habitat is known, occur in the air-passages, pharynx,

or anterior part of the cesophagus of their hosts. This appears

to be the first time that one of this group has been found

occupying the body-cavity.

An interesting feature in the new species is the absence of

spines on the body. The cuticle is quite smooth, unlike any of

the other members of the genus. However, in some specimens

spines can be seen below the surface of the cuticle, but not

reaching to the outside, which seems to indicate a gradual dis-

appearance of spines owing to the worm living in the body-cavity

of its host, and therefore being in no danger of getting swept

away as is the case in open passages. In all other ways it agrees

closely with the other members of the genus.

On account of the absence of spines I propose for this new

species the name Lechriorchis mermis.

LECHRIORCHIS INERMIS, sp.n. (PI, XCIII.)

Length of body 5-10 mm. Oral sucker 0°40-0°60 mm,

Ventral sucker 0°60-0°92 mm. Average ratio of oral to ventral

sucker 2:3. Prepharynx small, esophagus the same length as

pharynx. Intestinal ceca reaching a good way beyond testes,

* Communicated by the SpcRETARY,

¥ For explanation of the Plate see p. 936.

934 MISS M. V. LEBOUR ON

Testes behind centre of body, right in front of left. Cirrus-sac

reaching to posterior margin of ventral sucker. Ovary on right

side behind or slightly overlapping ventral sucker. Yolk-glands

reaching from just in front of the level of the anterior margin of

ventral sucker to centre of testes or behind them. Receptaculum

seminis uterinum present on right side of body behind right

testis.

The body (fig. 1) is convex dorsally and nearly flat ventrally, both

suckers being conspicuous and the cirrus usually protruding from

the genital pore. The length of mature worms varies from 5 to

10 mm.,the last-named being in the case of a specially well extended

specimen examined under pressure. Unfortunately, this large

specimen was damaged before being fixed. One immature specimen

occurred measuring 4.4 mm. in length which contained a few

eges and had a very short uterus. In breadth the worms

measured from 1:2 mm. to 1:9 mm., the greatest breadth being

in the region of the ventral sucker. ‘The average length is about

5:5 mm., breadth 1:4 mm.

A good deal of interesting variation occurred in the arrangement

of the organs. The suckers vary but little, the ratio of the oral

and ventral suckers being almost exactly 2:3. Both are circular in

outline with circular apertures. 'The aperture of the oral sucker

measures about 0°20 mm. across; the aperture of the ventral

sucker is more than half the width of the sucker and very regular

and conspicuous. The usual position of the ventral sucker: is

with its centre almost exactly at the anterior third of the body.

It may, however, be a little further back. In the young specimen

it is much nearer the centre of the body, a fact to be expected as

the hind portion of the body always increases to a much greater

extent than the fore part.

The prepharynx is exceedingly short but quite distinct; the

pharynx is broad and muscular, the breadth slightly greater than

the length (length 0°20 mm.); the esophagus is about the same

length as the pharynx.. Surrounding the pharynx are large

glandular cells which are continued for a short distance down the

ceesophagus. Pharyngeal glands are mentioned by Stafford in

L. primus (1905). The intestinal ceca vary in thickness in

different parts and reach usually about 0°60 mm. beyond the

testes; beginning near the centre of the body, they gradually

become more lateral and slightly vential in position until they

reach the testes, when they become quite irregular and may be

both dorsal to the testes or one ventral and one dorsal (fig. 3); ‘and

one may be longer than the other, in one case quite twice as much

extending beyond the left testis as the right. The usual length

beyond the testes is 0°60 mm. Sometimes the ceca bend inwards

so as to be completely internal to the testes. The ends are

swollen and very much broader in the posterior than in the

anterior part.

The excretory vesicle is large and opens at the extreme posterior

A NEW REPTILIAN TREMATODE. 935

end, where it is surrounded by large deeply staining cells; just

in front of the opening the vesicle is much folded, giving it

the appearance of having been “‘ puckered in” before the narrow

opening. In front of this pleating it widens out as a large tube

running dorsal to and parallel with the ascending limb of the

uterus; in front of the testes and behind the ovary it forks, each

limb reaching far forward beyond the ventral sucker to about

midway between the ventral sucker and pharynx.

The testes are oval bodies more or less lobed on their inner

margins, usually lying obliquely with the right testis in front of

the left; the usual situation is for the right to be about 0°15 mm.

in front of the left, but they may be almost on a level, and in

two cases the left testis was in front of the right. Curiously

enough, these variations occurred in the small immature worm

and in the largest specimen. It is just possible that we may

have to do here with a different species. In a specimen 5:2 mm.

in length the testes measured, right 0°90 mm. x 0:44 mm., left

0°84 mmm. x 0°44 mm., the greater axis lying longitudinally. On

the inner margin there are two or three lobes. The cirrus-sac is

long and reaches almost to the posterior level of the ventval

sucker. Sometimes it reaches the extreme posterior end, and in

two cases (again the immature worm and the largest) it was

much shorter and only reached a short way behind the anterior

margin of the ventral sucker; in this respect being much nearer

L. validus Nicoll. The vesicula seminalis is much convoluted

and occupies the posterior third of the cirrus-sac, the ductus

ejaculatorius being surrounded by an enormous mass of prostatic

cells and ending ina short protrusible cirrus often to be seen

exserted from the genital pore. The male opening is on the

right side of the genital pore, which is situated on a level with

the centre of the pharynx and midway between it and the lateral

margin of the body-wall. The ovary is almost round, and lies

dorsally on the right side of the body immediately in front of the

right testis. It overlaps the ventral sucker for about a third of

its diameter; its position varies however, and in some cases it

may be rather more than this and in others not so much. The

portion of the ovary that overlaps the cirrus- pouch is on its right

side. The oviduct receivesa large shell-gland, gives off a Laurer’s

canal, and receives the vitellarian duct, then runs down as the

uterus in many coils to nearly the posterior end of the body,

when it doubles back as a much broader tube, runs forward

gradually straightening and narrowing and passes the ventral

sucker dorsally. In the course of the descending portion behind

the right testis is a receptaculum seminis uterinum. When the

uterus reaches the level of the anterior ends of the excretory

vesicle its walls form a vagina with a thick cuticular lining, and

surrounded by a layer of deeply staining large cells which

accompany it for the whole of its course until it opens on the

left side of the genital pore. The eggs are very numerous and

of adeep brown colour when old, the young eggs being much paler.

936 ON A NEW REPTILIAN TREMATODE.

The cap is not very conspicuous until the egg is open. The eggs

(fig. 2) measure ‘033 mm.—'036 mm. x ‘021 mm.—-023 mm. The

vitellaria are variable in length, but never reach beyond an ex-

tremely short distance in front of the ventral sucker, 0°02 mm. is

quite the furthest they ever reach and usually it is not so much.

Posteriorly they reach to about the middle of the testes, but are

extremely variable in this region, and the left may reach to

behind the left testes and the right hardly reach to the anterior

portion of the right, or both may reach to the level of the middle

of the testes. The vitellarian follicles are in small groups lying

laterally to the outside of the intestinal ceca; small ducts from

them unite in a transverse duct each side lying just behind the

ovary, and these unite in a small reservoir in the centre of the

body which gives off the vitellarian duct to the oviduct.

Lechriorchis validus Nicoll is undoubtedly the most nearly

related to the present species. JL. inermis differs from it chiefly

in its much smaller eggs and position of the vitellaria. The

position of the testes (right in front of left instead of the reverse)

and of the vesicula seminalis being rather more variable, do not

seem to be such important specific characters.

References.

Nicouu, W. (1911).—On Three New Trematodes from Reptiles.

Proceedings of the Zoological Society of London, pp. 677—

681.

SrarrorD, J. (1905).—Trematodes from Canadian Vertebrates.

Zoologischer Anzeiger, xxvii. p. 691.

EXPLANATION OF PLATE XCIII.

Lechriorchis inernis, sp. 0.

The following letters apply to all the figures.

C.S. Cirrus-sac. | R.S.U. Receptaculum seminis

K.P. Excretory pore. | uterinum.

E.V. Excretory vesicle. | 8.GL. Shell-gland.

G.P. Genital pore. T. Testis.

I. Intestine. | U. Uterus.

O. Ovary. | V. Vitellaria.

(ES. Gsophagus. | VG. Vagina.

O.S. Oral sucker. | V.S. Ventral sucker.

PH. Pharynx. V.SEM. Vesicula seminalis.

P.PH. Prepharynx.

Fig. 1. Ventral view. Length 5°4 mm.

2. Eggs, length °033 mm.

3. Transverse section through unpaired portion of excretory vesicle and testes,

showing intestinal cca irregularly placed.

4, Transverse section through ventral sucker,

IBZ gS) MEMS) Je ZIOINY,

A.H.Searle del.et lth, Huth imp.

ZBUGMATOLEPAS MOCKLERI gen.et spn.

Mb Lat

Ob Uy

ATA,

ANS

IF) ops LOS) NPIL 2G’

Huth imp.

A.H.Searle del. et lith.

CALANTICA (TITANOLEPAS) TUBERCULATA, Darwin sp.

ON FOSSIL CIRRIPEDES. 937

54. Cirripedes from the Cenomanian Chalk Marl of

Cambridge. By THomas H. Wiruers, F.G.S.*

[Received September 25, 1913: Read November 25, 1913.]

(Plates XCIV. & XCV.7)

INDEX.

Page

Structure of Zeugmatolepas and Titanolepas ..................941, 946

Lines of Evolution in Pollicipedide .......... ace GOT

Distribution (Geological), Cretaceous of * Europe, Gengiate

lepas and Calantica (Litanolepas) . Berane raat .. 939, 944

Zeugmatolepas mocklert, gen. et SP. N.... 2.0... 2ee cee ree cee cee vee eas 938

Z. crete Steenstrup sp. .... 94.1

Scalpellum crete Steenstr up sp., ‘referred to Zeugmatolepas . . 942

Zeugmatolepas compared with Pollicipes ....... . 941

Scalpellum tuberculatum Darwin, referred to Titanolepas . 943

Titanolepas subgen. nov. of Calantica, for Scalpellum

tuberculatum Darwin ...... ISIS aveh oy aceon eee eke

Calantica, see Titanolepas and Scillelepas BSCE ace d oaauce ons 947

Scillelepas compared with Titanolepas ..............0.c000 eee. GAT

Through the industry of the late Mr. F. Mockler, I have been

enabled to examine a large series of Cirripede remains which he

obtained from the Cenomanian Chalk Marl in the neighbourhood

of Cambridge. By far the greater number can be referred to two

species, which add materially to our knowledge of the phylogeny

of the pedunculate Cirripedes.

One, Zeugmatolepas mockleri, gen. et sp. n., 1s represented by

two nearly complete capitula and portions of nine others, as well

as a large number of isolated valves. It is interesting from the

fact that, while it agrees with: the species of Pollicipes in the

number of valves of the capitulum, it differs not only in the

more specialized form of scutum, but also in the size and position

of the upper latera, which are nearly as large as the scuta, and

have become elevated to occupy the whole of the space between

the scuta and terga. This form certainly serves more than any

other to connect the genus Pollicipes to Scalpellum, and, together

with S. (?) crete Steenstrup sp., from the Upper Senonian of

Denmark, is now included in the new genus Zeugmatolepas.

The second species, Scalpellum tuberculatwm Darwin, is repre-

sented by a large number of separated valves, including several

not previously known. These enable us not only to reconstruet

the capitulum, but also to show that the species must be referred

to the genus Calantica Gray. Representatives of that genus

are now found living, and the geologically oldest species of its

subgenus Scillelepas, to which Scalpellum tuberculatum comes

nearest, occurs in the Miocene of Italy. The recent and

Tertiary species have the umbones of all the valves apical, so

that it is surprising to find in the Cretaceous representative,

* Communicated by Dr. W. T. Canman, F.Z.S,

+ For explanation of the Plates see p. 948.

938 MR. T. H. WITHERS ON

S. tuberculatum, a more specialized form of scutum, the umbo

being subcentral. On account of its higher evolutionary deve-

lopment, S. tuberculatwm is placed in anew subgenus of Calantica,

namely Z7vtanolepas, and it is here suggested that this form may

represent an offshoot from the ancestral Scillelepas line, and may

eventually have given rise to the genus Oxynaspis.

Both species are remarkable in having an advanced form of

scutum in which the umbo is subcentral, and, together with a

small undescribed species of which only the carina is known,

constitute the oldest known examples of this specialized type of

valve. It is quite evident that Zeugmatolepas mocklert and

Titanolepas tuberculata are derived from either the genus

Pollicipes or from a Pollicipes-like ancestor, and both have

evolved the same type of scutum. These forms seem to show

that the transition of the umbo in the scutum from an apical

to a subcentral position, has occurred independently in unrelated

forms belonging to distinct lines of development, and recent

work leads me to believe that this is equally true also of the

carina in the more advanced forms of Scalpelluwm (sensu lato).

In 7. tuberculata the position of the umbo in the scutum is

constant; but in Z. mockleri it changes apparently according to

age, although some individuals seem to be more advanced in this

respect. It may be, therefore, that in the scutum of Z. mocklert

the subcentral umbo was quite a recent development.

The two species must have been comparatively common in the

Chalk sea, for of Zewgmatolepas mockleri we have remains of at

least 100 capitula, and of Z%tanolepas tuberculata more than 30.

Family POLLICIPEDIDS.

ZEUGMATOLEPAS *, gen. nov.

Capitulum longer than broad, comprising at least 34 valves,

composed of scuta, terga, upper latera, and carina, and three or

more whorls of subtriangular lower latera, with a rostrum,

subrostrum, and subcarina ; upper latera almost as large as the

scuta, occupying the whole of the space between the sorta and

terga, the apices of the upper latera contiguous with those of the

scuta ; umbo of scuta subcentral, and of the remaining valves

apical,

Genotype.—Zeugmatolepas mocklert.

ZEUGMATOLEPAS MOCKLERI, sp. n. (Plate XCIV. figs. 1-14.)

Diagnosis.— Valves thin and generally smooth. Carina narrow,

usually with a strong, median, rounded keel, not divided off into

parietes and intraparietes, tage margin acutely angular. Scutum

subtriangular to trapezoidal in shape, with the umbo varying in

position from just below the apex to about one-third the length

of the valve from the apex ; there is in most scuta a deep trough

* Zedyua = a connecting link.

FOSSIL CIRRIPEDES. 939

above the umbo extending almost parallel to the upper occludent

margin ; apical portion usually constricted and acute ; basilateral

angle shortly and obliquely truncated. Tergum subrhomboidal,

<i its upper half curved towards the eeu basal angle

sometimes acute, but more often shortly and obliquely truncated.

Material.—Two almost complete capitula and nine portions of

others. All of these appear to be immature, and there is also a

large series of isolated valves.

Holotype.—The nearly complete capitulum, I. 15830 (Pl. XCIV.

sae, ALT).

Distribution.—Cambridge Greensand: Cambridge; Cenomanian,

Chalk Marl: near Cambridge, and Burham, Kent ; Cenomanian,

A. plena-marls : Oxted, Surrey, and Wantage, Berkshire ; [?Ceno-

manian |, Chalk detritus: Charing, Kent.

Measurements.—The two nearly complete capitula (Pl. XCIV.

figs. 11, 13) each measure 4 mm. from the apex of the scutum to

the base of the lowest whorl of latera, and the uncrushed

expitulum (Pl. XCIV. fig. 11) measures 4-2 mm. in breadth.

Both are immature examples. Among the detached valves the

largest examples of the upper series measure respectively :—

Carina, I. 15820. Length (valve shghtly broken at apex)

5-9 mm.; breadth 1:7 mm.

Scutum, I. 15822. Length (from apex to lowest point of basal

margin) 6°4 mm.: breadth 4°5 mm.

Tergum, I. 15833. Length 7°6 mm.; breadth 3:4 mm.

Upper latus, I. 15834. Length 4:5 mm.; breadth 3-2 mm.

There are, however, several fragments which show that these

valves attained somewhat larger dimensions. A comparison of

these with the proportions of the valves, as shown in the nearly

complete capitulum, seems to show that the capitulum of the

species reached a length of at least 15 mm.

Description of valves.—Valves thin, smooth, or ornamented

with excessively fine ridges radiating from their umbones.

Carina (Pl. XCIV. figs. 1, 2) narrow, widening gradually

downwards from the apex to ‘the basal margin, either slightly

bowed inwards, straight, or bowed outwards, noe ‘divided off into

parietes and intraparietes ; basal margin acutely angular; tectum

slightly to moderately convex transversely ; in some carine there

is a Strong, rounded, median, longitudinal keel, nearly as wide as

the portion of the valve on either side, but in some specimens

it is less strongly marked and in others barely perceptible. Inner

surface somewhat thickened near the apex and marked with

oblique growth-lines ; the extent of the surface thus thickened

varies, and at the most is one-third of the length of the valve;

the valve therefore projected freely to a variable extent.

Scutum (Pl. XCIV. figs. 3-6) varying in shape from sub-

triangular to trapezoidal, moderately convex transversely, the

surface of the valve sloping more steeply from the umbo to the

lateral margin; umbo situated just below the apex in young

specimens, and in mature examples usually occupying a position

940) MR. T. H. WITHERS ON

about one-third the extent of the valve from the apex ; above the

umbo the valve is usually produced abruptly into an acute angle ;

lateral margin longer than the basal margin, almost straight in

its lower part and usually sharply bent inwards and upwards

above a line with the umbo; basal margin indistinctly marked off

into three almost equal parts, the two outer parts being obliquely

inclined towards the apex; lower occludent margin slightly

convex, about the length of the basal margin; upper occludent

margin about half the length of the lower occludent margin,

sometimes nearly in line with it, but more often abruptly bent

from the umbo towards the lateral margin, and in some cases

almost in line with the umbo and nearly at right angles to the

lower occludent margin. Almost all the specimens have a strong

ridge extending from the umbo to a point just above it on the

lateral margin, and above this ridge the narrow portion of the

valve is sharply bent downwards and upwards, and forms a. deep

trough which is bounded by the raised upper occludent margin ;

three further ridges, not to be seen in some specimens, extend

from the umbo to the basal margin, one to the basilateral angle,

a second to one-third of the distance from the rostral angle, and

a third to midway between the two. On the inner surface there

is a deep pit for the adductor scutorum; the inner occludent

mar. gin is much thickened, and above the nae a portion of the

valve is bent downwards at right angles to the outer surface in

some specimens, and in others it is sharply bent inwards; when

the occludent margin is almost in line with the umbo, this

downwardly and inwardly bent portion of the valve almost

entirely overhangs the pit for the adductor scutorum, and forms

a roof to the upper part of the inner surface.

Tergum (Pl. XCIV. fig. 7) subrhomboidal, slightly convex

transversely, with a feebly marked curved ridge extending from

the apex to the basal angle at about one-fifth the width of the

valve from the carinal margin, and from this ridge the valve

slopes steeply to the carinal margin ; upper half of valve curled

towards the scuta, and in most valves away from the opposing

tergum; apex acute; basal angle sometimes acute, and sometimes

slightly and obliquely truncated; carinal margin convex when

continuous, but when formed of two almost equal portions, as is

usually the case, the upper margin is convex and the lower margin

almost straight; scutal or lateral margin straight, almost equal

in length to the concave occludent margin, which has a slightly

raised border; the shape of the upper carinal and occludent

margins is influenced by the extent to which the upper half of

the valve is curved towards the scuta, a character which is

very variable. Along the occludent margin, especially in those

examples conspicuously bowed away from the opposing tergum,

a portion of the valve is bent downwards and sharply inwards to

form an obtusely angular ledge on the inside of the valve; this

ledge, as well as that formed on the inner occludent margin of

the scutum, was no doubt for the protection of the animal’s body,

FOSSIL CIRRIPEDES. 941

for these ledges would serve to fill up the hiatus made through

the upper portions of the scuta and terga being bowed away

from each other. In some of the terga from the Chalk Marl of

Cambridge and Burham, the adoccludent portion of the valve is

not bent downwards and inwards to form a ledge on the inner

surface, but extends outwards to form a second raised ridge

almost parallel with the raised occludent margin. The terga from

the B. plena- marls have this outer ridge finely denticulated on

its margin.

Upper latus (Pl. XCIV. fig. 8) subtriangular, slightly curved

towards the scuta, almost flat : umbo slightly projecting ; at the

scutal margin a narrow slip of the valve is bent downwards, and

during the thickening and growth of the valve a slight ledge has

been formed which has a tendency to project beyond the umbo ;

tergal margin slightly convex; scutal margin slightly concave,

and about the same length as the tergal margin: basal margin

marked off into three almost equal lines, and to the angles thus

made two feebly marked ridges extend from the umbo.

Rostrum (Pl. XCLV. fig. 9) diamond-shape, bowed inwards; a

strong rounded keel extends from the apex, widens gradually to

the basal angle, and the portion of the valve on each side slopes

steeply towards the lateral margins ; apical portion acute; basal

portion rather less acute than the apical portion, with the basal

angle slightly rounded. The inside of the valve is deeply concave.

Structure and Affinities—The capitulum of Zeugmatolepas

mockleri (see restoration, Pl. XCIV. fig. 14) closely resembles

that of Pollicipes, especially i in the munmiber of valves, but differs

(1) in the more specialized form of seutum which has a subcentral

umbo, (2) in the size and position of the upper latera, which are

elevated to occupy the whole of the interval between the scuta

and terga, and (3) in the capitulum being longer than broad, and

therefore of a more erect and Scalpellwm-like shape. In this

form the umbo of the scutum appears generally to be almost

apical in young valves, and to become further removed from the

apex with age, until it reaches about one-third of the distance of

the valve from the apex. Some individuals, however, have the

umbo further removed from the apex than others of the same

size. All the valves are exceedingly variable, much more so than

in any fossil pedunculate Cirripede with which I am acquainted ;

these variations are noted under the descriptions of the several

valves.

Zeugmatolepas mockleri agrees more closely with S. (?) crete

Steenstrup* sp. from the Upper Senonian of Denmark than with

any other species, and, indeed, S. (?) crete may quite well have

been directly developed from 7. mockleri. Through the kindness

of Dr. J. P. J. Ravn I have been able to examine examples of

the known valves (scuta, terga, and carina) of S. (2) crete from

* Steenstrup, J., Kroyer’s Neturinete Tidsskrift, Bd. i. 1837, p. 359; Bd. i,

1839, p. 399, pl. v. figs. 1-3; Darwin, C. R., Pal. Soc. Mon Foss. Lepadidee, 1851,

p. 45, pl. i. figs. Ll a-e.

Proc. Zoou. Soc.—1913, No, LXITI, 63

942 MR. T. H. WITHERS ON

the collection of the Mineralogical Museum of the University of

Copenhagen, and Dr. Ravn informs me that this species has been

found only in the Upper Senonian (Zone of Scaphites constrictus).

Darwin doubtfully referred S. (%) crete to the genus Scalpellum,

and observed (1851, p. 45): “ Protessor Steenstrup was originally

inclined to believe that the capitulum [of SS. (?) crete] was formed

of only five valves; could this be proved, the species would very

naturally rank with a small recent one from the Island of

Madeira, which, owing to the upward growth of the scuta, and to

certain peculiarities in the animal’s body, I have felt myself com-

pelled to raise to the rank of a genus, under the name Oxynaspis.”

An examination of the valves from Copenhagen University, and

of some valves obtained by myself from the matrix of a specimen

presented to the Geological Department of the British Museum

by Copenhagen University, shows, at any rate, that we can no

longer hold Steenstrup’s view that S. (?) crete had only five valves.

Among the above valves are some small lower lateral plates,

which, although not so well preserved as those from the English

Cenomanian Chalk, obviously belong to the same general type as

those of Zeugmatolepas mockleri. WS. (%) eretce, together with the

species now described, is included, for the reasons previously

given, in the new genus Zeugmatolepas.

The scutum of Z. crete differs from that of Z. mocklert in that

the umbo is further removed from the apex, being situated almost |

midway on the occludent margin, and in a greater portion of the

valve near the basi-lateral angle being sharply upturned almost at

right angles to the middle part of the basal margin. The lateral

(or tergal) portion of the valve is not steeply inclined from the

umbo, and there is no deep trough, as in’ Z. mockleri, running

parallel with the upper occludent margin. These and other difter-

ences result, no doubt, from the difference in position of the umbo,

and the consequent greater development of the tergal or lateral

portion of the valve. The carina in Z. cret@ is relatively wider

than in Z. mockleri, the central ridge is much more obscure, and

in the tergum there is a much wider groove near the occludent

margin. The whole of the valves in Z. crete are appreciably

thinner. These differences, however, while deserving of specific

distinction, appear to result from the further development of

valves like those of Z. mockleri, and consequently I consider these

two species to be closely and probably directly related.

Genus CALANTICA.

1825. Calantica Gray, ‘Annals of Philosophy’ (n. s.), vol. x.

jo OM

1907. Calantica Gray: Pilsbry, Bull. U.S. Nat. Mus. no. 60,

Oh (oh

1908. Calantica Gray : Pilsbry, Proc. Acad. Nat. Sci. Philadelphia,

p. 106.

Capitulum with two whorls of valves, the upper comprising

paired scuta, terga, and a carina, the terga occupying the whole

FOSSIL CIRRIPEDES. 943

of the space between the scuta and carina; lower whorl com-

prising three pairs of latera, a rostrum, and a subcarina. Umbo

of all valves apical.

The Oriental group, called by Pilsbry (1908) Calantica s. str.,

has the valves of the lower whorl low and wide, small, not

concealing the bases of the valves of the upper whorl.

TITANOLEPAS *, subgen. nov.

Valves of lower whorl large, high, but not incurved, and over-

lapping the bases of the valves of the upper whorl, Umbo of

scutum subcentral and of the remaining valves apical.

Genotype.—Titanolepas tuberculata.

CaLAnTICA (TITANOLEPAS) TUBERCULATA Darwin sp. (PI. XCY.

figs. 1-10.)

1851. Scalpellum tuberculatuwm C. R. Darwin, Pal. Soc. Monogr.

Foss. Lepadide, p. 43, pl. 1. fig. 10.

1854. Scalpellum tuberculatum C. R. Darwin, Ray Soe. Monogr.

Sub-class Cirripedia, Balanidz, Synopsis et Index Syste-

maticus, p. 634.

1854. Scalpellum tuberculatum C. R. Darwin: J. Morris, Cat.

Brit. Foss. 2nd ed. p. 97. :

1865. Scalpellum tuberculatum C. R, Darwin: J. W. Salter &

H.. Woodward, Cat. & Chart Foss. Crustacea, p. 27, pl. 1.

ime. 0

Lot. Sealpelen tuberculatum COC. R, Darwin: H. Woodward,

Brit. Mus. Cat. Brit. Foss, Crustacea, p. 143.

1886. Scalpellum tuberculatum C, R. Darwin: J. Kafka, Sitz.-

Ber. k. bohm. Gesell. Wiss. Prag (1885), p. 565, pl, i,

Tae, “Lc

1887. Geico tuberculatum C, R, Darwin: A, J. Fritsch &

J. Kafka, Crust. Bohmischen Kreidef, p, 6, fig. 9.

1888. Scalpellum tuberculatum C. R. Darwin; A, Peron, Bull,

Soe. Sci, Yonne, vol. xli. (1887) p. 269.

Diagnosis.—Valves with fine ribs radiating from their apices

and standing out as blunt spines at each zone of growth, Carina

strongly convex transversely, with the parietes narrow and

striated. Scutum with the umbo situated about one-third the

extent of the valve from the apex, and with two ridges extending

from the umbo, one to the basi-lateral angle, and the other to near

the rostral angle, Tergum elongately diamond-shape, with a very

acute basal angle. Valves of lower whorl with the inner apical

half flat and strongly marked with growth-lines.

Remarking on this species, Darwin said (1851, p. 43): “Through

the kindness of Mr. Harris, J have examined several valves, which

1 believe to belong to the same species : the specimens were found

in the chalk detritus [of Charing, Kent], and, therefore, may have

come from the Upper or Lower Chalk or Chalk-marl; but more

probably from the Upper Chalk,”

* Tivavos = White earth, Chalk,

944 MR. T. H. WITHERS ON

T. tuberculata has since been recorded by J. Kafka (1886,

p- 965) from the Cenomanian of Kamajk, Bohemia, and by

A. Peron (1888, p. 269) from the Turonian of Yonne, France.

Kafka mentions a single tergum only, and if his figure is Aeon.

it differs in certain “ohamneiers from the original tergum from

Charing. Peron recorded a small and a very large carina, but

gave neither measurements nor figures ; he drew attention to the

fact that Darwin had supposed the species to come probably from

the Upper Chalk, and said that so far as he knew it came from

the ‘Turonian.

T’. tuberculata occurs fairly commonly in the Cenomanian Chalk

Marl near Cambridge, and we have over 150 valves, including

99 valves of the lower whorl. From the Turonian 7Jerebr apna

zone at Whyteleafe, Surrey, we have a single scutum, and from

the Cenomanian Chalk Marl of Burham, Kent, a single rostrum.

Type.—The collection of the late Mr. Wm. Harris from the

Chalk detritus of Charing, Kent, was acquired from his daughter

in 1881, by the British Museum (Natural History). Several

valves of 7’. tuberculata are represented in the collection, three

of which—a carina (I. 14657), scutum (I. 14659), and tergum

(I. 14658)—are mounted together on paper, and with them is a

label bearing the name ‘“ S. tuberculatum” in Darwin’s hand-

writing, These are probably the original specimens figured by

Darwin, but there is no means of proving this since Darwin gave

no measurements of the figured valves of this species. A further

label (not in Darwin’s handwriting) bearing the words “ very

valuable” was with the specimens, but whether this was meant

to indicate that they are Darwin’s original specimens must be

left to individual opinion. I consider them to be the types, and

since Darwin regarded the carina as the typical valve in Scalpellum,

that valve is here taken as the holotype.

Measurements.—The valves of this species are exceedingly

small, and to judge from these, the largest capitulum could not

have measured much more than 5 or 6 mm. in length. Of the

upper series of valves the largest examples measure respec-

tively :—

Carina, I. 15850. Length 4:1 mm.; breadth 1:2 mm.

Scutum, I. 15843. Length (from ‘apex to middle of basal

margin) 2°5 mm.; breadth 2°6 mm.

Tergum, I. 15851. Length 4:3 mm.; breadth 2°6 mm.

Distribution. [¢ Cenomanian]| Chalk detritus: Charing, Kent ;

Cenomanian, Chalk Marl: near Cambridge, and Burham, Kent ;

Cenomanian, Korytzaner Schichten: Kamajk, Bohemia; Turonian,

Terebratulina-zone: Whyteleafe, Surrey ; Holaster planus-zone :

Armeau and Saint-Julien-du-Sault (Yonne), France.

Description of Valves—Valves plainly ribbed longitudinally ;

the ribs are narrow, but become more prominent on the edge of

each zone of growth, where the ribs are thicker and usually stand

out as short blunt spines.

Carina (Pl. XCV. fig. 1) very narrow, either slightly bowed

FOSSIL CIRRIPEDES. 945

inwards, straight, or bowed outwards; tectum strongly convex

- transversely, not carinate; basal margin almost rounded to

bluntly angular; in young specimens, as shown by the periods

of growth, the basal margin was almost straight; parietes slightly

hens inwards, narrow, aie less than half ane width of the

tectum, and marked with almost parallel growth-lines ; inner

surface of valve deeply concave for the lower two-thirds, the

upper third of the valve being solid and marked with lines

of growth, indicating that the valve jrojected freely to that

extent.

Scutum (Pl. XCV. fig. 2) trapezoidal, umbo situated on the

occludent margin about one-third the extent of the valve from

the apex; almost flat to strongly convex ; four-sided, the lateral

or tergal margin slightly longer than the basal margin, with

which “it makes an angle of about 70°; the basal margin ‘much

longer than the lower part of the occludent margin, with which

it makes almost a right angle, the upper and lower portions of

the occludent margin rane in an angle of about 135°; the upper

occludent margin, said Darwin, “ homologically corr esponds with

the tergal margin of the other eretaceous species and with the

upper, nearly straight, portion of the occludent margin in the

tertiary S. magnum and the recent S. vulgare.” The edge of the

upper occludent margin is upturned and thickened, forming a

prominent ridge, with its apex projecting beyond the lateral

margin; a second prominent ridge extends from the umbo to the

basi-lateral angle; a third ridge, barely perceptible in some

specimens, extends from the umbo to the basal margin, near

to the rostral angle. On the inner surface there is a deep

hollow for the adductor muscle ; along the upper occludent margin

the inner surface is slightly raised, which corresponds to the

outer thickened ridge, the space between the two ridges being

concave ; the inner ridge extends to the umbo, whence it becomes

more pronounced and turns inwards, forming a prominent ridge

above the hollow for the adductor muscle.

Tergum (Pl. XCV. fig. 3) elongately diamond-shape, slightly

angular transversely ; the occludent margin is thickened owing

to the presence of a ridge on the outer and inner edge, corres-

ponding with those on the upper occludent margin of the scutum ;

close and parallel to the occludent margin is a ridge which

becomes prominent near the scutal margin and extends beyond

it; a very prominent straight ridge extends from the apex, and

thickens towards the very acute ‘basal angle; scutal and lower

carinal margins almost equal in length, and longer than the

occludent and upper carinal margins, which latter are also of

almost equal length and together make a right angle.

Valves of the lower whorl (Pl. XCV. figs. 4-8). These are

ornamented externally like the scutum, tergum, and carina.

The apical half of their inner surface is almost flat and marked

with lines of growth, showing that these valves must have pro-

jected freely and overlapped the bases of the valves of the upper

946 MR. 'T. H. WITHERS ON

whorl ; the lower haif is considerably concave. Umbo of valves

apical. The valves are much less modified than is the case with

the valves composing the lower whorl in the more typical forms

of Scalpellum, and consequently their position in the capitulum

is not so readily determinable. Seventeen of the valves are

symmetrical, and five asymmetrical. Of the symmetrical valves,

nine have an extremely concave basal margin with produced

lateral angles (Pl. XCV. fig. 4) and probably represent rostra ;

ae others have a much less coneave basal margin (Pl. XCV.

fig. 8) and may represent sub-carine. ‘The valves just mentioned

are thought to be rostra and sub-carine, owing to the pronounced

concavity of the inner part of the valve near the base. The

three remaining symmetrical valves probably represent median

lateral valves (Pl. XCV. fig. 6). They differ from the supposed

rostra and sub-carine in the smaller concavity of the inner basal

part of the valve, and in the growth periods being formed of

two downward curves which meet on the median line, gradually

becoming more obtuse towards the basal margin. Of the asym-

metrical valves, two are subtriangular in shape with a recurved

acute apex, and probably represent carinal latera (Pl. XCV.

fig. 7); the three others, which may be rostral latera (Pl. XCV.

fig. 5), are about twice as wide as long, and are divided un-

equally by a fold extending from the apex to a position on the

basal margin about two-thirds the distance from the rostral

angle, the ‘basal mar gin on each side being concave

Cirencinnre and A Fiiiion, —While Darwin had little doubt that

the scutum and tergumn figured by him belonged to the same

species, he did not feel so certain with regard to the carina, and

he pointed out that this valve had a close general resemblance to

the same valve in Pollicipes rigidus J. de C. Sowerby. A specimen

of 7. tuberculata in the British Museum (Natural History),

registered I. 14656, formerly in the Harris Collection, affords

valuable evidence on this point, for it consists of the carina and

the paired terga in position (PI. XCOV. fig. 9), and proves that

the carina really does belong to the same species. Darwin

stated also that the apex of the carina did not project freely in

this species, but an examination of the carine from Charing,

including the type, and of the numerous carine from the Chalk

Marl of Cambridge, shows that the upper part of the inside of

the valve is much thickened to one-third of its length from the

apex (Pl. XCV. fig. 1c), and this portion is marked with

growth-lines, showing that it must have projected freely to that

extent. Proof of this is afforded by the specimen I. 14656 from

Charing (Pl. XCV. fig. 9), in which the carina and terga are

in position, for the carina there projects freely.

The carina, scutum, and tergum only of 7. twberculata were

known to Darwin, but the specimens from Cambridge include

several other valves which enable us to add much to our know-

ledge of the capitulum. The ornamentation of these valves is so

characteristic that no doubt can be entertained as to their

belonging to this species, Although there areas many as twenty-

FOSSIL CIRRIPEDES. 947

two valves *, other than scuta, terga, and carine, to judge by

their structure, none appear to have formed upper latera, or at

all events to fit between the scuta and terga, as do the upper

latera in most forms of Scalpellum. Moreover, the form of the

scuta and terga and the structure of their inner surface shows

that the tergal margin of the scutum and the scutal margin of

the tergum were in close proximity to each other. In this con-

nection Darwin said (1851, p. 44): “I have no doubt that the

ridge along the upper occludent margin of the scuta, and

that on the occludent margin of the terga, together with

their projecting points, are related to each other, owing to

the close contact of these valves.” It follows, therefore, that if

no upper latera were present, the upper whorl of valves in

T. tuberculata compr ised only five valves, namely, a carina, a pair

of terga, and a pair of scuta. The valves of the lower whorl, as

mentioned in their description, must have overlapped the bases

of the valves of the upper whorl, and if we are correct in desig-

nating these valves as rostrum, rostral latus, median latus,

carvinal latus, and sub-carina, the lower whorl would consist of

at least eight valves, making thirteen valves for the whole

capitulum (see restoration, Pl. XCV. fig. 10).

In the number and arrangement of the valves of the capitulum

Titanolepas tuberculata approaches the forms of Scalpellum

grouped by H. A. Pilsbry 7 under the genus Calantica Gray, and

comes nearest to those oaieeledl by him in the section Scile-

lepas Seguenza, of which the type is S. carinata Phillipi sp.,

from the Pliocene of Messina, Sicily. The capitula of the species

grouped under Calantica closely resemble that of Pollicipes, but

differ in that there is only a single basal whorl of valves, com-

prising three pairs of latera, a rostrum, and a sub-carina. The

species included by Pilsbry under the section Scillelepas are

chiefly characterized by the absence of an upper lateral plate in

the upper whorl, which consequently consists only of five valves,

and by the bases of the valves of the upper whorl being over-

lapped by those of the lower whorl; the umbo in all the valves

is apical. While 7. tuberculata agrees with these species in

having five valves only in the upper whorl, and in the valves of

the lower whorl overlapping those of the upper whorl, it differs

from all the species included under the genus Calantica and the

section Scillelepas by the umbo of the scutum being in a sub-

central position.

We have in 7. tuberculata, therefore, a species agreeing in

all essential characters with the forms of Calantica (Scillcelepas),

except that it has a more specialized form of scutum. The

* Two further valves have since been noticed among the Cirripede material from

the Chalk detritus of Charing, Kent, in the British Museum (Natural History).

These are identical with the valves from Cambridge, which I consider to be rostral,

and are registered I. 14663, I. 14664. A further rostrum has been found in the

Chalk Marl of Burham, Kent.

+ H. A. Pilsbry, “On the Classification of Scapelliform Barnacles,” Proc. Acad.

Nat. Sci. Philadelphia, 1908, pp. 106-107; “The Barnacles _(Civripedia) contained

in the Collections of the U.S. National Museum,” Bull. U.S. Nat. Mus. No. 60,

1907, pp. 8-9.

948 ON FOSSIL CIRRIPEDES,

fossil species that can be definitely referred to Calantica (Scille-

lepas) are found in Tertiary rocks, viz. Scillelepas parone

from the Miocene of the neighbourhood of Turin, Italy, and

S. carinata and S. ornata from the Pliocene of Messina, Sicily,

so that it is an important advance to show that a representative

of the genus existed so far back in time as the Cenomanian. It

is certainly remarkable to find that 7’. twberculata had a more

highly specialized seutum than the Tertiary and recent species,

and to mark this advance in the evolution of the group, I place

it in a new sub-genus, 7itanolepas. This course is the more

desirable since 7. twberculata may represent a side line of develop-

ment from the ancestral Scillelepas line, which, by the sup-

pression of the lower whorl of valves, and by the umbo of the

carina becoming sub-central, may eventually have given rise to

the genus Oxynaspis.

For help in connection with this paper, I wish to express my

indebtedness to Dr. F. A. Bather, Dr. W. T. Calman, Mr. C. P.

Chatwin, and Dr. H. A. Pilsbry.

EXPLANATION OF THE PLATES.

All the specimens here figured are in the collection of the Geological

Departinent of the British Museum.

Prats XCIV.

Zeugmatolepas mockleri Withers, gen. et sp. n.

Fig. 1, Carina. Cenomanian, Chalk Marl: near Cambridge. I. 15820. a, outer

view; b, side view; c, inner view.

2. Carina, I. 15821. a, outer view; 6, side view ; c, inner view.

3, 4, 5, 6. Scuta (various forms), I. 15822-I. 15825. )

7. Tergum, I. 15826. (a, outer view; 6, inner

8. Upper latus, I. 15827. r view.

9. Rostrum, I. 15828. y

10. Several valves in position, including the carina, I. 15829.

11. Holotype. Nearly complete capitulum showing the left side, I. 15830.

12. Several valves in position, I]. 15831.

13. Nearly complete capitulum showing the right side, I. 15832.

14. Restoration of capitulum based on the above material from the Chalk Marl

of Cambridge. ¢., carina; 6., scutum; ¢., tergum ; u.l., upper latus ;

r., rostrum; /.7., lower latera; s.c., sub-carina.

Figs. 1, 2, 4, 9-18, 10 diameters; figs. 3, 5-8, 14, X 5 diameters.

———

Prate XCV.

Calantica (Titanolepas) tuberculata Darwin sp.

. Carina. Cenomanian, Chalk Marl: near Cambridge. I. 15842. a, outer

view ; 6, side view; c, inner view; d, transverse section hear apex.

. Scutum, I. 15843 (figure reversed).

. Tergum, I. 15844.

. Rostrum, I. 15845. |

. Rostral latus, I. 15846. r a, outer view ; 6, inner view.

Median latus, I. 15847. |

. Carinal latus, T. 15848 (figure rever rsed).

. Sub-carina, I. 15849.

. Carina and pair of terga in BART [? Cenomanian| Chalk detritus ;

Charing, Kent. J. 14656.

Restoration of capitulum, based on disconnected valves from the Chalk Marl

of Cambridge. c., carina; s., scutum; ¢., tergum; r., rostrum ; 7./., rostral

Jatus; m.7., median latus; e./., carinal latus; s.c., sub-carina.

Figures 1-10 X 10 diameters.

sae Gene re

Ins Zo So ISIS, Il, XCVI,

BATRACHIDERPETON LINEATUM.

Ins B. SB, UNS, IPM, SCV,

BATRACHIDERPETON LINEATUM.

ON A COAL-MEASURE STEGOCEPHALIAN. 949

55. Batrachiderpeton lineatum Hancock & Atthey, a Coal-

Measure Stegocephalian. By D. M.S. Watson, M.Sc.,

Lecturer on Vertebrate Palzeontology in University

College, London *.

[Received May 20, 1913: Read October 28, 1913.]

(Plates XCVI. & XCVII.,f and Text-figures 162-167.)

INDEX. Page

Miorpliol 0 oye tanta tra his. negroes juice | OO.

Systemiatl ee caer askaeecesd wesc eee ee eee OOO

Amongst the wonderful series of vertebrate fossils in the

Hancock Museum, Neweastle-on-Tyne, derived from the Low

Main Seam of Newsham Colliery, a small stegocephalian Batrachi-

derpeton is represented by excellent material, which is described

as completely as possible in this paper.

SxuLu.—WMaterial.

I. A nearly complete skull with the ornament and palate well

preserved. The type specimen. (Atthey collection.)

II. A complete skull showing with great clearness all the

sutures on the top of the head. (Atthey collection.)

III. A much disarticulated skull of importance in ascertaining

the structure of the palate. (Atthey collection.)

IV. Half of a badly-preserved skull, showing well the maxillary

and palatine teeth. (Dinning collection.)

Shape of the Skull.

All the specimens are crushed flat, so that the shape of the

skull has to be settled by indirect means. It is triangular in

plan, the postero-lateral corners being much produced into long

‘cornua.” Specimen II. shows that there were almost vertical

cheeks, as on the right side this part was turned under and

crushed on to the lower surface of the roof of the skull, whilst

the left side is pressed outwards. This crushing has only very

slightly distorted the palate, and the distance between the two

quadrates is almost identical with that between the spot where

the cheek is turned under on the right side and the corresponding

point on the left side.

There is a pineal foramen of medium size almost exactly half-

way between the point of the nose and the back of the head in

the middle line. The orbits are laterally placed and lie entirely

in front of the middle of the skull. The small anterior nares are

also laterally placed.

The structure of the cranial roof will be best understood from

* Communicated by Prof. J. P. Hin, D.Sc., F.R.S., F.Z.S.

+ For explanation of the Plates see p. 962.

950 MR. D. M. S. WATSON ON A

text-figs. 162 & 164. All the sutures drawn with solid lines in

these figures can be clearly seen in specimen IT.

The interesting features are the great development of the

Text-fig. 162.

Batrachiderpeton lineatum. Dorsal aspect of the restored skull and clavicular

apparatus. X13. Skull mainly founded on specimen II., from which all the

sutures are taken, and the pectoral girdle restored from the bones of specimen III.,

which is a smaller individual, enlarged to correspond in size with the skull.

Reference letters :—Cu., Clavicle; Cm., Cleithrum; Fr., Frontal; I.Cu., Inter-

clavicle; Lac., Lachrymal; Mx., Maxilla; P.Mx., Premaxilla; P.O., Post-

orbital; P.Par., Postparietal; Par., Parietal; Po.FR., Postfrontal; PR. FR.,

Prefrontal ; Sq., Squamosal; Tas., Tabulare.

COAL-MEASURE STEGOCEPHALIAN, 951

horns, formed entirely by the squamosals and tabulares, and

the correspondingly large size of the post-parietals.

The extremely small size of the nasals is an interesting feature.

The sutures separating the lachrymals from the premaxille are

not certain, but there is a separation and displacement of the

bones at the dotted line which has probably taken place along

the suture.

Text-fig. 1635.

Batrachiderpeton lineatum. , Ventral aspect of the restored skull and clavicular

apparatus. X13. Skull mainly from specimen I., sutures from specimen III.,

clavicles, etc., from specimen III.

Reference letters as before with :—Quv., Quadrate; Pat., Palatine; PaR.Oc., Pare

occipital process; Par.Sp., Parasphenoid; Pr.V., Prevomer; Pr., Pterygoid.

952 MR. D. M. 8. WATSON ON A

The palate is of great interest; in the restoration the general

features are drawn from specimens I., I1.,and 1V., and the sutures

are introduced from specimen ITI., where the pterygoids, palatines,

and prevomers are disarticulated.

Basis cranii.—The basis cranii is much crushed in all the

specimens and was apparently very thin and delicate.

The two large exoccipital condyles are clearly shown in I.

and Il. They are rounded from front to back and to a less

extent from side to side. The bone between them is extremely

thin, but no doubt is partly composed of basioccipital. The basi-

sphenoid in I. is slightly better preserved ; it bears two very small

basipterygoid processes only very obscurely indicated. In front

of this region it is produced into a narrow rostrum undoubtedly

parasphenoid.

Extending outwards and backwards from the sides of the basis

cranii are two long, very delicate processes which are shown,

much crushed and broken, in specimens I., II., and III. In

specimen I. on the right side, a small foramen is rather obscurely

indicated on the process about a third of its length from the

middle line, this opening can only be the fenestra ovale. The

process is probably mainly opisthotic, with the prootic forming

some of its front border. Immediately above the exoccipital

condyle in the same skull is a foramen for cranial nerves ix.

and x.

The outer end of the opisthotic articulates as usual with a facet

on the under surface of the tabulare.

The quadrates are well shown in specimens J. and It. They

are small bones with a well-developed trochlear condyle, the outer

border is articulated with the quadrato-jugal, there being

probably no foramen between the two bones.

The upper end of the quadrate is supported by a downwardly

directed flange from the squamosal, well seen in specimens I.

and III. The inner border of the quadrate is continued by

a short thin pterygoid process (specimen I.).

The pterygoid isa large bone which articulates by a small facet

with the basipterygoid process of the basisphenoid (specimen I.).

Behind this it is continued as a deep but thin plate, the posterior

ramus, which articulates distally with the quadrate and whose

upper border has a suture with a continuation of the descending

flange of the squamosal mentioned above. The anterior part of

the bone forms a broad flat plate covered with a shagreen of small

sharp teeth. The anterior border of the plate is in contact with

the prevomer and the lateral border with the palatine.

The palatine bears a single row of nine medium-sized swollen

teeth, and its outer border articulates with the jugal and maxilla.

There is apparently no transpalatine.

The prevomer is a small bone with a median symphysis with

its fellow; it forms a good deal of the inner border of the posterior

nares and bears two teeth placed transversely ; its exact relations

to the palatine are not certain.

COAL-MEASURE STEGOCEPHALIAN. 953

The premaxilla appears to bear three teeth, but the suture

separating it from the maxilla with six teeth is not quite certain.

Text-fig. 164.

Batrachiderpeton lineatum. Weft lateral aspect of the restored skull and clavicular

apparatus. Skull from specimen IT., clavicle, etc., from specimen III. 1.

Reference letters as in former figures with :—Jv., Jugal; Qu.Ju., Quadrato-jugal.

Lower Jaw.—Waterial.

Specimen III. The perfect left ramus and disarticulated parts

of the right ramus of the same individual. Although the

posterior part of the right ramus, composed of the angular,

surangular, and prearticular, is not well preserved, this specimen

is of great importance because it gives the most satisfactory

evidence of the front of the mandible, the dentary and splenial

being slightly separated and completely free from matrix on

the outer and largely so on the inner side.

Specimen V. <A perfect right ramus showing well the suture

between the prearticular and the angular. (Atthey collection.)

Specimen VI. A perfect right ramus. (Barkas collection.)

Specimen VII. An isolated dentary. (Atthey collection.)

The various specimens are crushed in different directions and

suggest that the jaw had a considerable cavity and was perhaps

nearly circular in cross section.

There is a distinct though short postarticular process.

The articular is well ossified though small; it bears an articular

facet adapted to the trochlea quadrati.

The suture between the surangular and angular is not clearly

shown in any specimen. The angular is a very large bone

extending nearly up to the symphysis; it forms the lower part

of the jaw, and its inner border articulates by a suture (spe-

cimen V.) with the prearticular, from which it is separated by

a smali oval internal mandibular vacuity.

‘The prearticular 1s a large bone covering the inner side of the

articular and extending well forward.

The dentary is a fairly large bone overlapping the surangular

and angular behind; it bears eleven or twelve swollen teeth and

has a symphysis with its fellow; its lower border has a suture

954 MR. D. M. S. WATSON ON A

with the splenial. The splenial has a symphysis with its fellow

and is entirely a bone of the outer side of the jaw, as is

conclusively shown in specimen ITI.

Text-fig. 165.

Batrachiderpeton lineatum. Right ramus of the lower jaw, outer aspect. X 2.

Outline and sutures from specimen ITT.

Reference letters:—Ane., Angular; Den., Dentary; Sp., Splenial; Sur.Ane.,

Surangular.

The complete jaws show extremely well a great patch of small

conical teeth lying inside those of the dentary. Specimen III.

shows conclusively that these do not belong to the dentary, the

splenial, the angular, or the prearticular, so that it is quite

certain that there is another element, which can only be the

coronoid.

Text-fig. 166.

Batrachiderpeton lineatum. Left ramus of the lower jaw, inner aspect. X 2.

Drawn from specimens III., V., & VI.

Reference letters as on text-fig. 165 with :—Cr., Coronoid; Pr.Arv., Prearticular.

Between the coronoid, prearticular, and angular there is a small

vacuity.

Pectoral girdle.

Specimen ITI. retains both clavicles and a cleithrum.

The clavicle is composed of a flat ventral plate with an orna-

ment of strong radiating ridges, from the upper surface of which

within the lateral border an almost cylindrical process rises

abruptly, this slopes slightly backwards and outwards and gives

attachment to the cleithrum. The dorsal surface of the ventral

plate of the clavicle shows clearly the impression of the part of

COAL-MEASURE STEGOCEPHALIAN. 955

the interclavicle which overlaid it, and enables the width of the

latter bone to be accurately determined.

Text-fig. 167.

Ceraterpeton reticulatum H. & A. Restored dorsal surface of skull and

clavicular arch. X 4.

The cleithrum is a T-shaped bone, flat, and ornamented by

fine grooves, the upright of the T is affixed to the vertical process

of the clavicle.

Discussion of the Skull.

One of the most interesting features of the skull is the reduc-

tion of the number of bones in the temporal region. The

primitive amphibian has in the temporal region three bones, the

squamosal, supratemporal, and intertemporal, all of which occur

in the primitive reptile Seymouria. It is important to discover

which of these bones is really present in Batrachiderpeton.

In the primitive amphibians “ Lowomma” and Pteroplaa all

three bones are present in a typical manner and also the

squamosal; that is, the outer bone, which articulates with

the quadrato-jugal, is well characterized and differentiated frem

the other two by the fact that its posterior border is bent down

in front of the otic notch and articulates with the upper edge of

the quadrate ramus of the pterygoid, it also passes down behind

the quadrate itself, overlapping a good deal of its posterior

surface. Neither the intertemporal nor supratemporal has any

connection whatever with the pterygoid or quadrate.

The conditions in these primitive amphibia are,those which

956 MR. D. M. S. WATSON ON A

always occur in the large Stegocephalia, modified of course by

changes in the shape of the skull. I have seen them in :—

“« Loxomma,” Pteroplax, Batrachosuchus, Cyclotosaurus, Plagio-

sternum, Trematosaurus, Micropholis, Capitosaurus, Aphaneramma,

Bothriceps, Archegosaurus.

In Batrachiderpeton the squamosal is united to the pterygoid,

and covers the back of the quadrate, not by the turning down

of its hinder edge but by a special descending flange, a difference

depending entirely on the development of the ‘‘ horn” which has

carried the otic notch, which, in all amphibia and reptiles where

it is present, lies between the tabulare and the squamosal, far

out and backwards. The great sheet of squamosal which les

behind the quadrate is in fact a new development, the original

posterior border being now represented by the flange.

There is hence no doubt that the single temporal bone re-

maining in Batrachiderpeton is the outer of the original three.

That this is really the homologue of the mammalian squamosal,

I hope to be able to show in a subsequent paper. The other

bones of the top of the skull do not call for special notice. ;

The palate of Batrachiderpeton is of very great interest. I

pointed out recently that the type of palate found in ““Loxomma,”

with a large single basioccipital condyle, basipterygoid processes

on the basisphenoid, and large pterygoids, which leave only a very

small interpterygoid vacuity divided by a narrow parasphenoid, is

the primitive type of amphibian palate, because it agrees very

closely with that of the Crossopterygian fish, and because no

palate with large parasphenoid and interpterygoid vacuities is

known to occur in the Coal-Measures or Lower Carboniferous.

From known types it is possible to pick a morphological although

non-phylogenetic series showing exactly how the change from the

primitive palate to the typical palate of Capitosaurus has taken

place. Hryops, which, as shown by v. Huene’s recent extremely

interesting paper, has a quite large basioccipital and basipterygoid

processes of the basisphenoid, and in which the parasphenoid and

interpterygoid vacuities are only of moderate size, is exactly

intermediate between the primitive type and that found in the

Triassic forms, where the basioceipital and basisphenoid are very

much reduced and the pterygoids are supported by sutural

union with the edges of the parasphenoid. The palate of Batra-

chiderpeton agrees with the primitive type, in having large

pterygoids meeting in the middle line and articulating with

basipterygoid processes of the basisphenoid, It differs, however,

in that it has already completely replaced the basioccipital con-

dyle by a pair of exoccipital condyles, just as have the later

large Stegocephalia.

Iscussion of the Lower Jaw.

Except that it is not known whether an epicoronoid was present

or not, the lower jaw of Batrachiderpeton is extremely like that

COAL-MEASURE STEGOCEPHALIAN. 957

of “Loxomma” in its structure, particularly in the fact that the

splenial is entirely a bone of the outer side of the jaw, which

is proved in the most conclusive manner by specimen III.

In later Stegocephalia the splenial (as shown, for example,

in Lryops, Anaschisma, Bothriceps, and “ Labyrinthodon ” lepto-

gnathus), although it still has a good exposure on the outer

surface, has also a large flange running up on the inner side of

the jaw to the coror.oid

In Reptilia the internal part of the splenial is always larger

than the external, and in later reptiles this latter part disappears

altogether.

It thus seems certain that the trend of evolution of the splenial

has been directed to gradual growth of an internal flange at the

expense of the external part of the bone which alone occurs in

the primitive amphibia.

In correlation with this development is a reduction of the

coronoid and a complete loss of the epicoronoid.

As the angular is to all appearance a serial homologue of the

splenial, it is probable that it may have tended in early Amphibia

to pursue a similar course of change, in which case we may

suppose that in primitive forms the suture between the angular

and the prearticular probably ran along the lower border of the

jaw just as does the suture (when there is one) between the

coronoid and the splenial.

This type of jaw, which we have arrived at hypothetically, is

actually realized in Crossopterygian fishes, Megalichthys being the

best type.

The structure of the jaw in Paleozoic Crossopterygians was first

accurately made out by Traquair in Rhizodus and Rhizodopsis.

I give here an account of the jaw of Megalichthys and have also

examined those of Holoptychius and Osteolepis, which agree in

general structure.

In Megalichthys the dentary is a large bone narrowing from

front to back, where it ends in a point. It bears throughout its

border a single row of small pointed teeth, within which at the

anterior end are one or two large “laniary” teeth. Its lower

border overlaps three bones, the splenial, angular, and sur-

angular.

The splenial has an articulation with its fellow; it lies entirely

on the outer side of the jaw and its lower border is entirely free.

Its oblique posterior edge overlaps the angular.

The angular is a bone similar to the splenial, except that it

has of course no symphysis and its lower border articulates with

the coronoid. a

The surangular is overlapped by the angular, which it greatly

resembles ; it covers the outer side of the well-ossified articular.

* The combined prearticular and coronoid is a very large bone

running forward from the inner side of the articular nearly to

* A specimen of Glyptolepis paucidens in the Manchester Museum seems to

show a clear suture between the short prearticular and the coronoid.

Proc. Zoou., Soc.—i913, No. LXIV. 64

958 - MR. D. M. 8. WATSON ON A

the front of the jaw ; in front it ends freely in a point, its lower

border articulates with the angular and surangular, and its

upper border with the epicoronoids; the inner surface of the

bone is covered with a shagreen of small denticles.

The epicoronoids are two small bones lying between the upper

edge of the coronoid and’ the dentary. The posterior is a short

bone forming the front of the suprameckelian vacuity, and

bearing two Tar ge teeth and the pits for their replacing teeth. It

is separated from the anterior epicoronoid by a large round

foramen. ‘The anterior bone bears one tooth and its replacing

tooth-socket.

The jaw of Megalichthys differs slightly from that of Ahizodus,

Rhizodopsis, etc., in there being only three bones besides the

dentary on the outerside of the jaw, instead of four, and only two

epicoronoids instead of three.

Comparison of this Jaw with that of “ Zovomma” will, I think,

justify the nomenclature used here, even the direction of overlap

of the bones being the same in the two types.

Systematic position of Batrachiderpeton.

The classification of the smaller stegocephalian Amphibia, so

abundant in the Coal-Measures and Permian Rocks of Europe

and North America, is in such confusion, to which some

recent work has added, that it is at present only possible to

proceed by reference to individual specimens which have been

well described. The difficulty of identifying species and the

rashness with which genera often founded on very imperfect

materials have been extended to include other types often of

very distinct structure, combined with the great technical

difficulties of working on small and often badly-preserved

animals, make any wide divisions at present of very doubtful

value.

Batrachiderpeton in the forward position of its orbits and the

ereat ‘“cornua” at once recalls Ceraterpeton, originally described

by Huxley from Kilkenny. The type-specimen of this latter

genus is badly preserved, so that the sutures of the cranial roof

are not clearly shown, although it seems probable that the

apparent sutures are real, in which case there are many resem-

blances between the two types.

To the type-species Ceraterpeton galvani two other specimens

have been assigned—Ward’s specimen from North Staffordshire

described by C. W. Andrews, and a specimen from Kilkenny

figured by A. Smith Woodward. These two specimens agree

extremely closely in size and in the shape of the skull, but they

differ from the type in being only about one-half of the size and

in having a narrower skull. In the Hancock Museum there are

three specimens closely similar from the Low Main seam of

Newsham, one of which is the type of Hancock and Atthey’s

COAL-MEASURE STEGOCEPHALIAN. 959

Urocordylus reticulatus. These three specimens agree very closely

in measurements and in all characters which can be observed ;

they are undoubtedly cospecific. They also agree exactly with

Andrews’ and Smith Woodwaxd’s specimens in the proportions of

the skull and body, and in size, but differ from Andrews’ specimen

very markedly in the character of their cranial ornament, which

is almost entirely composed of ridges instead of pits. They must

hence be regarded as specifically distinct.

In text-fig. 167 (p. 955) I have given a drawing of the top of

the head with such sutures as are clearly visible in one of the

Neweastle specimens preserved in counterpart; this should be

compared with Dr, Andrews’ figure. It is certain that the ptery-

goids met in the middle line, as in Batrachiderpeton, although

the detailed structure of the palate cannot be made out.

One important feature, shown clearly by the type-specimen of

‘“« Urocordylus” reticulatus, is the T-shaped cleithrum, also shown

in Dr. Woodward’s figure, where it is interpreted as scapula.

Such a shaped bone is known in no other animal except Batrachi-

derpeton, and seems to show definitely that ‘* Urocordylus”’

reticulatus and Datrachiderpeton are closely related. ‘The very

remarkable clavicle is also identical in the two types.

Whether Urocordylus reticulatus H. & A. really belongs to the

genus Ceraterpeton is uncertain, but it is probable that it does ;

in any case, by calling it Ceraterpeton reticulatwm in full, no

trouble will arise ; it seems probable that it is not Urocordylus.

One of the most marked features of Ceraterpeton is the ex-

pansion and corrugation of the distal ends of the neural and

hemal spines; the meaning of this condition, which occurs

throughout the column, is obscure. It also occurs in Ceraterpeton

galvant, Urocordylus wandesfordii, Scincosaurus crassus, Ptyonius

estocephalus, and other types. As there is no definite evidence

that these types cannot be related, 1¢ is natural to assume that

they are; but it must be remembered that any such relation

can only be slight, for if Fritsch’s restoration of the skull of

Scincosaurus crassus is at all correct, it differs very greatly from

C. reticulatum and Batrachiderpeton.

Another type which appears to be related is Diceratosaurus

punctolineatus Cope, recently deseribed by Jekel. The palate

of this type is extremely like that of Batruchiderpeion, differing

in minor features of the dentition, in the larger interpterygoid

vacuity, and in the presence of a transverse bone.

If we suppose, as is quite probable, that some of the sutures

of the cranial roof were invisible, it is probable that the skull

structure was very like that of Ceraterpeton reticulatum. Ou the

other hand, this type has not the T-shaped cleithrum and peculiar

neural arches of the latter form.

Assuming, as seems justifiable, that Batrachiderpeton is allied

to Ceraterpeton reticulatum, it is of interest to see the differences

between the two types. Ceraterpeton reticulatwm is obviously the

64*

960 MR. D. M. 8. WATSON ON A

less modified, and the following list shows the direction of change

in passing to Batrachiderpeton :—

1. Increase in the relative size of the “ cornua”’ involving

larger squamosals, postparietals, and tabulares.

2. More anterior position of the orbits.

3. Great decrease in size of the nasals.

4, Broadening of the clavicles and interclavicle.

The most interesting comparison, however, is with Diplo-

caulus, a type which Jzkel has already brought into relation

to Ceraterpeton.

Comparison of text-fig. 162 (p. 950), the dorsal surface of the

skull of Batrachiderpeton, with the figures of the skull of

Diplocaulus given by Williston & Case, shows a very striking

resemblance, particularly in the way in which the horn is

developed, by enlargement of the squamosal (prosquamosal of

Williston & Case), tabulares, and postparietals (supra-occipital

plate of Williston & Case).

If the bone regarded by Williston as a nasal is compared with

the undoubted lachrymal of Batrachiderpeton no doubt of its

identity can arise, it being evident that the nasals, extremely

small in Batrachiderpeton, are quite lost in Diplocaulus.

It also seems extremely probable that the bone called

squamosal by Williston & Case is really the postorbital dragged

out from the border of the orbit by the development of the

horn.

The only other important difference shown on the top of the

skull lies in the fact that in the earlier type the frontal is

excluded from the orbit, whereas in the American form it forms

much of the border of that opening. ‘This depends on the

different position of the orbits, which in Batrachiderpeton are

laterally directed, whilst in Dzplocaulus they look directly

upwards. It is in every way probable that this difference

depends entirely on the habits of the animals in question.

The skull of Batrachiderpeton shows no grooves for lateral line

organs, and it seems likely that the animal was not very

aquatic ; and as its skull was probably of only moderate size, it

is possible that it pursued an ordinary life.

Case, apparently justly, regards Diplocaulus as living in the

mud at the bottom of ponds, and its head is so enormous

that it must have been carried flat on the ground. Such a habit

of life is inconsistent with laterally placed eyes, which would

therefore necessarily migrate to the dorsal surface as in all flat-

bodied bottom-dwelling fishes, e. g., Lophius, Raia, Nolea.

The clavicular apparatus of Diplocaulus is very similar to that

of Batrachiderpeton. The clavicles in both types consist of a flat

ventral plate, from the upper surface of which a cylindrical

process rises dorsally within the lateral edge.

The vertebral column of Diplocaulus is now well known. The

COAL-MEASURE STEGOCEPHALIAN, 96]

presacral vertebra, 16 in number, have an elongate hour glass-

shaped “centrum ” and elongated neural arches with low elong gate

spines. There are two processes for the rib, the upper Grmied by

the arch, the lower by the “centrum.” There are hypapophysial

in addition to the ordinary zygapophysial articulations. The

caudal vertebree have no rib articulations and expanded hemal

arches fused into the “ centrum.”

The vertebral column of Ceraterpeton is not well known, but

the Newcastle specimen of C. reticulatwm seems to show the

following characters:—'There are about 18 presacral vertebre

with hour-glass-shaped centra, long neural arches and spines

almost eontainly with hypayophysial i in addition to the ordinary

articulations, and a single process neaily in the centre of the

vertebra for the rib; this may be partly carried by the arch.

The later caudals differ in having no rib articulation and in

having large expanded hemapophysial spines fused onto the

centrum.

The vertebral columns of the two types are, in fact, very

similar, differing apparently only in the fact that in Diplocaulus

the head of the cba is double, whereas in Ceraterpeton reticulatwm

it 1s single.

The shortness of the neural and hemal spines of Diplocaulus

may be due merely to the flatness of that animal.

The lower jaw of Diplocaulus as described by v. Huene agrees

very well with that of Batrachiderpeton, the inner row of teeth

at the front being probably borne on the real coronoid, and

. Huene’s complementare being an epicoronoid.

In fact, the only important difference between the two types

lies in the palate. Dzplocaulus differs in its palate from Batrachi-

derpeton exactly as does Hryops from Loxomma.

The dentitions in the two types are identical in general lines.

The differences are that in Batrachiderpeton there are large

pterygoids meeting in the middle lne and supported by the

basisphenoid, in Diplocaulus there are large interpterygoid

vacuities and the pterygoids are supported by the parasphenoid.

I have already endeavoured to show that in the large Stego-

cephaha this change actually takes place, a view which if correct

removes all ditticulty of regarding Batrachiderpeton and Diplo-

caulus as very closely allied.

Finally, it may be noticed that the characters in which

Diplocaulus differs from Batrachiderpeton are exactly those in

which the latter differs from Ceraterpeton :—

1. Increase in relative size of the cornua, involving larger

squamosals, postparietals, and tabulares.

More anterior position of the orbits (and their dorsal

position).

3. Complete loss of the nasals.

4. Broadening of the clavicles and interclavicles.

962 ON A COAL-MEASURE STEGOCEPHALIAN.

I wish to express my thanks to the Committee of the

Northumberland and Durham Natural History Society and to

K. L. Gill, Esq., Curator of the Hancock Museum, for the

kindness with which they have allowed me to work on the-

wonderful material in their Museum.

BIBLIOGRAPHY.

AnpreEws, C. W.—Note on a Specimen of Keraterpeton galvani

Huxley, from Staffordshire. Geol. Mag. new ser. Dee. iv.

vol. 11. pp. 81-84. 1895.

Bror, EF. v.—Hin Beitrag zur Kenntniss von Diplocaulus

Cope. Centr. f. Min. Geol. u. Pal. 1902, pp. 536-541.

Casn, KE. C.—Revision of the Amphibia of the Permian of

North America. Carnegie Institution, Washington, pub.

no. 146, 1911.

J @KEL, O.— Ueber Ceraterpeton, Diceratosaurus u. Diplocaulus.

Neues Jahrb. f. Min. Geol. u. Pal. 1903, Heft 1, pp. 109-

134.

Huxtry, 'T. H.— Description of the Vertebrate Remains from the

Jarrow Colliery. Proc. Roy. Irish Acad. vol. xxiv. p. 353.

1867.

Traquair, R. H.—On the Structure of the lower jaw in Rhizo-

dopsis and Rhizodus. Ann. & Mag. Nat. Hist. ser. 4,

vol. xix. pp. 299-306. 1877.

Watson, D. M. 8.—-The larger Coal-Measure Amphibia.

Manch. Mem. vol. lvl. pt. 1, pp. 1-13, pl. & figs. 1912.

Wuuurston, 8. W.—The Skull and Extremities of Diplocaulus.

Trans. Kansas Acad. Sci. 1909, pp. 123, 131.

Woopwarp, A. S.—On a New Specimen of the Stegocephalian

Ceraterpeton galvani. Geol. Mag. new ser. Dee. iv. vol. iv.

1 AB, LOA

Hancock, A., and Arrnry, T.—Description of a Labyrinthodont

Amphibian, a New Generic Form, obtained in the Coal

Shale at Newsham, near Newcastle-upon-Tyne. Ann. &

Mag. Nat. Hist. ser. 4, vol. vi. pp. 56-65, pl. 1. 1870.

EXPLANATION OF THE PLATES.

Prats XCVI.

Fig. 1. Skull of Batrachiderpeton lineatum H. & A. Specimen I. (type-specimen),

14. Dorsal aspect.

2. Same specimen as fig. 1. Palatal aspect.

3. Rami of the lower jaw of Batrachiderpeton lineatum, X 2.

A. Specimen VI. Right ramus, outer aspect.

B. Specimen III. Left ramus, inner aspect, showing very clearly the patch

of teeth on the coronoid.

Prare XCVII.

Fig. 4. Skull of Batrachiderpeton lineatum H. & A. Specimen II., X 1. Dorsal

aspect.

5, Same specimen as fig. 4. Palatal view

West, Newman

1-4. PEACHIA QUINQUECAPITATA.

5-7. BICIDIUM AQUOREZ 8.B.PARASITICUM.

ON NEW ACTINIANS FROM BRITISH COLUMBIA. 963

56. On two New Actinians from the Coast of British

Columbia. By J. PLayrarr McMourrtcu, C.M.ZS.

[Received August 1, 1913: Read November 11, 1913.]

(Plate XCVITI.*)

INDEX.

Systematic :— Page

FZCACHIOAGUUNO MECH ILC, ISP ep Mey ersceee steelers eects seieeeect = 963

Bi cidenmeguoncces Spa Um ceseeeentaee eee een OO

The two forms described below were obtained during a visit to

the Marine Station maintained by the Canadian Government at

Departure Bay, Vancouver Island. They are of especial interest

from the probability that they represent stages of a single species,

and belong to a group that has not yet been described as occurring

on the west coast of North America. Since the specific identity of

the two forms is only a probability, it has seemed well to regard

them for the present as distinct, and even, for reasons given

below, to assign them provisionally to different genera.

PEACHIA QUINQUECAPITATA, sp. n. (PI. XCVITI. figs. 1-4.)

This form (fig. 1) was dredged by Dr. C. McLean Fraser

in Nanoose Bay, Vancouver Island, in 15-20 fathoms. Jn the

majority of the individuals the base is depressed in the centre

and smaller than the column. It is thin, and in all cases

shows clearly the lines of insertion of the mesenteries as well-

marked invections, so that it can have possessed but very slight

adhesive powers, if any. In one individual the attachments of

the mesenteries did not quite reach the centre of the base, but

ended abruptly a short distance from it, leaving a circular central

~area which was exceedingly thin, but nevertheless not perforated.

In other examples this condition did not occur, the insertions

of the mesenteries extending quite to the centre, so that there

were no indications of a terminal pore, such as has been described

by Gosse (1860) in P. hastata.

The column (fig. 1) is almost cylindrical, though usually

contracting somewhat towards either extremity and, in some

examples, is grooved by twelve well-marked invections, although

in others these are quite indistinct. Examined under a lens the

surface is seen to be studded with minute elevations, which are

all the more noticeable by being of a paler colour than the

general surface. No foreign particles were adhering to these

elevations in any of the examples studied, and, as I had no

opportunity for examining living specimens, I cannot say whether

they had the power of adhering to surfaces with which they

came into contact, as seems to be the case with the similar

* Wor explanation of the Plate see p. 972.

964 PROF. J. P. McMURRICH ON

structures of P. hastata (Haddon and Dixon, 1885) and P. tri-

capitata (Andres, 1883).

There was no differentiation of the upper part of the column to

form a capitulum, nor was there any distinct margin.

The tentacles are twelve in number, arranged in a single cycle.

They are of moderate length and stoutness, obtuse or tapering

slightly at the tips, but not capitate, and frequently with longi-

tudinal grooves.

The lips are elevated considerably above the level of the

disk, and show only a single siphonoglyph (fig. 2), which is

completely separated from the rest of the stomatodeeum by the

fusion of its lips throughout the greater part of their extent.

At least this was the case in three individuals that I examined

anatomically, and it would seem, therefore, to be of general

occurrence. Sedgwick (1884) has described the same peculiarity

as occasional in P. hastata, but it is not mentioned by other

authors who have studied the anatomy of that species (Haddon,

1885, 1889; Faurot, 1890, 1895).

The general surface of the stomatodeum is longitudinally

grooved along the lines corresponding to the insertions of the

mesenteries, and each lip is raised in the intervals between the

grooves into five rounded elevations, which correspond with the

intervals between the perfect mesenteries, with the exceptions of

those between the directives. One of these latter is quite small

and the other is represented by the margins of the siphonoglyph,

which, although not elevated to any great extent beyond the rest

of the lips, nevertheless bear fine tuberculiform or digitiform

processes (figs. 2, 3) which correspond with the conchular lobes

of other forms. Two of these processes form a pair lying one

on either side of the more dorsal portion of the siphonoglyph

opening, and the three others are situated more ventrally, the

median one being in the sagittal plane of the body, opposite

the bottom of the siphonoglyph. In three individuals examined,

of different sizes, I found no variation in the form or arrange-

ment of these conchular processes, and would therefore conclude

that this is their final adult arrangement.

Size.—The height of the column, measured in individuals pre-

served in formalin, was in the larger forms about 2°0 em., the

diameter at about the middle being about 0-9 cm., and at the

limbus 0°6 cm. The length of the tentacles was 0-4 cm.

Colour.—Examples preserved in formalin retain a considerable

amount of the original coloration, and from these it is possible to

state that the general colour of the column is a more or less

intense brownish red. In some cases it is distributed over the

entire extent of the column and in others it is limited to the

distal portion, the proximal or basal part being almost colourless.

A closer examination shows that the pigment is not uniformly

distributed throughout the area in which it occurs, but presents

the appearance of a brownish red ground-colour upon which

are scattered numerous minute dots of a paler shade, these

NEW ACTINIANS FROM BRITISH COLUMTIIA. 965

representing the small elevations already described as scattered

over the entire surface of the column. The tentacles seem to have

been of a paler colour than the upper part of the column and

marked by four or five rings of a more or less intense brownish

red (fig. 2). The conchular processes seem to have been of the

deeper shade, with paler apices.

Structure.—The column mesoglea is much thicker than the

ectoderm and is of a finely fibrillar structure, cells being scattered

among the fibrils. There are no ectodermal muscles, but

associated with the presence of ectodermal gland-cells there is a

well-defined layer of nerve-fibres. The adhesive organs appear

in sections as elevated areas of the ectoderm, composed of slender

cells considerably longer than those of the adjacent ectoderm ;

the areas contain no gland-cells and, consequently, appear much

paler than the general ectoderm in stained preparations. In

structure, therefore, the organs differ from typical verruce only

in being elevations of the surface instead of depressions. The

endodermal musculature is supported on short simple processes,

and towards the distal portion of the column it becomes weaker,

the cells being arranged in an almost simple layer, and there is

no indication of a sphincter.

The musculature of the tentacles and disk is very moderately

developed, and there are no muscle-fibres on either the ectodermal

or endodermal surface of the stomatodeum. The structure of

the siphonoglyph is quite characteristic. Its ectoderm is much

higher than that of the rest of the stomatodeum and destitute of

gland-cells. The surface bears very numerous strong cilia and is

provided with a distinct cuticular layer, beneath which the nuclei

of the cells are arranged in very many layers, the basal portion of

the epithelium containing but few, and appearing as 1f composed

of a very fine reticulum, which is traversed by a band of nerve-

fibres. The endoderm is also much higher than it is over the

rest of the stomatodeeum and its basal portion is also reticular in

structure, though somewhat coarser than the ectoderm. The

mesogloea also differs from that of the general stomatodzeum, its

ground-substance being almost homogeneous and assuming a

clear blue tint with hematoxylin-erythrosin, while elsewhere the

colour is decidedly violet. The fusion of the lips, mentioned

above, is merely an epithelial union; indeed, it appears to be

due to adhesion of the cuticular layers of the two adjacent

surfaces.

The mesenteries are arranged in ten pairs, six of which

are perfect, the remaining four pairs occupying the lateral

and sulco-lateral interspaces. Of the perfect mesenteries, two

pairs are directives, and all have well-developed diffuse muscle-

pennons (fig. 4) consisting of strong though but slightly

branched lamellae. A well-marked parieto-basilar muscle is also

present forming usually a slight fold. The reproductive cells

could not be distinguished in the individuals examined, but

the endoderm of the mesenteries in the region proximal to the

966 PROF. J. P. McMURRICH ON

mesenterial filament is greatly thickened and very granular, as

if from the presence of ingested food-material. ‘The eight im-

perfect mesenteries are destitute of mesenterial filaments and,

indeed, represent only the muscular portion of the perfect

mesenteries. Their muscle-processes ave but few in number and

they do not possess any distinct pavieto-basilar muscle. There

is no difference in the development of the various perfect

mesenteries such as Faurot has described for P. hastata; all are

provided with mesenterial filaments and are alike in all

particulars.

The first description of a form that may definitely be assigned

to the genus Peachia is that furnished by Reid (1848) of a species

washed ashore in the Bay of St. Andrews and named Actinia

cylindrica. ‘This name had, however, already been employed by

Renier (1804) for Certanthus membranaceus, and has therefore

given place to the term Peachia hastata, proposed by Gosse (1855)

for a form with which Reid’s species is evidently identical. In

the following year Koren and Danielssen described under the

name Siphonactinia boeckit a form that is certainly congeneric

with the Peachia of Gosse, and that author in 1860 described two

additional members of the genus, P. uwndata and P. triphylla.

No further additions to the list of species were made until 1879,

when Hutton described as P. carnea a form cast up on the beach

at Dunedin, New Zealand; a little later Andies (1883) described

as Siphonactinia tricapitata a form from Naples that he had

originally (1881) considered identical with Gosse’s P. triphylla,

and in 1893 I described P. korent from off the coast of the

Argentine Republic. There seems to be no room for doubt as to

the distinctness of P. guinqguecapitata from P. hastata: the form

of the conchula, the equality of all the perfect mesenteries, and

the feebler development of the longitudinal musculature of the

imperfect ones furnish sufficient bases for their separation, in-

dependently of their coloration and geographical distribution.

P. koreni, with only eight tentacles and a simple conchula, may

also be regarded as distinct, and although the description of

P. carnea is very incomplete, its geographical distribution is

prima facie evidence of its distinctness also.

With regard to the remaining four species, all of which are

Kuropean, the evidence is not so clear, sce no anatomical data

concerning them are available. P. wndata, from the Channel

Islands, is believed by Haddon (1889) and G. Y. & A, F. Dixon

(1891) to be merely a young example of P. hastata, its conchula

resembling that of immature examples of the latter species.

P. triphylla, also from the Channel Islands and the Firth of

Clyde (Robertson, 1869, 1875), on the other hand, has a conchula

composed of three lamellar processes, and in this resembles

P. boeckii; the latter, however, bearing the lamellae upon the

summit of a tubular prolongation of the siphonoglyph, whereas

in Gosse’s figure of P. triphylla they are represented as sessile.

The difenencs may, however, be due toa difference in contraction,

NEW ACTINIANS FROM BRITISH COLUMBIA. 967

and it is not improbable that the two forms are identical. It

must be noted, however, that Faurot (1895) identifies P. boeckir

with P. hastata, and until anatomical data are furnished it will

be well to regard it as a distinet form. Finally, in the Medi-

terranean P, iricapitata the conchula is composed of tubercles ;

and while the correctness of its original identification by Andres

(1881) with P. triphylla must still be regarded as a possibility,

it seems advisable for the present to regard it as a good

species.

A provisional arrangement of the known species of Peachia

may then be stated thus :—

Tentacles 12,

Conchular lobes 6-20, irregular ...............0..0........ PB. hastata.

Couchular lobes 3, lamellar,

sessile ......... 06.04 P. triphylla.

borne on tubular prolongation of siphonoglyph. P. boeckii.

Conchular lobes tuberculiform,

three in number sodoosdeaoabdsBeosocsonensce | Leo HPUCRY DIAM RES

five im mumaber 2...) sccseseeesececeenseicre esses ee Quinguecapitata.

? Poets dsb Aun ote Soetoqupenidn cool Me ra AOU TORO

MentaclestSii Mi tasts sa yserche copies nba weuietse nian stb ere MONEIUTs

BIcIDIUM EZQUOREA, sp.n. (Pl. XCVIII. figs. 5-7.)

This form is not uncommon as a parasite upon the bell of the

Leptomedusa “quorea forekalii, which is of very common occur-

rence in the waters of British Columbia. The column (fig. 5)

has a rounded base, is somewhat conical or urn-shaped and

wrinkled transversely by contraction in examples preserved in

formalin. Proximally it usually shows some faint longitudinal

grooves which mark the lines of insertion of mesenteries, but no

signs of verruce or other adhesive organs could be seen. ‘The

margin, which is indistinct, is occupied by a single series of twelve

short and obtuse tentacles. The mouth (fig. 6) is usually

widely expanded, so as to completely hide the disk, and shows a

single siphonoglyph ; no conchula was present.

Size.— Length about 7 mm., greatest diameter 4 mm.

Colowr.—There was no trace of colour in any of the examples

seen.

Structure—T'be column mesoglea is thinner than either the

ectoderm or endoderm and has a homogeneous or finely fibrillar

structure, cells being scattered throughout the ground-substance.

The ectoderm is of uniform structure throughout its entire

extent, no verrucal areas being distinguishable. There are no

ectodermal muscle-fibres, and those of the endodermal surface are

arranged ina simple layer, there being no mesogleeal lamellee for

their support and no indications of a sphincter muscle. The

musculature of the tentacles is also very feeble, the ectodermal

fibres being arranged in a single uviform layer, while the endo-

dermal ones are hardly distinguishable.

The single siphonoglyph is prolonged somewhat below the level

968 PROF, J. P. McMURRICH ON

of the rest of the stomatodeum and forms a well-marked deep

groove, whose endoderm is much higher than that of the rest

of the stomatodeum, although it does not present a reticular

formation such as occurs in Peachia quinquecapitata. ‘There are

six pairs of mesenteries, all of which are perfect and all possess

mesenterial filaments; no indications of additional mesenteries

of the second cycle were to be found. ‘Two of the six pairs

were directives. ‘The longitudinal musculature (fig. 7) formed

a low pennon extending throughout the entire muscular area

of the mesentery, the supporting lamelle being palisade-like

in their arrangement, and is higher than the endoderm that

covered them. No parieto-basilar muscle was present in my

preparations, which did not, however, include the most proximal

portion of the column.

A number of immature Actinians, a list of which has been

given by Haddon (1887), have been described as parasites in

various species of Medusz, and all present structural features

sufficiently similar to suggest that they are generically identical

with one another and with the form described above. They have

been assigned in part to the genus Halcampa and in part to

Peachia, but none of these has yet been actually shown to trans-

form into the one or the other of these genera, and until the

adult condition is definitely known it seems well to retain

for them the genus Dicidium established by L. Agassiz (1859),

recognizing, however, that this is but provisional and that the

forms assigned to it are larval or at least Immature.

The evidence furnished by the structure, although not con-

clusive, does however give some indications of the probable

position of the adult form and, I believe, points towards the

genus Peachia, rather than to Halcampa. Differences between

the two genera that are pertinent to this question are to be found

in the presence of a conchula in Peachia and its absence in

Halcampa; in the single deep siphonoglyph of Peachia as

compared with the two shallow ones of Hulcampa; and in the

somewhat diffuse form of the muscle pennons in Peachia as

compared with the compact and circumscribed ones of Halcampa.

The first of these differences is not apparent in all species of

Bicidium, the conchula probably being late in its development,

but the other characteristics are constant in all known forms

whose anatomy has been recorded.

Since Haddon (1887) reviewed the various species of Bicidium

some additions have been made to our knowledge of them, and in

support of my contention as to their affinities to Peachia rather

than to Haleampa, a reconsideration of them may not be amiss.

The absence of a conchula in B. equoree is, it is true, opposed to

its reference to Peachia, but, as has been pointed out, this

structure may very well be late in developing, and in the nature

of the siphonoglyph and the muscle pennons the similarity to

that form and the difference from Halecampa are pronounced.

NEW ACTINIANS FROM BRITISH COLUMBIA, 969

The occurrence of P. quinquecapitata in the same locality is also

suggestive, but the differences between the two forms are too

marked to warrant their identification, although it is not im-

probable that these differences are due to age rather than specific

differences.

The structure of Actinia clavus of Quoy and Gaimard (1833),

recently studied by Pax (1912), shows, I believe, that it must be

considered congeneric with B. equoree, although Pax refers it to

the genus Halcampa. As in the British Columbian species, there

are no indications of a conchula; but again, the form of the

siphonoglyph and the muscle pennons suggest Peachia rather than

Halcampa, and until further evidence as to its exact position

is available, it would seem well that it should be known as

Bicidium clavus.*

The evidence as to the affinities of B. parasiticwm is more

definite. It was first described by L. Agassiz (1859) from Cyanea

arctica and later by Verrill (1864), who showed that it possessed

a well-marked trilobed conchula, and, still later (1866), assigned

it to the genus Peachia. Preparations that I have of this species

show it to have a striking general similarity to 5. equoree, the

mesenteries being twelve in number, all perfect and all bearing

mesenterial filaments, and the siphonoglyph single and deep.

The muscle pennons also are of the same diffuse type, but the

lamelle (fig. 8) are not arranged in the manner of a palisade,

but are decidedly branched in a dendritic manner so that

they present an appearance of being airanged in groups. It

is to be noted that Verrill (1874) mentions the capture of two

very large examples of this species, imbedded in gravel at low-

water mark at Eastport, Maine; he does not, however, give any

anatomical data concerning them, and until it is definitely known

that the form develops the four pairs of secondary mesenteries

. (zygocnemes), it seems-advisable to allow it to remain in the

genus Sicidium.

The form which F. Miller (1860) deseribed very completely as

Philomedusa vogtii is also undoubtedly a Bicidium. It occurred

upon the meduse Olindias and Chrysaora and was provided with

a trilobed conchula, twelve tentacles, twelve mesenteries, all of

which were perfect and furnished with mesenterial filaments, and

had asingle siphonoglyph. The form described by Graeffe (1884)

* The observations of Pax make it quite certain that R. Hertwig (1882) was in

error in identifying a Halcampa from the Kerguelen Islands with Quoy and

Gaimard’s species. The possibility of Hertwig’s forms being young examples of

Studer’s Hdwardsia kerguelensis, which Kwietniewski (1896) has shown to hea

Halcampid, should not be lost sight of, although the differences in the descriptions

as they stand are too great to permit of a definite identification. Furthermore, the

contention of Pax that the H. clawus of Tizard and Murray (1881) and Appelléf

(1897) is distinct from that described by Hertwig is undoubtedly correct, but since

the forms so named by Appelléf were the actual types of Danielssen’s Halcampoides

ahyssorwm, it is difficult to understand the necessity for the new name, seplentri-

onalis, that Pax bestows upon them.

970 PROF. J. P. McMURRICH ON

as Haleampa medusophila, occurring on various meduse (Zina,

Octorchis, and Mquorea) is probably identical with Miller’s

species, the absence of a conchula being probably due to its

greater immaturity.

In 1860 Wright described as Peachia fultoni * a form that

he found parasitic on a species of Thawmantias (Phialidium %),

later (1861) changing its name to Haleampa fultoni. This 1s

probably the same as the form described by Haddon (1887), and

erroneously, as has been pointed out by Carlgren (1904), regarded

by that author as the larva of Haleampa chrysanthellum. In

1887, McIntosh recorded the occurrence of actinian larvee on

various Thaumantiad meduse occurring at St. Andrews, identi-

fying them as the young of P. hastata, an opinion in which

Haddon (1888) concurred after an examination of their structure,

still maintaining, however, their distinctness from the examples

previously described as the young of HH. chrysanthellum. They

possessed eight tentacles and eight large mesenteries together

with four smaller deuterocnemic ones. It may be that Haddon

is correct in regarding the two forms he examined as distinct

species, but even if so it seems clear that they are to be referred

to the same genus, and since it is still uncertain that they

actually do develop into Peachia, it will be preferable to place

them in Biciduin.

Finally, Dendy (1888) has described from medusz obtained

at Port Philip a parasitic actinian with twelve tentacles, within

which there was ‘“‘an inner circle of 12 cushion-like swellings,”

which in older individuals become saccular outgrowths. It is

not possible to ascertain from Dendy’s account, whether or not

these outgrowths represent a conchula, and the exact affinities of

the form must remain doubtful, although with a probability that

it represents a Peachia’.

The available evidence seems, accordingly, to point strongly in

favour of these various medusophilous forms being young stages

in the development of Peachia rather than Halcampa, but a

direct linking up of the immature examples of Sicidiwm with

their respective adults is necessary to settle the question.

* Some confusion exists in the references to the literature of this species. Andres

(1883) gives the date of its first description as 1859 and as references the Pro-

ceedings of the Physical Society of Edinburgh, vol. 11. 1859, and the New Edinburgh

Philosophical Journal, vol. xii. 1860. Haddon (1887) repeats the latter reference

but substitutes for the former, Proceedings of the Royal Society of Edinburgh,

vol. ii. 1860. There is no paper in the Proceedings of the Royal Society of

Edinburgh by Wright referring to this species, and the date of vol. i. of the

Proceedings of the Royal Physical Society of Edinburgh, which does contain such

a paper, is 1861 and not 1859. ‘The reference to the Edinburgh New Plilosophical

Journal (this being its correct title) should read “ New Series, vol. xii. 1860.”

+ Carlgren (1904) mentions, without descriptions or names, two additional forms

that should probably be referred to this genus, one occurring in the medusa

Eutimalphes indicans on the Swedish coast and the other on a large medusa from

Valparaiso.

“NEW ACTINIANS FROM BRITISH COLUMBIA. 971

LITERATURE.

Aaass1z, L.—On some new Actinoid Polyps of the United States.

Proc. Boston Soc. Nat. Hist. vi., 1859.

Anpres, A.—Prodromus neapolitane actiniarum faune ete.

Mitth. Zool. Stat. Neapel, ii., 1881.

Anpres, A.—Die Aktinien. arma und Flora des Golfes von

Neapel, ix., 1883.

APPELLOF, Die Actiniengattunzen Fenja, Algir und Hal-

campoides, Dan. Bergens Mus. Aarb. x1., 1881.

CaRLGREN, O.—Kurze Mittheilungen tiber Anthozoen. Zool,

Anz. xxvii, 1904.

Dantetssen, D. G.—« Actinida.” Norwegian North Atlantic

Exped. xix., 1890.

Denpy, A.—Note on some Actinian larve parasitic upon a

medusa from Port Philip. Proc. Roy. Soc. Victoria,

1888.

Dixon, G. Y. and A. F.—Report on the Marine Invertebrate

Fauna near Dublin. Proc. Roy. Trish Acad., ser. 3, i1.,

1891.

Favrot, L.—Sur la disposition des cloisons mésenteroides chez

la Peachia hastata. C. R. Acad. Paris, ex., 1890.

Favror, L—Etudes sur l’anatomie, l’histologie et la développe-

ment des Actinies. Arch. Zool. exp. et gén., ser. 3, ii1.,

1895.

Gossn, P. H.—Description of Peachia hastata, ete. Trans.

Linnean Soe. xxi., 1855.

Gosss, P. H— Actinologia Britannica. London, 1860.

Graerre, E.—Uebersicht der Seethierfauna des Golfes von Trieste.

ILI. Celenteraten. Arb. zool. Inst. Wien, v., 1884.

Happon, A. C., and Drxon, G. Y.—The structure and habits

of Peachia hastata (Gosse). Sci. Proc. Roy. Dublin Soc.,

n. S. iv., 1885.

Happon, A. C.—On the arrangement of the mesenteries in the

parasitic larva of Halcampa chrysanthellum (Peach).

Proc. Roy. Dublin Soe. n. s. iv., 1887.

Happon, A. C.—On larval Actiniz parasitic in Hydromeduse

at St. Andrews. Ann. Mag. Nat. Hist. ser. 6, 11., 1888.

Happon, A. C.—A Revision of “She eae Netinie. "Part It,

Sci. Trans. Roy. Dublin Soe. ser. 2, iv., 1889.

Hurron, F. W.—Contributions to a (Galenterate Fauna of

New Zealand. Trans. New Zealand Inst. xii., 1879.

Koren, J., and Dantetssen, D.C.—Nye Actinier. Fauna htt.

Norvegie, i1., 1856.

Kwirtniewski, C. R.—Revision der Actinien welche von

ierrnee Profs Sbuder. ayn gesammelt worden. Jena.

Zeitschr. xxx., 1896.

McIntosu, W. C.—On the commensalistic habits of the larval

forms of Peachia. Ann. Mag. Nat. Hist. ser. 5, xx.,

1887.

972 ON NEW ACTINIANS FROM BRITISH COLUMBIA.

McMourricu, J. P.—Report on the Actiniz collected by the

U.S. Fish Commission Steamer ‘ Albatross’ during the

winter of 1887-1888. Proc. U.S. Nat. Mus. xvi., 1893.

Miter, F.—Ueber Philomedusa Vogtii. Arch. f. Naturg. xxvi.,

1860.

Pax, F.—Revision des types des Actinies décrites par Quoy et

Gaimard. Ann. Sci. Nat., Zool. sér. 9, xvi., 1912.

Quoy et GAImMARD.—Zoologie du Voyage de la corvette ‘Astrolabe.’

Paris, 1833.

Ret, J.—Account of a new species of Actinia (A. cylindrica).

Aun. Mag. Nat. Hist. ser. 2, 1., 1848.

Rosertson, D.—Living specimen of a rare Sea Anemone, Peachia

triphylla. Proc. Nat. Hist. Soc. Glasgow, i., 1869.

Rosertson, D.—On the Sea-Anemones of the shores of the

Cumbraes. Proc. Nat. Hist. Soc. Glasgow, i1., 1875.

Sepewrck, A.—On the Origin of Metameric Segmentation ete.

Quart. Journ. Micr. Sci. ser. 2, xxiv., 1884.

Tizarp and Murray.—Exploration of the Faroe Channel during

the summer of 1880 in H.M. hired ship ‘ Knight Errant.’

Proc. Roy. Soc. Edinburgh, x1., 1881.

Verritt, A. E.—Revision of the Polypi of the Eastern Coast of

the United States. Mem. Boston Soc. Nat. Hist. 1.,

1864.

Wraieut, 1’. S.—Ohservations on British Zoophytes. Edinburgh

New Phil. Journ. n. s. xii., 1860.

Wricut, T. 8.— Observations on British Zoophytes. On

Halcampa Fultoni, a parasitic Actinia. Proc. Roy. Phys.

Soe. Edinburgh, u1., 1861.

EXPLANATION OF PLATE XCVIII.

r. 1. Peachia quinquecapitata. Nat. size.

2. Oral view of P. quinquecapitata expanded. X 2?

3. Oral view of partially contracted individual of P. quinquecapitata,

showing arrangement of conchular lobes.

4, Transverse sections of mesenteries of P. guinquecapitata.

5. Side view of individual of Bicidiwm equoree. X A.

6. Oral surface of B. equoree. X 4.

7. Transverse sections of mesenteries of B. equoree.

8. Transverse sections of mesenteries of B. parasiticum.

ON SPONGES IN WATERWORKS. 973

57. Sponges in Waterworks. By W. N. Parner, Ph.D.,

F.Z.S., Professor of Zoology, University College,

Cardiff,

[ Received September 17, 1913 ; Read November 11, 1913.]

INDEX. Pages

Occurrence in Cardiff Waterworks and elsewhere ... 973, 974

TBRHL IYO) GSA dion aoa las Buados abe Reb donors nosoad aad pao uaaEos doaaaens oom KIC Cee)

Reproduction Neato ano PRA nod cass ete rosacachEcceuannseemey ei ONO

Methods of treatment adopted .....................255 ae 976

Some three or four years ago I received through my colleague,

Dr. Scholberg, some specimens of a branched, yellowish-brown

organism which had been found growing in some of the pipes

at the Cardiff Waterworks, and which proved to be very fine

examples of Spongilla lacustris. Subsequently I learnt that

‘there had been an extensive growth of this sponge in the pipes

leading to one of the series of filter-beds, which had caused much

trouble and difficulty; and that, although the growth had

apparently disappeared for a time after treatment with sulphate

of copper, it, as | expected, soon appeared again, and became as

flourishing as ever. In the summer of 1911 the growth was so

extensive as to cause anxiety, not only as regards reducing the

flow in the pipes, but also on account of the unpleasant odour

which had been noticed in the water, due doubtless to the

living sponge as well as to its decomposition. In December

of the same year I was requested by the Waterworks Engineer,

Committee; and now that the treatment I subsequently pro-

posed has—at any rate for the present—been successful, I have

the permission of the Committee to publish an account of our

experiences. [I should lke to express my indebtedness to

Mr. Priestley and his staff for their cooperation and for giving

me every facility in prosecuting the enquiry; I have also to

thank Dr. Kemna, of Antwerp, Dr. Harmer, Mr. Kirkpatrick,

and Professor Minchin for valuable information given at the

beginning of the investigation.

_ Although sponges of various species have been recorded at

numerous other Waterworks *, they do not appear to have been

the chief source of trouble, as at Cardiff.

The interesting subject of the fauna of Waterworks has quite

recently been so fully dealt with by Dr. Harmer in these

“ Proceedings” (19138, p. 426), that it is unnecessary for me

* Hg. at Boston, Mass. (Bowerbank. Proc: Zool. Soc. 1863, & Potts, Acad. Nat.

Sci. Philadelphia, 1881, 1882, and 1887); Hamburg (Kraepelin, Abh. Naturwiss.

Ver. Hamburg, 1886); Torquay (Chapman, “ Animal Growths in Water-pipes,”

Transactions of the Institute of Water Hngineers, 1913, & Kirkpatrick—see Harmer,

P. Z. 8. 1913, p. 486) ; and Aberdeen (Kirkpatrick, loc. cit. p. 489).

Proc. Zoou. Soc.— 1913, No. LXV. 65

O74 PROF. W. N. PARKER ON

to refer in detail to the history and literature of the subject.

Dr. Harmer’s account of the difficulties which have recently

occurred in various English Waterworks owing to the presence

of Polyzoa, shows the importance of biological investigation in

dealing with such cases, as was pointed out in this country by

Professor Hickson in ms presidential address to Section D of the

British Association at Southport in 1903.

The Cardiff district is supplied with excellent soft water from

the Taff Fawr reservoirs in Brecknockshire, midway between

Merthyr and Brecon. From these higher reservoirs the water

is conducted by conduits into various other storage-reservoir's

at a lower level, in the near neighbourhood of Cardiff, and

from those into filter-beds: the latter, of course, prevent the

access of organic particles and spicules into the service- pipes

supplied from them. The sponge had hitherto only been found in

the pipes leading to certain of the filters on the northern border

of Cardiff known as the “‘ Heath” filters, from which the greater

part of the city is supplied. The water to these filters comes

from two storage-reservoirs, at Llanishen and Lisvane, situated

about a mile farther north. Careful examination has so far not

revealed any trace of the sponge in other parts of the system,

which therefore I need not describe further here*.

The water from Llanishen reservoir passes through metal

screens with fine meshes, so as to strain off all but minute solid

particles, into a valve-shaft from which it is conducted by under-

ground pipes to a valve-chamber at the “ Heath.” It is then

again screened before flowing into a 3-ft. pipe, with which lateral

pipes are connected ending in bell-mouths, one to each of the six

filter-beds.

The chief trouble occurred in this pipe, which was thickty lined

with a luxurious growth of the sponge, consisting of dense

incrustations giving off numerous finger-like processes which

branched and branched again, and some of which reached a

length of 8 inches or more, frequently showing concrescence 7.

Other growths were found in the valve-chamber and _ screen-

chamber, in the corners and crevices of which it was not easy to

get access to them.

On visiting the “ Heath” on December 12th, 1911, the sponge,

packed with gemmules, showed no signs of dying down for the

winter, and it appeared to me that had it been desired to culti-

vate the organism, probably no better plan could have been

devised than that furnished by the water-chambers and pipes

* For further details with regard to the Cardiff Waterworks, see a paper by

C. H. Priestley, M.Inst.C.K., on the “ Development of the Cardiff Water Supply,”

read at the Sessional meeting of the Royal Sanitary Institute at Cardiff on April

12th, 1912; also a description of the Waterworks by the same author issued from the

Cardiff Waterworks Engineer’s Otfice, 1908.

+ Cf. figures by Bowerbank, “A Monograph of the British Spongiadz,” Ray

Society, 1864-1882; and Johnston, “A History of British Sponges and Lithophytes

(Corallines),” London, Edinburgh, and Dublin, 1882.

SPONGES IN WATERWORKS. LD)

(cf. Kraepelin, loc. cit.). It can grow as well in the dark, with-

out zoochlorelle, as in the light ; and being effectively protected

from frost and other adverse conditions, it seemed possible that

jt might continue alive during the whole winter. On the same

day, we made a cursory examination at Llanishen and Lisvane,

with no result; but on the following day, the reservoir-keeper,

after emptying the valve-shaft at Llanishen, found a considerable

growth at a depth of between 10 and 20 feet.

Since then, a careful look-out has been kept for the sponge

throughout the entire system, but there has been no positive

evidence of its occurrence in other parts, either in the open or in

the underground pipes.

The two most important problems which presented themselves

were—the destruction of the sponge in the infected area, and the

prevention of re-infection throughout the whole system. As

regards the latter, the only sound method would be the intro-

duction of pre-filtration through sand, so as to starve the sponge

by preventing the microscopic organisms (on which it and other

‘‘microphagous”” organisms which might at any time appear

depend for their nutriment) from passing into the pipes and

reservoirs (cf. Kemna, quoted by Harmer on p. 432), as well as

to keep out larve and gemmules. This method would take

a long time to carry out and entail considerable expense ;

moreover, 1t would not be effective unless it could be adopted at

the lower reservoirs (where there are difficulties as regards level)

as well as the higher ones, as there would be a reappearance of

organisms in them. As the matter was urgent, I therefore felt

it would be better to confine attention at first to the destruction

of the organism in the infected area and the prevention of its

regeneration and of reinfection by the gemmules.

Treatment with any destroying agent except such as would be

dangerous to human life would be ineffective if applied to the

adult sponge, as it would not destroy the myriads of gemmules

protected by their resistant coats. It was therefore necessary in

the first instance to make a periodical examination of the sponge

for a year or more, so as to ascertain (1) whether sexual repro-

duction occurred, (2) when the gemmules germinated, and

(8) whether the adult growth perished partially or entirely

during the course of the winter*.

An examination of specimens at various times of the year has

so far revealed no larve or any signs of sexual reproduction ; but

the enormous production of gemmules throughout the crusts and

branches is quite sufficient to start new growths in any part to

which the gemmules could gain access. There can be no doubt

that a sexual stage is unnecessary for the reproduction of the

sponge from year to year.

* Cf. Marshall (SB. Naturf. Gesellsch., Leipzig, 1884, & Journ. Roy. Micros.

Soc. v. 1885) ; Weltner (Archiv f. Entwicklungsmechanik, Bd. xxxiii. Heft 3 u. 4).

6d*

976 ON SPONGES IN WATERWORKS.

On visiting the “ Heath” and Llanishen on February 3rd, 1912,

when the filter-beds were covered with ice 13 ins. thick, we "found

that the adult growth was dying down, but that it was still

closely packed with gemmules, some of which had begun to

germinate and start new growths among the old. On March 23rd

gemmation had proceeded further, new growths being abundant,

and I suggested that in the course of the spring the pipes and

chambers should be scraped and treated with strong brine, so

applied as to reach any small spaces and crevices in which

gemmules could lodge, and thus to attack the sponge when it was

least capable of resistance. This was done on May 2nd, and

with the sponge were removed a large quantity of iron corrosive

nodules—another trial to the Water Engineer, referred to by

Harmer on pp. 430, 437, & 438 of his paper; the brine was then

washed out through a bye-pass. The result was apparently satis-

factory, butin the course of the summer a few small new growths

were observed by the Foreman: they were very much less

abundant than in the previous summer, and no smell had been

noticed since the treatment. An examination on September 24th

showed that these comparatively few young incrusting sponges

had not yet given rise to any outgrowths, and that they con-

tained no gemmules: they were in all probability due to a few

gemmules which had not germinated when the treatment was

carried out. I therefore suggested a second application of brine.

By November 14th, when this was done, the growths had

increased in size considerably, but apparently no gemmules had

been formed. Instead of leaving the pipes empty for only a few

hours, as on the previous occasion, it was possible this time to

give them three days before being again used, and since then no

trace of the sponge has been seen,

It is interesting to note that no Polyzoa were found, but that

a number of colourless Hydree like those described by Kraepelin

at Hamburg were obtained by the reservoir-keeper from the

bottom of the valve-shaft at Llanishen on May 25th; unfor-

tunately these were all dead by the time they reached me*.

* Since the above was written, a few young sponge-growths were again observed

and at once destroyed. Colonies of Cristatella, with numerous statoblasts, had

also appeared : fortunately this is not one of the “moss’’-forming Polyzoa. (Oct.

25th, 1913.)

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ON CYPRINODONT FISHES. 977

58. A Revision of the Cyprinodont Fishes of the Subfamily

Peeciliine. By O. Tate Ree@an, M.A., F.Z.S.

[Received October 11, 1913 ; Read November 11, 1913. ]

(Plates XCIX,-CI.* and Text-figures 168-173.)

InDEX. :

SYSTEMATIC ;— Page

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The Peeciliine are viviparous Cyprinodonts in which the anal

fin of the male is advanced and modified into an intromittent

organ, They are American, ranging from Carolina to Argentina

and from Arizona to Ecuador ; many of the species live in the

sea as well as in fresh water. Most of the species are quite

small, two or three inches long; the giant of the group, Belonesox

belizanus, attains a length of eight inches, In some forms the

males are much smaller than the females, and the adult male of

Heterandria formosa, 15 to 20 mm, long, is one of the smallest

fishes known; in several species males are rather scarce,

In Giinther’s classification (Cat. Fish. vol. vi.) the Cyprinodonts

were divided into Carnivore, with the intestine short or but

little convoluted and the bones of the lower jaw firmly united,

and Limnophage, with the intestine long and convoluted and

the bones of the lower jaw loosely connected. It is now known

that both types oceur in two distinct subfamilies, Characodontinza

and Peeciliine, and there is good evidence that in the latter the

* For explanation of the Plates see p, 1018,

978 MR. C. TATE REGAN ON

limnophagous structure has been evolved three or four times

independently.

The intromittent organ is freely movable and is supported

internally by bony stays, 2 to 5 in number, that project down-

wards and forwards from the bridges that connect the parapo-

physes of the posterior precaudal vertebre. It is formed by the

prolongation of the third, fourth, and fifth anal rays, the first

and last of which are stout and are more or less expanded trans-'

versely so as to margin a groove on one side or the other, or they

may even meet so as to form a tube. The urogenital orifice is

directly in front of the base of the fin, and may often be covered

by the pelvic fins, which, doubtless, help to conduct the seminal

fluid into the groove, or in certain forms where the pelvic fins of

the male are considerably enlarged they may be closely applied

to the intromittent organ, converting the grooves into closed

tubes. The distal segments of the prolonged rays may be

variously modified into spines, hooks, barbs, ete., which may help

to retain the organ in position during coition.

The differences in structure of the intromittent organ are of

great systematic importance; hitherto they have been almost

neglected, except by Higenmann (Proc. U.S. Nat. Mus. xxxil.

1907, p. 425), who has described them in the genera Lebistes,

Girardinus, Glaridichthys, Phalloceros, and Phalloptychus *.

Many of the species eat Mosquito larve, and the ‘“ Millions ”

fish (Lebistes reticulatus) has been introduced into many countries

in the hope that it will check malarial fever t. On account of

their small size, pretty appearance, and interesting habits the

Peeciliine are great favourites with European aquarium-lovers,

especially in Germany; several of the new species described

below have been sent to me by my valued correspondents in

Hamburg, Herr J. P. Arnold and Herr A. Rachow, as newly

imported aquarium-fishes. In the German aquarium-journals

and in two booklets by Stansch (Die lebendgebiirende Zahnfarkfen,

Leipzig, Wenzel, 1910, 1911) have appeared many accounts of the

habits of different species: from these one gathers that the

broods follow each other at intervals of only a few weeks, but

that the number of young varies greatly according to the species ;

thus a pair of Pecilia pare had broods of 8, 11, and 10 at

intervals of four weeks, but in P. vivipara 124 were counted in a

single brood. ‘The new-born young are usually from 5 to 10 mm.

long and grow rapidly, in many species attaining sexual maturity

inabout three months. The breeding-habits are of great interest :

in species where the males are brilliantly ornamented (J/ol-

lienisia, Xiphophorus, Pecilia brannert) they dart about displaying

their beauty ; in these and in related forms the females appear

to encourage their advances, but in others they are quite shy

and their mates have to exercise cunning to get near them. It

* Some additional figures are given by Langer in a paper on the morphology of

these fishes, just published (Morph. Jahrb. xlvii.).

+ See P.Z.S. 1910, p. 146; 1912, p. 906,

CYPRINODONT FISHES. 979

is of some interest to note that in Pecilia and related genera

the females welcome the attentions of the males and the intro-

mittent organ is quite short, but as in Gambusia and Phalloceros

the females try to make the males keep their distance it is

much longer.

Synopsis of the Genera *.

I. Lower edge of caudal peduncle sharp, without a

median series of scales; bones of lower jaw firmly

united ; teeth conical or villitorm.

Body rather deep ; pelvic fins well developed, behind base

of pectorals; anal tin of male without long appen-

COlEKe =I} Mea nan Gun ttn Sagon some ud Sema TOES Can Coe eee cae

Body elongate; pelvic fins absent in female, minute and

below gill-openings in male; anal fin of male ending

in a pair of long appendages with spine-like processes 2. Temeurus.

1. Alfare.

II. Lower edge of caudal peduncle rounded or obtuse,

with median series of scales.

A. Pelvic fins similar in both sexes.

1. First produced ray of anal fin of male without

long processes or appendages.

a. Third and posterior branch of second pro-

duced ray of anal fin of male each ending

in a retrorse spine or hook.

Jsws not produced; teeth conical or villiform; anal fin

of male with distal part of first produced ray serrated,

the terminal serrations directed towards its tip;

third produced ray and posterior branch of second

each ending in a retrorse hook. Dorsal 6-12; origin

belindithatiotanalieeeeneee case reece ee eres Bonn Guaaebustian

Jaws produced ; teeth slender, pointed, depressible; anal

fin of male with terminal segments of first produced

ray not forming serrations; third produced ray and

posterior branch of second each ending in a retrorse

SPUN CBee eer See ese aac COREE ep ae eloweson:

6. Third and posterior branch of second pro-

duced rays of anal fin of male not bearing

retrorse hooks or spines,

a, Anal fin of male ending in a more or less

distinct antrorse hook, the anterior branch

of the second produced ray curved for-

ward.

* Mouth moderate, with distinct lateral

cleft ; teeth conical or yilliform,

+ Dorsal 7-12; origin behind that of anal.

Extremity of anal fin of male supported equally by third

and anterior branch of second produced rays ......... 5. Priapichthys,

Extremity of anal fin of male supported equally by first

and anterior branch of second produced rays ......... 6, Priapella.

+t Dorsal 11-17; origin in advance of

that of anal; extremity of male anal

fin a strong hook formed by the un-

segmented end of the anterior branch

of the second produced ray ............... 7, Pseudoxiphophoyus,

** Mouth small, without distinct lateral

cleft; hook at end of anal fin of male

formed by anterior branch of second

produced ray.

* The differences in the structure of the male intromittent organ used in this

synopsis are illustrated in text-tigs. 168, 169 (genera 1 to 4 and 11), 170 (genera 5 to

10), 171 (genera 10 and 13), 172 (genera 14 to 22), and 178 (genera 23 to 26).

”

980 MR. C. TATE REGAN ON

Teeth somewhat compressed, pointed; anal fin of male

with first produced ray not serrated, third longer

than posterior branch of second .

Teeth broad incisors; anal fin of male with first pro-

duced ray serrated not far from tip, third shorter

than posterior branch of second .. bere ee

*** Mouth small, fanevere teeth TnI

oar-shaped. Extremity of anal fin of

male supported equally by first and

anterior branch of second produced

rays; first not serrated ..................055

B. Anal fin of male short, ending in a small

retrorse hook formed by second and

third produced rays; teeth conical or

villiform

y. Anal fin of male Tone erg ‘dlandie, me

ending in an antrorse hook.

Bones of lower jaw firmly united; teeth conical or villi-

form, fixed .. :

Bones of lower jaw ‘loosely connected ; ‘teeth more or less

ease and compressed, mov able :

. First produced ray of anal fin of male maine

in an antrorse appendage; mouth small, but

bones of lower jaw rather firmly joined.

Appendage forked, each fork antler-like; teeth oar-

shaped .. a de rsedcr sae edeateri aaah eernbbicia

Appendage not forked, ‘long, doubly curved, belnteds

teeth chisel-shaped abit

3. First produced ray of exell fin a et ean

a pair of curved horn-like processes not far

from its end.

a. Bones of lower jaw firmly united.

Teeth chisel-shaped ; outer series close-set

Teeth spear-shaped ; outer series spaced UNS AS

6. Bones of lower jaw ey connected ; teeth

ROWAN, sccnonadocvne

B. Pelvic fins enlarged in the “amallas ‘dhe ceuol | ray

longest.

1. Bones of lower jaw firmly united; outer series

of teeth slender, pointed ..................... 000005

2. Bones of lower jaw rather firmly united; outer

series of teeth broad incisors .............0.00.0..

3. Bones of lower jaw loosely connected; outer

series of teeth slender, curved, oar- or spoon-

shaped ; intromittent organ short.

a. Extremity of intromittent organ unpro-

tected; first produced ray with terminal

hook and strong subterminal serrations;

anterior branch of second hooked forward ;

posterior branch of second with distal seg-

ments ‘produced into serrations beyond

extremity of third.

Third produced ray without hook; anterior branch of

second unsegmented distally ; caudal similar in both

SEXES) PL OFSAl OIA Re ate re mmeNins Ree eP A a) a epee Ser

Third produced ray ending in a retrorse hook ; anterior

branch of second segmented throughout; caudal

of male with lower rays a into a long

ee appendage. Dorsal 11-15.. ee F

. Extremity of intromittent organ Spretened Ty

a hood of thick skin that can be slipped off

frontwards, being attached only at its base

to the first prolonged ray.

a, First prolonged ray tapering evenly, sharply

serrated distally.

8. Heterandria.,

9. Pseudopecilia.

10. Peciliopsis.

. 11. Brachyrhaphis.

12. Leptorhaphis.

. 13. Phalloptychus.

. 14. Phalloceros.

. 15. Cnesterodon.

... 16. Glaridichthys.

17. Toxus.

. 18. Girardinus.

19. Pamphorichthys.

20. Pamphoria.

21. Platypecilus.

22, Xiphophorus.

COYPRINODONT FISHES, 981

Kirst prolonged ray without antrorse spine; last without

LOUIE! FORCES ocd. ado condor sho adecaoocsougoscqdecnd sea doaneeeos ap JECHEC CG?

Kirst prolonged ray without antrorse spine; end of last

bearing a pair of processes directed outwards and

towards)theybasevor the) sm) sesscueee ce sess sn cedasejaccies ses: 24. Lebistes.

First prolonged ray bearing a small antrorse spine at or

near its end; last with processes as in Lebistes ...... 25, Mollienisia.

8. First prolonged ray becoming abruptly

slender at some distance from end, not

sharply serrated, bearing a small antrorse

spine at or near its end; last prolonged

‘ray without terminal processes ............ 26. Limia.

. 1. Atraro Meek, 1912.

Petalosoma (non Lewis) Regan, Ann. Mag. Nat. Hist. (8) i.

1908, p. 462.

Alfaro Meek, Field Mus. Publ. Zool. x., Sept. 1912, p. 72.

Petalurichthys Regan, Ann. Mag. Nat. Hist. (8) x., Nov. 1912,

p. 494.

This genus resembles Gambusia, except for the sharp lower edge

of the tail. The intromittent organ (text-fig. 169 F, p. 990) is

simple in structure, and the pelvic fins are enlarged in the male.

1, ALFARO CULTRATUS.

Petalosoma cultratwum Regan, Ann. Mag. Nat. Hist. (8) ii.

1908, p. 462.

Alfaro acutiventralis Meek, Field Mus. Publ., Zool, x.1912, p.72.

Depth 33 in the length. Dorsal 7-8; when laid back nearly

reaching caudal. Anal 9-10, in advance of dorsal. 32 to 35

scales in a longitudinal series. Total length 45 to 90 mm.

Costa Rica.

2. ALFARO AMAZONUM.

Petalosoma amazonum Regan, Ann. Mag. Nat. Hist. (8) viii.

1911, p. 659, figs.

Depth 32 in the length. Dorsal 8-9, when laid back not

reaching caudal. Anal 10, in advance of dorsal. 33 or 34 scales

in a longitudinal series. Total length 35 to 51 mm.

R. Amazon at Obidos.

2. Tomreurus Eigenm., 1909.

Ann. Carnegie Mus. vi. p. 53.

TOMEURUS GRACILIS.

‘Kigenm. |. c. and Mem. Carnegie Mus. v. 1912, p. 460, pl. Ixv.

fies) 7—8:

“British Guiana.

3. GamBusiA Poey, 1855 *.

Mem. i. p. 382.

Gambusia (part.) Giinth. Cat. Fish, vi. p. 333 (1866); Garman,

* G. picturata Poey (‘Synopsis,’ p. 410, 1868) is a doubtful species of uncertain

position.

982 MR. CG. TATE REGAN ON

Mem. Mus, Comp. Zool. xix. 1895, p. 82; Regan, Biol. Centr.-

Amer., Pisces, p. 93 (1907).

Paragambusia Meek, Publ. Columbian Mus., Zool. v. 1904, p. 133.

This genus, as now restricted, includes species in which the

male intromittent organ is moderately long, nearly 4 of the length

of the fish (without caudal), and is formed on a definite plan

which is subject to considerable modification in the different

species. The distal segments of the first prolonged ray are pro-

duced into processes directed more or less towards its apex ; the

anterior branch of the second prolonged ray is slender distally

and at some distance from the “end has an antrorse projection

which may be termed the “ elbow”; the posterior branch of this

ray ends in a retrorse pointed hook or barb, and the segments

immediately proximal to the elbow of the anterior branch are

produced backwards into serre ; the third prolonged ray ends in

a hook more or less similar to that of the second.

Of the 17 species here recognized I have seen males of 9, the

distal part of the anal fin of each of them is figured. Of Gam-

busia holbrookii (text-fig. 169 A) [ have examined two adult males,

of G. oligosticta (text-fig. 169 B) four, of G. nicaraguensis (text-fig.

168 A) two, of G. wrayi (text-fig. 168 B) four, of G. gracilior

(text-fig. 168 C) three, of G. dominicensis (text-fig. 169 C) two,

of G. nigropunctata (text-fig. 168 F) one, of G. punctuta (text-

fig. 168 D) three, and of G. senilis (text-fig. 168. E) one. These

suffice to prove that the differences shown in the figures are really

specific; the number, form, and size of the modified segments

of the first prolonged ray, the number of segments distal to

he elbow, the form of the hook, and the number and length

of the serre of the second prolonged ray, and the form of the

hook of the third, vary but slightly for each species. The most

noticeable variation is in G. oligosticta, the other examples

differing from the one figured in the addition of a segment

to each hook, so that the anterior hook has a short stem and

the posterior quite a long one.

The following table gives the number of dorsal rays, of scales

in a longitudinal series, and the distribution of the species :—

Dz. be Se | Distribution.

Ws Cho OOUDROOIPIE 250255 580 200406 8 30 | Virginia to Alabama.

Ds (Cis FOOPEIOUIS oc0650 000005 sn6060 a 31-32 | Florida to Texas.

Bb CR GIWES. don ceo s0ecncs00ecs00¢ 6-7 30-31 | Florida to Tampico.

215 (CR EAOUMS Soadoasasosaccn soon 8-9 30 Chihuahua, Mexico.

| 5. G. nicaraguensis............ 7-8 28 |S. Mexico to Nicaragua.

| RE WO ed ccantacadace bag sagees sl iq 32 | Nicaragua.

| 7. G. caudovittata ............ | 8 31 | R. San Juan, Colombia.

Waanclymumctate me (cue emlln aLO 31-33 | Cuba.

| O, @ nigropunctata ......... 9 30 | 9

| 10. G. melanosticta ............ | BNO AVS) gg

| ll. G. puncticulata : 9 | =830 hg

| 12. G. melanopleura............| 11-12 | 30-81 | Jamaica.

IS “Cro GUAGE Tooc'sed soe x60 oon seu deh 8 | 31-33 5

\ 1a. Gy Oligosticta pen. ees ee) SH9 29-31 | eS

WG (Cro GPRACH VOR a5 30408 600500006 8-9 31-33 5¢

16. G. dominicensis ............ 9 | - 28-29 | Haiti.

17. G. caymanensis ............ 9 | 30-31 ) Grand Cayman.

983

CYPRINODONT FISHES,

Text-fig. 168.

Distal part of intromittent organ o

C. G. gracilior

Py 10), Et,

p- 61

334 (1866) ;

Len}

lor

pwc’

Dee

Do eS

Cate

as,

« fo:

hee,

FG te

eres

= Se

lea

Se ae

oS) 2s

a ace

x AO

Sl ee .

Ses

ee 2

kit

t holbroo

Cambusic

1. GAMBUSIA HOLBROOKII.

Fleterandria holbrooktt Gir:

Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 82, pl. x1.

figs. 4-13.

Haplochilus melanops Cope, Proc. Amer. Phil. Soc. xi. 1877,

p. 457,

Zygonectes atrilatus Jord. & Brayton, Bull. U.S. Nat. Mus. x11.

1878, p. 84.

@. Depth of body 34 to 4 in the length, length of head 4.

Diameter of eye 3 to 34.in length of head, interorbital width 2.

30 scales in a longitudinal series, Dorsal 8; origin above

posterior end of anal, equidistant from base of pectoral and base

of caudal; longest rays 3 length of head, Anal 10. Pectoral

8 length of head; pelvics extending to origin of anal. Least

depth of caudal peduncle 2 length of head. Olivaceous ; usually

a, dark bar below the eye; a blackish spot above the vent; dorsal

and caudal fins with small dark spots.

¢. Origin of dorsal fin equidistant from eye and base of

caudal.

Virginia to Alabama.

Here described from specimens from North Carolina and

Virginia, 3 females 38-48 mm, in total length and 2 males of

26 mm.

2. GAMBUSIA PATRUELIS,

Heterandria patruelis Baird & Girard, Proc. Acad. Philad. 1854,

p. 390.

Gambusia patruelis Girard, U.S. Mex. Bound, Surv., Fish. p. Us

pl. xxxix. figs, 1-7.

2. Depth of body 3 in the length, length ef head 4. Diameter

of eye 34 in length of head, interorbital width 2. 31 or 32

scales in a longitudinal series. Dorsal 7; origin behind end of

base of anal, nearer to base of caudal than to base of pectoral ;

longest rays 2 the length of head. Anal 10; first branched ray

longest. Pectoral 4 length of head; pelvies extending to vent.

Least depth of caudal peduncle # length of head, Brownish

above, yellowish below ; dorsal and caudal fins with small dark

spots.

Florida to Texas.

Here described from two specimens from Pensacola (Jordan),

AT and 50 mm, in total length,

3. GAMBUSIA AFFINIS.

Heterandria afinis Baird & Girard, Proe. Acad. Philad, 1854,

| B05

js Gambusia afinis Girard, U.S. Mex. Bound. Surv., Fish. p. 72,

pl. xxxix. figs. 12-15 (1859); Giinth. Cat. Fish. vi. p. 336

(1866).

Gambusia speciosa Girard, Proc, Acad. Philad, 1859, p. 121.

Gambusia gracihs Girard, |. ¢.

Gambusia humilis Giinth. t. ¢, p. 334.

CYPRINODONT FISHES. 985

Zygonectes brachypterus Cope, Bull. U.S. Nat. Mus. xx. 1880,

p- 34.

Zygonectes inurus Jord. & Gilb. Proc. U.S. Nat. Mus. 1882,

p- 043.

2. Depth of body 33 to 44 in the length, length of head

31 to 41, Diameter of eye 32 to 4 in length “of Head) interorbital

width re to 2. 30 or 31 scales in a longitudinal series. Dorsal

6-7; origin above posterior part or end of anal, equidistant from

some part of operculum or base of pectoral and base of caudal ;

longest rays 4 to $ length of head. Anal 10-11. Pectoral from

2 to nearly as lone as head; pelvics reaching vent or origin of

omell Brownish or olivaceous above, yellowish or silvery below ;

scales dark-edged; usually a dark suborbital bar; dorsal and

caudal fins with small dark spots.

Florida to Tampico; Mississippi.

Numerous females, measuring up to 50 mm. in total length,

from Florida and Louisiana; one from Tampico.

4, GAMBUSIA SENILIS. (Text-fig. 168 E.)

? Heterandria nobilis Baird & Girard, Proc. Acad. Philad. 18538,

ps 390;

¢Gambusia nobilis Girard, U.S. Mex. Bound. Surv., Fish. p. 71,

pl. xxxix. figs. 8-11 (1859).

Gambusia senilis Girar d, Proc. Acad. Philad. 1859, p. 122

2. Depth of body 3 to 3} in the length, length of head 32 6 4,

Diameter of eye 34 in length of head, interorbital width 2.

30 scales in a longitudinal series. Dorsal 8-9; origin above

middle or posterior ‘part of anal, equidistant from base of pectoral

and base of caudal ; longest rays 3 length of head. Anal 9-10.

Pectoral 2 length of head ; pelvics reaching vent. Least depth

of caudal ‘peduncle 2 a length of head. Brownish above, yellowish

below; scales dark- ed ged, especially on sides of abdomen ; a dark

suborbital bar and a “dusky. lateral band from eye to base of

caudal.

3. Origin of dorsal fin equidistant from preeoperculum and

base of caudal.

Chihuahua, Mexico,

Several females, 45 to 50 mm. in total length, and one

male of 25 mm.

5. GAMBUSIA NICARAGUENSIS. (‘Text-fig. 168 A.)

Gambusia nicaraguensis Ginth, Cat. Fish. vi. p. 336 (1866), and

Trans. Zool, Soc. vi. 1868, p. 483, pl. irxxii. fig. 3; Regan, Biol.

Centr.-Amer., Pisces, p. 96 (1907).

Paragambusia nicaraguensis Meek, Publ. Columbian Mus., Zool.

v. 1904, p. 135.

2. Depth of body about 3 in the length, length of head

31 to 4. Diameter of eye 3 to 3% in length of head, inter-

orbital width 2. 28 scales‘in a longitudinal ° series. Dorsal 7-8 ;

986 MR. C. TALE REGAN ON

origin above or a little behind posterior end of anal, twice as far

from middle of eye as from base of caudal; longest rays # or 2

length of head. Anal 11; second branched ray longest, edge of

fin emarginate. Pectoral a little shorter than head; _pelvics

reaching vent or origin of anal. Least depth of caudal peduncle

2 length of head. Upper part of body with 2 or 3 series of small

dark spots along the rows of scales; dorsal and caudal fins

spotted.

3. Origin of dorsal fin twice as distant from end of snout as

from base of caudal.

Southern Mexico; Nicaragua.

Several females, 40 to 55 mm. in total length, and two males

of 30 mm., including the types from Lake Nicaragua, and speci-

mens from El Hule (Aeek) and Coaxacoaleos (Arnold).

6. GAMBUSIA DOVII, sp. n.

2. Depth of body 3? in the length, length of head 3}. Dia-

meter of eye 31 in length of head, sneer bital width 13. 32 scales

in a longitudinal series. Darel 7; origin behind ond of anal,

twice as “ion from end of snout as from base of caudal. Anal 10;

anterior branched rays longest, but edge not emarginate. Least

depth of caudal peduncle slightly more than } length of head.

Olivaceous ; traces of spots on back and on caudal fin.

Lake Nicaragua.

A single specimen of 33 mm., collected by Captain Dow.

7. GAMBUSIA CAUDOVITTATA.

Regan, Ann. Mag. Nat. Hist. (8) xii. 1913, p. 471.

Recently described from a single specimen from the Condoto,

a tributary of the San Juan, W. Colombia.

8. GAMBUSIA PUNCTATA. (Text-fig. 168 D.)

Gambusia punctata Poey, Mem. i. p. 384 (1855); Giinth.

Cat. Fish. vi. p. 334 (1866); Jord. & Everm. Bull. U.S. Nat.

Mus. xlvii. 1896, p. 680.

2. Depth of body 3 to 4 in the length, length of head 34

to 4. Diameter of eye 3 to 31 in length of head, interorbital

width 2. 31 to 33 scales in a ‘longitudinal series. Dorsal 10;

origin above posterior end of anal, nearer to base of caudal than

to head; longest rays 2 to 1 the length of head. Anal 11; last

simple or first. branched ray longest ; edge of fin straight or

slightly emarginate. Pectoral # length of head pelvics re aching

vent. Least “depth of caudal ‘peduncle tt o 8 length of nena

Upper part of side with a dark spot on each scale, forming 3 to 5

regular longitudinal series; usually small dark spots on dorsal

and caudal.

3. Dorsal origin about equidistant from head and base of

caudal ; longest rays } length of head.

Cuba.

Here described from 16 females, measuring up to 85 mm. in

total length, and 3 males of 40 to 50 mm.

CYPRINODONT FISHES, 987

9. GAMBUSIA NIGROPUNCTATA, sp. n. (Text-fig. 168 F.)

2. Depth of body 34 to 32 in the length, length of head

32 to 32, Diameter of eye 3 to 3+ in length of head, interorbital

width 2 to 21. 30 scales in a longitudinal series. Dorsal 9:

origin above posterior part of anal, equidistant from base of

pectoral and base of caudal. Anall0-11. Least depth of caudal

peduncle less than # length of head. Coloration as in related

species.

3. Depth of body 4 in the length, length of head 32.

Diameter of eye 3 in length of ea mieror pital width mL,

Dorsal origin equidistant from middle of opercle and base of

caudal ; longest rays > length of head. Pectoral 2 length of

head.

Cuba.

Here described from two females of 34 and 50 mm. anda male

of 25 mm. from Fermina, Bemba.

10. GAMBUSIA MELANOSTICTA, Sp. 0.

2. Depth of body 3 in the length, length of head 34 to 3!.

Diameter of eye 3 to 31 in the Tenesh of head, interorbital

width 2. 29 to 30 scales in a longitudinal series. Dorsal 9-10;

origin ALOE end of anal, equidistant from base of pectoral ain

base of caudal ; longest rays } to 2 engin of head. Anal 10-11;

first. branched ray Tlongest, Pectoral 2 to 2 length of head ;

pelvics reaching vent. ‘Least depth of cma peduncle 2 a leneth

of head. Brownish above, golden below; upper par ts with

seattered small dark spots; dorsal and caudal fins spotted.

Cuba.

Three females from Havana, 35 to 42 mm. in tctal length,

received from Dr. D.S Jordan.

11. GAMBUSIA PUNCTICULATA.

Gambusia puncticulata Poey, Mem. 1. pp. 386, 390, pl. xxxi.

figs. 6, 7 (1855); Gunth. Cat. Fish. vi. p. 334 (1866) ; Garman,

Mem. Mus. Comp. Zool. xix. 1895, p. 87.

@. Depth of body 34+ to 34 in the length, length of head

4 to 41. Diameter of eye 33 in length of head, interorbital

width 2. Mouth smaller than in any other species of the genus.

29 or 30 scales in a longitudinal series. Dorsal 9; origin above

middle or posterior part of anal, equidistant from head and base

of caudal; fin rounded, the longest rays 3 length of head.

Anal 11, pointed. Pectoral 2 or # length of head ; pelvics small.

Least depth of caudal peduncle 3 " length of head. Upper parts

with scattered small dark spots ; 2 or 3 series of small black spots

on dorsal and caudal fins.

3. Origin o dorsal equidistant from eye and base of caudal;

longest rays 3 length of head.

Cuba.

Here described from two females, 35 mm. in total length, pre-

sented by Capta:n Vipan.

988 MR. CG. TATE REGAN ON

12. GAMBUSIA MELANOPLEURA.

Pecilia melanopleura Gosse, Soj. in Jamaica, p. 84, pl. i. fig. 3

(1851).

Haplochilus melanopleurus Giinth. Cat. Fish. vi. p. 317 (1866).

Gambusia melanopleura Garman, Mem. Mus. Comp. Zool. xix.

1895, p. 88.

Depth of body 3 in the length, length of head 33. Diameter

of eye 3 to 31 in length of head, interorbital width 1}. 30 or 31

scales in a longitudinal series. Dorsal 11-12; origin above

posterior part of anal, equidistant from base of pectoral and base

of caudal; longest rays 4 length of head. Anal 10-11; branched

rays slightly decreasing from first or second. Pectoral 7 length

of head; pelvics reaching vent. Least depth of caudal peduncle

3 or 2 length of head. Traces of spots on dorsal and anal fins

and in some of a few spots on the back.

Jamaica. :

Several females, the largest 47 mm. in total length, types of the

species.

13. GAMBUSIA WRAYT, sp. n. (PI. XCIX. figs. 3, 4, and Text-

fig. 168 B.)

2. Depth of body 33 to 4 in the length, length of head

32 to 34. Diameter of eye 3 to 33 in length of head, inter-

orbital width 2. 31 to 383 scales in a longitudinal series.

Dorsal 8; origin above posterior part of anal, equidistant from

base of pectoral and base of caudal; longest rays 3 length of

head. Anal 10-11; second branched ray longest. Pectoral

8 length of head; pelvics reaching vent. Caudal rounded or

subtruneate. Least depth of caudal peduncle 3 to 2 length of

head. Coloration probably as in G. gracilior.

3. Dorsal origin equidistant from middle or posterior part of

operculum and base of caudal; longest ray # length of head.

Least depth of caudal peduncle # or 2 length of head.

Jamaica.

Hight females measuring up to 55 mm. in total length, and

four adult males of 35 to 40 mm., collected by C. A. Wray.

14. GamBustIA oLicosticta, sp.n. (Pl. XCIX. figs. 1, 2, and

Text-fig. 169 B.)

2. Depth of body 22 to 34 in the length, length of head

3 to 34. Diameter of eye 3 to 34 in length of head, inter-

orbital width 2. 29 to 31 scales in a longitudinal series. Dorsal

8-9; origin above last 2 or 3 rays of anal, equidistant from base

of caudal and base of pectoral; fin rounded, longest rays 4 length

of head. Anal 10-11; first branched ray longest, the rest

regularly decreasing. Pectoral length of head ; pelvics reaching

vent. Caudal rounded or subtruncate. Least depth of caudal

peduncle 2 or 2 the length of head. Brownish above, yellowish

below ; edges of scales darker; usually a dark lateral stripe and

a few scattered spots; dorsal and caudal with small spots.

CYPRINODONT FISHES. 989

$. Dorsal origin equidistant from eye or postorbital part of

head and base of caudal; longest rays 2 the length of head.

Jamaica.

Six females, measuring up to 47 mm. in total length, and seven

males, the largest 33 mm. Some of these were collected by

C. A. Wray, others by the Rev. J. Seed Roberts.

15. GAMBUSIA GRACILIOR, sp. n. (PI. XCIX. figs. 5, 6, and

Text-fig. 168 C.)

2. Depth of body equal to or a little more than length of

head, which is 3} (young) to 4 in the length of the fish.

Diameter of eye 3 in length of head, interorbital width 2.

31 to 33 scales in a longitudinal series. Dorsal 8-9; origin

above end of anal, a little nearer to base of caudal than to base

of pectoral ; fin rounded, longest rays } length of head. Anal

LO=11; first or second br: snched rays longest, the rest regularly

decreasing. Pectoral as long as head without snout; pelvics

reaching vent. Caudal rounded or subtruneate. Caudal ‘peduncle

slender, its least depth + the length of head. Brownish above,

yellowish below; usually a narrow dark lateral stripe, some-

times a few scattered dark spots; dorsal and caudal with small

spots.

3S. Dorsal further forward and more elevated ; origin about

equidistant from middle of opercle and base of caudal; longest

vays about 2 the length of head. Least depth of caudal peduncle

3 the length of head.

_ Jamaica. —

Hight females, measuring up to 50 mm. in total length, and

five males, the largest 37 mm., collected by C. Wray. These were

mixed up with the examples of G. oligosticta, and could be picked

out by their more slender form and smaller head, characters

found to be associated with differences in the structure of the

intromittent organ. ‘

16. GAMBUSIA DOMINICENSIS, sp. n. (PI. XCIX. fig. 7, and

Text-fig. 169 C.)

2. Depth of body 34 in the length, length of head 3%,

Diameter of eye 34 in length of head, interorbital width 2.

28 or 29 scales in a longitudinal series. Dorsal 9; origin above

posterior end of anal, nearer base of caudal than base of pectoral ;

longest rays 3 length of head. Anal 10-11; first branched ray

longest. Pectoral # length of head; pelvics reaching origin of

anal. Least depth of caudal peduncle 2 length of head.

Brownish, scales dark-edged; a faint lateral stripe; a few

scattered dark spots on upper parts; abdomen golden; dorsaland

caudal fins with series of small dark spots.

¢. Dorsal origin equidistant from head and base of caudal.

Haiti.

A female of 52 mm., and two males, each 25 mm. in total

length, presented by Herr A. Rachow.

Proc, Zoou. Soc.—1913, No. LX VI. 66

990 MR. C. TATE REGAN ON

17. GAMBUSIA CAYMANENSIS, Sp. n.

Depth of body 3} to 4 in the length, length of head 3 to 31,

Diameter of eye 3 to 34 in length of head, interorbital width

Text-fig. 169.

}

== Spy

male

Distal part of intromittent organ of A. Gambusia holbrookii. B. G. oligosticta.

C. G. dominicensis. D. Brachyrhaphis rhabdophora. HK. Belonesow beli-

zanus. FE. Alfaro amazonum.

CYPRINODONT FISHES. 991

2 to 24. 30 or 31 scales in a longitudinal series. Dorsal 9 ;

origin above middle of anal, equidistant from operculum and base

of caudal ; longest rays nearly $ length of head. Anal 10; first

branched ray longest. Pectoral # length of head; pelvics reaching

vent. Least depth of caudal peduncle 4 length of head. Oliva-

ceous ; scales dark-edged; a dark bar below eye; dorsal fin with

two series of dark spots; caudal faintly spotted.

Grand Cayman.

Two females, 22 and 26 mm. in total length.

4. Brtonzesox Kner, 1860.

Sitzungsb. Akad. Wien, xl. p. 419.

Related to Gambusia, but the jaws produced, with broad bands

of slender, pointed, depressible teeth. Anal fin of male very

similar to that of Gambusia, differing in that the terminal seg-

ments of the first produced ray are not serrated, while rather

small retrorse spines at the ends of the third and the posterior

branch of the second produced rays represent the hooks of

Gambusia.

A single species.

BELONESOX BELIZANUS. (Text-fig. 169 E.)

Belonesox belizanus Kner, Sitzungsb. Akad. Wien, xl. 1860,

p- 419, fig.; Giinth. Cat. Fish. vi. p. 333 (1866); Meek, Publ.

Columbian Mus., Zool. v. 1£04, p. 135; Regan, Biol. Centr.-Amer.,

Pisces, p. 93 (1907).

Depth of body 4 to 6 in the length, length of head nearly 3.

Snout 4 the length of head. 55 to 65 scales in a longitudinal

series. Dorsal 9-10. Anal 10-12, in advance of dorsal.

Atlantic Slope of Vera Cruz, British Honduras, and Guatemala.

The largest species of the group, the males attaining 100 mm.,

the females 200 mm. in total length. Here described from ten

specimens from Perez, Belize, and Lake Peten.

5). PRIAPICHTHYS, gen. nov.

Differs from Gambusia in that the intromittent organ is longer,

when laid back nearly reaching the caudal fin, and is quite

differently formed ; the terminal part is hooked forward and the

prolonged rays taper distally and have no specially modified

segments, except the usual serrz of the posterior branch of the

second ; the first prolonged ray does not extend to the apex, and

the hook is supported by the anterior branch of the second and

the third (text-fig. 170 B).

It is here assumed that five species from Central America and

Colombia are congeneric with P. annectens, the only species of

which I have examined males; in all but P. parismina males are

known and agree with those of P. annectens in their long intro-

mittent organ.

66*

992 MR. C. TATE REGAN ON

Synopsis of the Species.

T. Origin of dorsal fin in, or a little in advance of, middle of

entire length (including caudal fin), above anterior part or

middle of anal.

A. D. 10-12; no black spot on anal fin ...................:...... anmnectens.

B. D. 8-9; a blackish spot on anal fin.

Length of head 33 to 32 in length of fish (without caudal) ......... episcopi.

Length of head 43 to 4% in length of fish (without caudal) ......... nigroventralis.

II. Origin of dorsal fin much nearer to end of caudal fin than

to tip of snout, above posterior part or posterior end of

anal. D. 7-8.

A blackish spot on anal fin .. ue nagonogovevasccaoonca fXGMPOSTOONR.

7 or 8 dark vertical bars on posterior part. of body... spnosnaancosse — UPC AOCOEPRo

8 or 9 dark bars on anterior # of body ee eee eR eI CLUE IULOCULCTUS7 Ss

1. PRIAPICHTHYS ANNECTENS. (Text-fig. 170 B.)

Gambusia annectens Regan, Ann. Mag. N. H. (7) a, IOOG,

p. 259, and Biol. Centr.- Amer., I2nCSS, [9s Oi, jk omy, IGE, Ds 6

(1907).

Costa Rica.

2. PRIAPICHTHYS EPISCOPI.

Gambusia episcopi Steind. Sitzungsb. Akad. Wien, Ixxvul. 1878,

p. 387, pl. 11, figs. 3, 4.

Panama.

3. PRIAPICHTHYS NIGROVENTRALIS.

Gambusia nigroventralis Kigenm. Indiana Univ. Studies, 1912,

No. 8, p. 26.

Rio San Juan, Colombia.

4, PRIAPICHTHYS PARISMINA.

Gambusia parismina Meek, Publ. Field. Mus., Zool. x. 1912,

Oa Mlk:

Costa Rica.

5. PRIAPICHTHYS TRIDENTIGER.

Gambusia tridentiger Garman, Mem. Mus. Comp. Zool. xix.

1895, p. 89.

Panama.

. PRIAPICHTHYS TURRUBARENSIS.

phage turrubarensis Meek, Publ. Field Mus., Zool. x. 1912,

joa Ibs

Costa Rica.

Perhaps a synonym of P. tridentiger.

6. PRIAPELLA, gen. nov.

This genus shows relationship to Priapichthys in the structure

of the long intromittent organ as well as in other characters.

CYPRINODONT FISHES. 993

PRIAPELLA BONITA. (Text-fig. 170 E.)

Gambusia bonita Meek, Publ. Columbian Mus., Zool. v. 1904,

p. 132, fig. 39; Regan, Biol. Centr.-Amer., Pisces, p. 95

(1907).

Rio Papaloapam in Mexico.

7. PskupoxreHoPHorws Bleek., 1863.

Atl. Ichth. iii. p. 140; Giinth. Cat. Fish. vi. p. 332 (1866).

Differs from Priapichthys in the longer dorsal fin originating

in advance of the anal (9) and in the structure of the intro-

mittent organ (text-fig. 170 C), with the anterior branch of the

second prolonged ray ending in an unsegmented antrorse hook,

at the base of which the first and third prolonged rays terminate.

There are three species from Mexico and Central America, but

males of one only (P. bimaculatus) have been described.

1. PSEUDOXIPHOPHORUS TERRABENSIS.

Gambusia terrabensis Regan, Ann. Mag. N. H. (7) xix. 1907,

p- 260; Biol. Centr.-Amer., Pisces, p. 97, pl. xii. fig. 7 (1907).

Dorsal 12-14; origin nearly equidistant from tip of snout and

base of caudal. Anal 9-10; origin nearly below middle of dorsal.

No dark spot above pectoral.

Rio Grande de Terraba, Costa Rica.

2. PSEUDOXIPHOPHORUS JONESII.

Mollienisia jones Giinth. Ann. Mag. N. H. (4) xiv. 1874,

otal

: Pseudoxiphophorus pauciradiatus Regan, Ann. Mag. N. H. (7)

xii. 1904, p. 256 and xvi. 1905, p. 362.

Gambusia jonesti Regan, Biol. Centr.-Amer., Pisces, p. 97, pl. xii.

fig. 8 (1907).

Dorsal 11-13; origin equidistant from tip of snout and middle

or posterior part of caudal. Anal 9-11, below anterior part of

dorsal. A dark spot above pectoral.

Mountain lakes and streams of Central Vera Cruz, Mexico.

3. PSEUDOXIPHOPHORUS BIMACULATUS. (Text-fig. 170 C.)

Aiphophorus bimaculatus Heck. Sitzungsb. Akad. Wien, i. 1848,

[> ASO, Jol, me sles, My A,

Pseudoxiphophorus bimaculatus Ginth, Cat. Fish. vi. p. 332

(1866) ; Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 81.

Gambusia bimaculata Regan, Biol. Centr.-Amer., Pisces, p. 98,

pl. xiv. fig. 4.

Dorsal 13-17 ; origin nearly equidistant from tip of snout and

994 MR. C. TATE REGAN ON

base of caudal. Anal 9-11, below anterior part of dorsal. A

dark spot above pectoral.

Southern Mexico, British Honduras, and Guatemala.

Text-fig. 170.

Distal part of intromittent organ of A. Heterandria formosa. B. Priapichthys

annectens. C. Pseudoxiphophorus bimaculatus. D. Poeciliopsis presidionis.

KE. Priapella bonita. ¥. Pseudopeeilia feste.

CYPRINODONT FISHES. 995

8. Hereranpria, Agass., 1853.

Amer. Journ. Sci. xvi. p. 135.

Girardinus (part.) Giinth. Cat. Fish. vi. p. 351 (1866).

Heterandria (part.) Garman, Mem. Mus. Comp. Zool. xix. 1895,

[Ds BOs

Differs from Gambusia and Priapichthys in the small transverse

mouth and compressed pointed teeth, the enlarged outer teeth

forming a close-set series. The intromittent organ is elongate,

as in Priapichthys, and its structure indicates a close relationship

to that genus. It differs in that the third prolonged ray is

scarcely longer than the posterior branch of the second, so that

the hook is supported only by the anterior branch of the second.

Two species from the United States and Mexico.

1. HereranpriA Formosa. (Text-fig. 170 A.)

Heterandria formosa Agass. Amer, Journ, Sci. xix. 1855, p. 136 ;

Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 91, pls. iv. fig. 7,

viii. fig. 8, and xi. figs. 1-3 ; Jord. & Everm. Bull. U.S. Nat. Mus.

xlvii. 1896, p. 687.

Girardinus formosus Giinth. Cat. Fish. vi. p. 354 (1866).

Depth of body 32 to 4 in the length, length of head 33

to 4. Diameter of eye 3 to 32 in length of head, interorbital

width 2. 28 to 30 scales in a longitudinal series. Dorsal 7-8 ;

origin above middle of anal, scarcely nearer to end of caudal than

to tip of snout. Anall0. A dark lateral band crossed by several

vertical bars ;,a blackish spot on basal part of dorsal, another on

anal.

South Carolina to Florida.

Hight specimens, the largest female 30 imm., the males 15 to

20 mm. in total length.

2. HETERANDRIA FASCIATA.

Gambusia fasciata Meek, Publ. Columbian Mus., Zool. v. 1904,

p. 129, fig. 37; Regan, Biol. Centr.-Amer., Pisces, p. 95 (1907).

Very similar to the preceding in form and coloration, except

for the absence of the lateral band, differing especially in the

more posterior dorsal fin, its origin above end of anal and much

nearer to end of caudal than to tip of snout. Dorsal 8. Anal

9-10. 32 scales in a longitudinal series.

Southern Mexico.

Total length 55 mm.

Six specimens from Oaxaca and Tequesixtlan (Gadow).

The male of this species has not yet been described.

9. PSEUDOPGCILIA, gen. nov.

Closely related to Heterandria, but with the teeth broad

truncated incisors and the intromittent organ somewhat different

in structure.

996 MR. C. TATE REGAN ON

PSEUDOPGCILIA FESTH. (Text-fig. 170 F.)

Pecilia feste Bouleng. Boll. Mus. Zool. Torino, xii. 1898,

No. 329, p. 13.

Santa Elena, Western Ecuador.

10. PasciLiopsis, gen. nov,

This genus has the mouth and dentition of Pecilia, but in the

males the pelvic fins are neither enlarged nor modified, and the

intromittent organ is long and slender. The first prolonged ray

and the anterior branch of the second extend to the end of the

fin and are somewhat curved forward distally; the posterior

branch of the second is short and the third still shorter and

slender distally, so that the serrations of the second project

beyond it (text-figs. 170 D and 171 B).

Mexico, Central America, and Colombia.

Synopsis of the Species.

1. Dorsal origin behind that of anal.

Sides of body with a series of vertical bars ......... 1. presidionis.

A lateral series of spots, pave more than 8 in number and

smaller than the eye 2. lutzi.

A lateral series of 4 to 8 spots, , each about as lar: oe | as the eye 3. pleurospilus.

No bars or spots Se Metteaaetaesanees rc puede 4. retropinna.

2. Dorsal origin in advance of that of anal.

Diameter of eye 3 to 34 in length of head, which is 3} to 32 in

the length of fish .......... .... 5. isthmensis.

Diameter of eye 2% to en in length of head, ‘which is 4 to 4h in

the length of fish . Las diaratinelst bisGemiaus Cannel et aa tenes eieenesen), OTM DUCUCeIAUs

1. Pa@cILiopsis PRESIDIONIS. (Text-fig. 170 D.)

Pecilia presidionis Jord. & Culver, Proc. Calif. Aead. (2) v.

1895, p. 413, pl. xxix.

Girardinus presidionis Regan, Biol. Centr.-Amer., Pisces, p. 99

(1907).

As I have stated in the ‘ Biologia’ this form and the next seem

to differ from P. pleurospilus only in coloration, and should perhaps

be regarded as subspecies of that species.

Rio Presidio in Sinaloa.

Heterandria colombianus Kigenm. (Indiana Univ. Studies, 1912

No. 8, p. 27), from brackish water at the mouth of the R. Dagua,

Colombia, seems to be extremely similar to P. presidionis.

2. Pa@cILIoPsIs LUTZI.

Heterandria lutzi Meek, Publ. Colombian Mus., Zool. v. 1904,

p. 148, fig. 47.

Girardinus lutzi Regan, Biol. Centr.-Amer., Pisces, p. 99

(1907).

Oaxaca, Mexico; R. Motagua in Guatemala.

CYPRINODONT FISHES. 997

3. PdCILIOPsIs PLEUROSPILUS.

Girardinus pleurospilus Giinth. Cat. Fish. vi. p. 355 (1866),

and Trans. Zool. Soc. vi. 1868, p. 486, pl. Ixxxvil. fig. 1; Regan,

Biol. Centr.-Amer., Pisces, p. 100 (1907).

Lakes Duenas and Nacasil in Guatemala.

4, PacILIOPsIS RETROPINNA.

Pecilia retropinna Regan, Ann. Mag. Nat. Hist. (8) 11. 1908,

p- 458.

Costa Rica.

. P@CILIOPSIS ISTHMENSIS, sp.n. (PI. C. figs.3, 4; and Text-

fig. 171 B.)

@. Depth of body 23 to 3 in the length, length of head 35 to 33.

Diameter of eye 3 to 34 in length of head, interorbital width

14 to 2. 26 to 28 scales in a longitudinal series. Dorsal 9-10;

origin equidistant from anterior edge of eye and base of caudal;

longest ray # length of head. Anal 10; origin below fourth or

fifth ray of dorsal: ; first branched ray longest, 3? length of head or

more. Pectoral a little shorter than head ; pelvics reaching anal.

Least depth of caudal peduncle 2 or ? length of head. Olivaceous ;

scales dark-edged ; a blackish spot aber ‘the vent; fins dusky, the

dorsal with blackish basal band and dark edge.

3. Dorsal origin equidistant from tip of snout and base of

caudal, or nearer former; intromittent organ, when laid back,

nearly or quite reaching caudal fin; indistinct cross-bars on body.

Colon, Panama,

8 females, 35 to 60 mm. in total length, and 4 males of 35 to

42 mm., presented by Herr A. Rachow.

6. PascILIOPsIs PITTIERT.

Pecilia pittierts Meek, Field Mus. Publ., Zool. x. 1912, p. 71.

Closely related to the preceding, but described as with a smaller

head and lax ger eye.

La Junta, Costa Rica.

Total length 43 to 65 mm., males to 53 mm.

11. BrRacHyRHAPHIS, gen. nov.

Differs from Gambusia in the shorter intromittent organ ending

ina small retrorse hook formed by the second and third pro-

duced rays.

BRACHYRHAPHIS RHABDOPHORA, (Text-fig. 169 D.)

Gambusia rhabdophora Regan, Ann. Mag. Nat. Hist. (8) ii.

1908, p. 457.

Costa Rica,

Specimens recently received show that in this species the origin

of the anal fin may be behind below or in advance of that of the

dorsal.

998 MR. C. TATE REGAN ON

Gambusia wmbratilis Meek (Publ. Field Mus., Zool. x. 1912,

p- 70) seems to differ from B. rhabdophora only in the larger

eye.

12. LeprorHaPHIS, gen. nov.

Differs from Gambusia only in the structure of the intromittent

organ, which is longer and more slender than in that genus and

slightly curved backwards distally. The first produced ray is not

serrated, gradually tapers distally, and extends nearly to the tip

of the fin; the second is slightly longer and has the posterior

branch almost as long as the anterior and with the usual serra-

tions, which project externally ; the third ray also seems to reach

nearly to the tip of the fin and appears to meet the first and form

a complete tube.

LEPTORHAPHIS INFANS.

® Xiphophorus gracilis Heck. Sitzungsb. Akad. Wien, i. 1848,

p. 300, pl. ix. figs. 3, 4.

Gambusia infans Woolman, Bull. U.S. Fish. Comm. 1894,

Text-fig. 171.

Distal part of intromittent organ of A. Phalloptychus januarius and

B. Peciliopsis isthmensis. 'The rays have been separated.

CYPRINODONT FISHES. 999

p- 62, pl. 11. fig. 3; Meek, Publ. Columbian Mus., Zool. v. 1904,

p- 131, fig. 38; Regan, Biol. Centr.-Amer., Pisces, p. 96 (1907).

Gambusia gracilis Meek, t. c. p. 130.

Rio Lerma and Rio Balsas in Mexico; ? Orizaba.

13. PHatitorprycHus Higenmann.

Proc. U.S. Nat. Mus. xxxii. 1907, p. 430.

Mouth and dentition of Pecilia. Intromittent organ long

and slender; third produced ray crossing second on right side

near base, meeting first and forming a closed tube; second

twisted distally so that the serrations are directed laterally or

even forwards.

PHALLOPTYCHUS JANUARIUS. (PI. C. figs. 7, 8, and text-

terse L/S)

Girardinus januarius Hensel, Arch. f. Nat. xxxiv. 1868,’

p. 360, and xxxv. 1869, p. 89.

Girardinus theringw Bouleng. Ann. & Mag. Nat. Hist. (6)

iv. 1889, p. 266.

Girardinus zonatus Schreiner, Arch. Mus. Rio Janeiro, xii.

LOSS ps We

S.E. Brazil; La Plata.

This species may at once be recognized by its coloration,

having several narrow blackish vertical stripes on each side of

the body. Dorsal 9; origin above ora little behind that of anal.

14. PHALLOcCEROS Higenm.

Proc. U.S. Nat. Mus. xxxii. 1907, p. 431.

Mouth small, but bones of lower jaw rather firmly joined ;

teeth oar-shaped, slender, curved, as in Pecilia. Intromittent

organ long (about # length of fish to base of caudal) ; last segment

of first produced ray modified into an antrorse appendage which

bifurcates, each fork antler-like in form ; anterior branch of

second ending just beyond the first in an antrorse process.

PHALLOCEROS CAUDOMACULATUS. (PI. C. figs. 5, 6, and text-

fig. 172 C.)

Girardinus caudimaculatus Hensel, Arch. f. Nat. xxxiv. 1868,

p. 362, and xxxv. 1869, p. 89.

Glaridodon janwarius Garman, Mem. Mus. Comp. Zool. xix.

1895, p. 42.

S.E. Brazil; La Plata.

The coloration readily distinguishes this species from Phallo-

ptychus januarius. Plain olivaceous, usually with a vertical

blackish spot on side below dorsal fin ; sometimes a lateral series

of dusky bars. Dorsal 7-8; origin above middle or posterior

part of anal.

1000 MR. C. TATE REGAN ON

15. CNESTERODON Garman.

Mem. Mus. Comp. Zool. xix. 1895, p. 43.

Gulapinnus Langer, Morph. Jahrb. xlvii. 1913, p. 207.

Mouth small, but rami of lower jaw rather firmly joined and

teeth chisel-shaped, broader than in Pecilia. Intromittent

organ long (nearly 2 length of fish to base of caudal); last

segment of first produced ray an appendage which is not forked,

but is very long, peculiarly curved, and pointed distally; second

ending in a process which is not or scarcely antrorse ; third

terminating in a hook, as in Gambusia.

CNESTERODON DECEMMACULATUS. (Text-fig. 172 B.)

Pacilia decemmaculata Jenyns, Zool. ‘ Beagle,’ Fish. p. 115,

pl. xxii. fig. 1 (1842).

Pecilia gracilis Cuv. & Val. Hist. Nat. Poiss. xviii. 1846,

p- 133.

Cnesterodon decemmaculatus Garman, Mem. Mus. Comp. Zool.

xix. 1895, p. 44.

Cnesterodon carnegiet Haseman, Ann. Carnegie Mus. vii. 1911,

p. 385, pl. Ixxxiil.

La Plata; Rio Grande do Sul.

A small species, easily recognized by the lateral series of dark,

rounded or vertically expanded spots. Dorsal 8-9; origin above

or a little behind that of anal.

16. GLARIDICHTHYS Garman, 1896.

Glaridodon (non Seeley, Proc. R. Soc. xliv. 1888, p. 135);

Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 40.

This genus is well distinguished by the dentition and by the

structure of the intromittent organ. It includes a single species

from Cuba and perhaps one from Chihuahua, but males of the

latter have not been described.

1. GLARIDICHTHYS UNINOTATUS.

Girardinus uninotatus Poey, Mem. ii. pp. 309, 383 (1861);

Giinth. Cat. Fish. vi. p. 351 (1866).

Glaridodon uninotatus Garman, Mem. Mus. Comp, Zool. xix.

1895, p. 41.

Glaridichthys faleatus Higenm. Bul]. U.S. Fish. Comm. xxii.

p. 224, fig. (1904).

Glaridichthys torralbasi Kigenm. t.c. p. 225, fig.

2. Depth of body 3} to 44 in length, length of head 4 to 5.

Hye large, more than 4 of length of head. 28 to 31 scales in a

longitudinal series. Dorsal 9-11; origin nearly equidistant from

head and base of caudal. Anal 10; origin well in advance of

that of dorsal. Olivaceous ; edges of scales usually darker ; often

1001

CYPRINODONT FISHES.

Text-fig. 172.

intromittent organ of A. Girardinus metallicus. B. Cnesterodon

Distal part of

caudomaculatus. D. Platypecilus macu-

latus. KE. Xiphophoru

1002 MR. C. TATE REGAN ON

a lateral series of short vertical bars; usually a blackish spot

above origin of anal fin.

3. Dorsal origin nearly equidistant from eye and base of

caudal. Bars on side well marked; lateral spot indistinct; a

blackish streak behind first produced ray of anal.

Here deseribed from several specimens up to 75 mm. in total

length from Cuba, including a male of 47 mm. extremely similar

to the type of G. torralbasi, and co-types of G. faleatus. The last

is a pale variety without the lateral spot, but structurally similar

to G. uninotatus; the slight production of the vertical fins is

found in several specimens with the typical coloration.

2. GLARIDICHTHYS LATIDENS.

Glaridodon latidens Garm. Mem. Mus. Comp. Zool. xix. 1895,

p. 42.

Chihuahua ; Mexico.

17. Toxus Higenm., 1904.

Bull. U.S. Fish. Comm. xxii. p. 226.

This genus differs from the preceding only in the dentition.

TOXUS RIDDLEI.

Higenm. |. c¢. figs.

Cuba.

18. GrrarpDrINus Poey, 1855.

Mem. i. pp. 383, 390; Garman, Mem. Mus. Comp. Zool. xix.

1895, p. 45.

Mouth and dentition as in Pecilia, but intromittent organ

formed exactly as in the two preceding genera.

A single species from Cuba.

GIRARDINUS METALLICUS. (Text-fig. 172 A.)

Poey, Mem. i. pp. 387, 391, pl. xxx1, figs. 8-11 (1855); Giinth.

Cat. Fish. vi. p. 351 (1866); Garm. Mem. Mus. Comp. Zool. xix.

1895, p. 46.

Girardinus denticulatus Garm, t.¢. p. 47.

Girardinus creolus Garm. |. ¢.

Girardinus garmani Higenm. Bull. U.S. Fish. Comm. xxii.

p. 226, fig. (1904).

Heterandria cubensis Kigenm. t. c. p. 227, fig.

Depth of body 3 to 4 in length, length of head 4 to 41.

Diameter of eye 23 to 3 in length of head. 27 to 31 scales in a

longitudinal series. Dorsal 9-10; origin nearly equidistant

from base of pectoral and base of caudal (a little nearer head

than caudal in fg). Anal 10-12; origin well in advance of

dorsal. Olivaceous; edges of scales darker; usually a lateral

CYPRINODONT FISHES. 1003

series of dark bars; dorsal with anterior edge dark and usually

a blackish spot or band posteriorly near base.

Cuba.

Several specimens up to 70 mm. in total length.

19, PAMPHORICHTHYS, gen. nov.

Mouth small; bones of lower jaw firmly united; a series of

firmly set, slender, pointed teeth and a band of much smaller

teeth within. Intestine long, coiled. Males with anal and

elvic fins below pectorals, the former a slender pointed organ

about + the length of the fish (without caudal), the latter also

modified, the outer rays produced, the inner about 4 as long and

forming a projection near their ends.

This genus is based on Garman’s Heterandria minor, which

seems to differ from Heterandria in the more slender teeth,

longer intestine, and shorter intromittent organ, but especially in

the modification of the pelvic fins in the male.

PAMPHORICHTHYS MINOR.

Heterandria minor Garman, Mem. Mus. Comp. Zool. xix.

1895, p. 92.

Amazon.

20. PAMPHORIA, gen. nov.

Mouth small; bones of lower jaw rather firmly joined; a series

of broad incisors and a band of minute tricuspid teeth within,

Intestine long.- Males with anal fin much advanced, forming an

intromittent organ that is longer than the head, and with the

second ray of the pelvics produced.

PAMPHORIA SCALPRIDENS.

Cnesterodon scalpridens Garman, Mem. Mus. Comp. Zool. xix.

1895, p. 45, pl. v. fig. 12. pl. viii. fig. 17.

Amazon.

21. Piatypacitus Giinth., 1866.

Cat. Fish. vi. p. 350.

Mouth small, transverse ; bones of lower jaw loosely connected ;

an outer series of slender curved teeth, somewhat expanded and

compressed towards their apices, spoon-shaped or oar-shaped.

Males with the pelvic fins enlarged, nearly as long as the intro-

mittent organ, which is unprotected by a hood, but has the

distal end remarkably modified (text-fig. 172 D).

Two species from Mexico, with the intromittent organ precisely

similar in structure.

1, PLATYP@CILUS COUCHIANUS.

Linvia couchiana Girard, Proc. Acad. Philad. 1859, p. 116.

Pecilia couchiana Regan, Biol. Centr.-Amer., Pisces, p. 104,

pleavarticoIN( 190m):

1004 MR. C. TATE REGAN ON

Depth of body about 2? in the length, length of head about 4.

23 to 27 scales in a longitudinal series. Dorsal 9-11; origin

equidistant from snout and anterior part of caudal. Anal 7-8,

opposite posterior 4} of dorsal. Blackish or brownish above,

yellowish below, the two colours sharply separated.

Rio San Juan at Monterey.

2, PLATYPa@cILUS MACcULATUS. (Text-fig. 172 D.)

Platypecilus maculatus Giinth. Cat. Fish. vi. p. 350 (1866).

Pecilia maculata Regan, Biol. Centr.-Amer., Pisces, p. 105

(1907).

Closely related to P. couchianus. Depth of body 2 to 22 in

the length, length of head 34 to 32. Anal 8-9. Olivaceous; a

dark spot or crescent or a pair of spots at base of caudal.

Atlantic Coast streams of Mexico and Guatemala.

22. XipHopHorus Heckel, 1848.

Sitzungsb. Akad. Wien, 1. p. 291.

Closely related to the preceding, as is shown by the great

similarity in the structure of the anal fins of the males (text-fig.

172K). The species occur in rivers of the Atlantic Slope from

Tamaulipas to Guatemala. After examination of a large series

of specimens I conclude that only 3 species are well established.

1. XIPHOPHORUS MONTEZUM2.

Jord. & Snyder, Bull. U.S. Fish. Comm. 1900, p. 131, fig. 11 ;

Regan, Biol. Centr.-Amer., Pisces, p. 107 (1907).

Dorsal 11-13. Anal 6-8. Scales 27 to 29. Depth about 3

in the length, length of head about 4. Yellowish; scales of

upper part of body dark-edged ; usually a dark lateral stripe.

Rio Panuco, Mexico.

2. XIPHOPHORUS HELLERI. (Text-fig. 172 EK.)

Heck. Sitzungsb. Akad. Wien, i. 1848, p. 291, pl. viil.; Giinth.

Cat. Fish. vi. p. 349 (1866); Regan, Biol. Centi.-Amer., Pisces,

p. 107 (1907).

Xiphophorus guentheri Jord. & Everm. Bull. U.S. Nat. Mus.

xlvii. 1896, p. 702.

Xiphophorus jalape Meek, Publ. Columbian Mus., Zool. v.

1903, p. 136, pl. x1.

NXiphophorus strigatus Regan, t.c. pl. xiv. fig. 7.

Dorsal 11-14. Anal 8-10. Scales 26 to 30. Depth 3 to 4

in the length, length of head 4 to 42. Sides of body blue, with

red stripes between the series of scales.

Southern Mexico ; Guatemala.

Mexican examples have nearly always a strong stripe from eye

to base of caudal (strigatis), the males sometimes with another

backwards from axil of pectoral (helleri, jalape); in specimens

CYPRINODONT FISHES, 1005

from Guatemala there is, as a rule, no strong lateral stripe

(guentheri):

Xiphophorus rachovii Regan (Ann. Mag. Nat. Hist. (8) viii.

1911, p. 373), from Puerto Barrios, Guatemala, has a pair of

alinge spots at the base of the caudal foe other examples received

later from Puerto Barrios lack these spots, but are not very

different in other respects. Whether this is another variety of

X. helleri or a hybrid with Platypecilus maculatus, as some

aquarium-writers in Germany believe, still remains to be settled.

3. XIPHOPHORUS BREVIS.

Regan, Ann. Mag. Nat. Hist. (7) xix. 1907, p. 65, and Biol.

Centr.-Amer., Pisces, p. 108, pl. xiv. figs. 8, 9.

Dorsal 13-15. Anal 9-10. Scales 27. Depth 24 to 22 in

length, length of head 32 to 32. No strong lateral stripe.

British Honduras.

23. Pacri1A Schneider, 1801.

Bloch’s Syst. Ichth. p. 452.

Pecilia (part.) Giinth. Cat. Fish. vi. p. 339 (1866); Garman,

Mem. Mus. Comp. Zool. xix. 1895, p. 52.

Mouth small, transverse; teeth slender, curved, expanded

towards the tip, spoon- or oar-shaped, forming a series with a

band of minute teeth within; bones of lower jaw loosely con-

nected. Pelvic fins in male enlarged, the second ray prolonged.

Intromittent organ short, its tip protected anteriorly by a cuta-

neous hood; first prolonged ray without terminal spine, with

several segments not far from the end forming acute serrations ;

last ray simple, without appendages (text-fig. 173 A-C).

As now restricted this genus only includes four (or five)

species from South America.

Synopsis of the Species.

I. Dorsal 7-9; origin nearly abovelthatotanalu ners esse eee 1. vivipara.

II. Dorsal 6-7.

Origin of dorsal above posterior end of anal; usually a dark spot

above pectoral, equidistant from head and dorsal fin ...... 2. pare.

Origin of dorsal above middle or posterior part of anal; a dark

spot on each scale and dark stripes between the series of

SGAIES Gin SIGS Or |WOGhY soosdonsasnonc0 souBoodon aouiddesduadéode Hoe unboRReDD 3. picta.

Origin of dorsal above middle of eee a blackish spot or bar

at base of caudal fin .. noddonee candog cet onan cdacoooanécasennasanaoas — Gly MRED

1. Pa@criia vivipaRA. (Text-fig. 173 C.)

Pecilia vivipara Schneid. Bloch’s Syst. Ichth. p. 452, pl. Ixxxvi.

fig. 2 (1801) ; Gunth. Cat. Fish. vi. p. 344 (1866); Garman, Mem.

Mus. Comp. Zool. xix. 1895, p. 53.

Pecilia schneideri Val. in Humboldt, Obs. Zool. ii. p. 159

(1828); Cuv. & Val. Hist. Nat. Poiss. xviii. p. 135 (1846).

Proc. Zoou. Soc.—1913, No. LX VII. 67

1006 Mk. C. TATE REGAN ON

Pecilia surinamensis Val. t. c. p. 158, pl. li. fig. 1; Cuv. & Val.

t.c. p. 120.

Pecilia unimaculata Val. t.c. p. 158, pl. li. figs. 2,5,6; Cuv. &

Val. t.c. p. 128; Giinth. t.c. p. 346.

2. Depth of body 22 to 4 in the length, length of head 34 to 44.

Diameter of eye 3 to 4 in length of head, mterorbital width

12 to 2. 25 to 27 scales in a longitudinal series. Dorsal 7-9 ;

origin nearly above that of anal and equidistant from some part

of operculum and base of caudal. Anal 8-10. Pectoral a little

shorter than head; pelvics reaching vent. Least depth of caudal

peduncle ? to # length of head. Olivaceous ; edges of scales usually

darker ; body often with dark cross-bars ; often a blackish spot,

which may be pale-edged, on upper part of side 2 or 3 scales in

front of dorsal fin; often a pair of blackish spots at base of caudal

above and below, extending along margins of fin ; fins sometimes

immaculate, but dorsal and caudal sometimes with series of small

spots, or dorsal with one or two blackish bands,

3. Dorsal a little further forward and higher than in 9.

Venezuela and Leeward Islands to the La Plata.

Numerous examples, the largest female 80 mm. and the

largest male 60 mm.

2. Pacinia PARE. (Text-fig. 173 B.)

Pecilia vivipara pare Kigenm. Ann. N.Y. Acad. vii. 1894,

p- 629.

? Pacilia amazonica Garman, Mem. Mus. Comp. Zool. xix.

1895, p. 64.

Acanthophacelus bifurcus Kigenm. Mem. Carnegie Mus. v.

1912, p. 459, pl. Ixv. figs. 4-6.

2. Depth of body 34 in the length, length of head 3}.

Diameter of eye 34 in length of head, interorbital width 2. 28

scales in a longitudinal series. Dorsal 6 ; origin above posterior

end of anal, nearly equidistant from head and base of caudal ;

middle rays longest, 4 length of head. Anal 8-9. Pectoral ?

length of head; pelvics extending to origin of anal. Least

depth of caudal peduncle 3 length of head. Olivaceous ; usually

a pale lateral stripe; a vertically expanded dark spot with pale

margin on upper part of side, nearly equidistant from head and

dorsal fin ; fins immaculate.

3. Dorsal origin nearer to head than to base of caudal ;

longest rays ? length of head. A dark spot above pectoral (as in

2), a larger one just above anal fin, a third on caudal peduncle ;

dorsal and caudal sometimes spotted, latter with dark upper edge.

Amazon; Guiana.

Four specimens, two females of 28 mm. and two males of 22

-and 24 mm. in total length, received from Prof. Eigenmann ;

these are co-types of the species from Para and of A. bifureus

from British Guiana. The intromittent organ is figured

(text-fig. 173 B).

Garman’s description of P. amazonica applies to P. pare,

eee

CYPRINODONT FISHES. 1007

except that the dorsal origin is said to be a little in advance of

that of anal.

3. Pacruia PrcTaA, sp. n. (PI. C. figs. 1, 2, and Text-fig. 173 A.)

Acanthophacelus melanzonus (part.) Kigenm. Ann. Carnegie

Mus. vi. 1909, p. 51; Mem. Carnegie Mus. v. 1912, p. 457,

pl. Ixiv. fig. 6.

2. Depth of body 53 to 4 in the length, length of head about 4.

Diameter of eye 33 to 4 in length of head, interorbital width 14.

26 to 29 scales ina longitudinal series. Dorsal 6-7, origin above

middle or posterior part of anal and equidistant from base of

pectoral and base of caudal; fin small, rounded. Anal 9-10,

pointed. Pectoral # length of head; pelvics reaching vent or

origin of anal. Least depth of caudal peduncle about # length of

head. Olivaceous; sides of body with a dark brown spot on each

scale and dark brown longitudinal stripes between the series of

scales.

3. Depth 3 to 34 in the length. Dorsal origin nearly equi-

distant from eye and base of caudal; fin elevated. Least depth

of caudal peduncle 3 to ? length of head. Coloration sometimes

as in female, but usually there are a few large dark spots on

posterior part of body; dorsal fin usually with blackish spots ;

often an ocellus on upper part of base of caudal fin.

Demerara.

Numerous examples, the females measuring up to 45 mm.,

the males to 30 mm. in total length, presented in 1872 by F. G.

Beckford, Esq.

A. melanzonus.is based on a male of Lebistes reticulatus (type)

and several females of P. picta.

In this species the intromittent organ differs somewhat from that

of P. vivipara, but is formed on the same plan (text-fig. 173 A, C).

4. P@cILIA BRANNERI.

Peeilia brannert Eigenm. Ann, N.Y. Acad. vii. 1894, p. 629.

Pecilia heteristia Regan, Ann. Mag. Nat. Hist. (8) iii. 1909,

p. 235.

2. Depth of body 33 in the length, length of head 4. Dia-

meter of eye 33 in length of head, interorbital width nearly 2.

27 or 28 scales in a longitudinal series. Dorsal 6-7; origin

above middle of anal and nearly equidistant from base of pectoral

and base of caudal. Anal 8, pointed. Pectoral ? length of head ;

pelvics extending to origin of anal. Least depth of caudal

peduncle 2 length of head. Olivaceous ; edges of scales darker ;

some blackish vertical streaks on the side and a blackish spot or

vertical bar at base of caudal fin.

3. Dorsal origin equidistant from middle of operculum and

base of caudal; two last rays produced into long filaments.

Caudal spot larger than in Q ; sometimes a blackish stripe near

upper edge of caudal fin.

67*

1008 MR. C. TATE REGAN ON

Para.

Here described from the types of P. heteristia, ¢ and 9, each

35 mm. in total length.

The intromittent organ is very like that of P. vivipara, but the

anterior branch of the second ray has fewer segments, as in

J OUCH

24, Lepistes Filippi, 1861.

Arch. Zool, Anat. Fisiol. i. p. 69.

Acanthophacelus Kigenmann, Proc. U.S. Nat. Mus. xxxii. 1907,

p- 426, fig. 1.

Differs from Pecilia only in that the third prolonged anal

ray of the male has its last segments modified into a pair of

projections which are directed obliquely outwards and towards

the base of the fin.

A single species from South America.

LEBISTES RETICULATUS. (Text-fig. 173 D.)

Pecilia reticulata Peters, Monatsb. Akad. Berlin, 1859, p. 412;

Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 458.

Lebistes pecilioides Filippi, Arch. Zool. Anat. Fisiol. 1. 1861,

p- 69, pl. iv. f. 6; Giinth. Cat. Fish. vi. p. 356 (1866).

Girardinus reticulatus Gunth. t. c. p. 352.

Girardinus guppy Ginth. t.c. p. 353; Regan, Proc. Zool. Soc.

1906, p. 390, pl. xxi. figs. 1, la.

Acanthophacelus reticulatus Kigenm. Proc. U.S. Nat. Mus.

xxxil. 1907, p. 426, f. 1; Mem. Carnegie Mus. v. 1912, p: 458,

pl. Ixv. figs. 1-3.

Acanthophacelus melanzonus (part.) Eigenm. Mem. Carnegie

Mus. v. 1912, p. 457, pl. Ixiv. fig. 5.

2. Depth of body 3 to 4 in the length, length of head 3 to 44.

Diameter of eye 3 to 33 in length of head, interorbital width 14

to 2. 26 to 28 scales in a longitudinal series. Dorsal 7-8,

rounded or obtuse; origin equidistant from eye or operculum and

base of caudal. Anal 9-10, pointed; origin below or a little in

advance of that of dorsal. Pectoral $ length of head ; pelvics ex-

tending to origin of anal. Least depth of caudal peduncle 2 2to ?

length of head, Olivaceous; edges of scales darker, forming a

agin: fins immaculate, exeEpe sometimes a small done spot on

middle of caudal,

3. Dorsal more elevated than in 9, when laid back nearly

reaching base of caudal fin; pelvics nearly as long as the short

intromittent organ. @oloration very variable ; often a blackish

spot above or behind base of pectoral, another above anal fin or

on middle of side, a third at base of caudal peduncle; often also

several bluish-silvery spots, which may unite to form a lateral

band, margined above and below by dark longitudinal stripes ;

dorsal and caudal sometimes spotted, often dark-edged.

Venezuela and St. Lucia to Santos.

1009

CYPRINODONT FISHES. |

Text-fig. 173.

Distal part of intromittent organ of :—A. Pecilia picta. B. P. pare. C. P. vivi-

EF. Mollienisia

KE. Limia versicolor.

s reticulatus.

D. Lebist

sphenops. ‘The pre

pare.

receding figures note that these ave reversed, the first

y being on the right, the third on the left.

uce-like hood is indieated in fig.

Tn comparing with p

prolenged 1

1010 MR. C. TATE REGAN ON

Numerous examples, including co-types of the species received

from the Berlin Museum and the types of Girardinus guppyi ;

females to 5d mm., males to 25 mm., in total length.

25. Moutrenis1a Le Sueur, 1821.

Journ. Acad. Philad. ii. p. 3; Giinth. Cat. Fish. vi. p. 347

(1866); Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 49.

This genus differs from Pacilia in having a small antrorse

spine at the end of the first prolonged ray of the male anal fin,

as in Limia,and at the end of the third a paired process, probably

the modified terminal segment, directed obliquely outwards and

towards the base of the fin, as in Lebistes (text-fig. 173 F),

Some species hitherto referred to Pecilia (M. sphenops,

M. gracilis) have an intromittent organ exactly similar to that of

Molhienisia latipinna and M. petenensis, and as the little-known

M. formosa is intermediate between the two groups in the size of

the dorsal fin, all may be placed in JMollienisia. Some other

species, with the dorsal fin further back than the anal, are pro-

visionally included ; all that is known of the males is Garman’s

statement that in P. cwneata the anal fin of the male is shorter

than the head, indicating that the species is neither a Peciliopsis

nor a Girardinus, although it may be a Limia.

Synopsis of the Species.

I. Dorsal 11-16; in adult males origin nearer end of snout than

base of caudal, base longer than distanee from caudal.

A. Caudal rounded or subtruncate (@ ) or lower angle slightly

produced (4); scales 28 to 80. Dorsal 15-16 ............. 1. petenensis.

B. Caudal rounded; scales 26 to 28.

Dorsal 14-16; in females origin equidistant from end of snout

and base of caudal (young) or nearer snout (adult); base equal

to (young) or more than (adult) distance from caudal .

Dorsal 11-13; in females origin a little nearer base of caudal than

end of snout, even in the adult ; base equal to (adult) or less

than (young) distance from cauiclalee Male ore ers ee Cee 3. formosa.

II. Dorsal 10-11; origin nearly equidistant from end of snout

and base of caudal fin; base shorter than distance from

caudal; anal origin below fourth ray of dorsal; scales 28

POND O)) Peper erate case seins sip ye ONE cave etiolee neaosmee Serteetee ue nae Meee CUCU LOSE

III. Dorsal 7-11; origin nearly equidistant from end of snout

and middle or posterior end of caudal fin; base shorter than

distance from caudal.

. latipinna.

A. Origin of anal below or a little behind that of dorsal ...... 5. sphenops.

B. Origin of anal in advance of that of dorsal.

Dorsal 8-10. Anal 8-10. Scales 29-33 . 6. elongata.

Dorsal 7-9. Anal 9-11. poe: 27-380.. 7. oceidentalis.

Dersal 8. Anal 9. Scales 2 8. spilwrus.

1. MoLuLIEeNIsiA PETENENSIS.

Giinth. Cat. Fish. vi. p. 348 (1866), and Trans. Zool. Soc. vi.

1868, p. 485, pl. cart. figs. 1-3; Regan, Biol. Centr.-Amer.,

Pisces, p. 106 (1907).

CYPRINODOND FISHES. LOLi

2. Depth of body 23 to 3 in the length, length of head 4.

Diameter of eye 4 in length of head, interorbital width 13. 28

to 30 scales in a longitudinal series. Dorsal 15-16; base about

as long as its distance from en of SOUS, longer than its distance

from caudal fin; longest rays 3 or 2 2 length of head. Anal 9-10;

origin below middle of dorsal. Pector al as long as head ; pelvics

reaching origin of anal. Caudal rounded or subtruncate. Least

depth of caudal peduncle nearly equal to length of head. A

small spot on each scale, most conspicuous posteriorly; dorsal

and caudal with series of small spots.

3. Base of dorsal 14 to 13 its distance from end of snout and

about twice its distance from caudal; longest rays longer than

head. Anal below anterior part of dorsal; produced rays 2 length

of head ; second pelvic ray about as long. Lower angle of caudal

slightly produced. Spots on lower part of dorsal forming undu-

lating stripes; a series of large spots, one on each interradial

membrane, along middle of fin. Lower margin of caudal blackish.

Lake Peten in Guatemala.

Three females, 100 to 110 wm., and three males, 100 to 130 mm.

in total length, types of the species.

. MOLLreNniIsiA LATIPINNA.

t Sueur, Journ. Acad. Philad. 11. 1821, p. 3, pl. ii.; Cuv. &

Val. Hist. Nat. Poiss. Sanu, [os Bos jole WAT 3 Giimth. Cat. Fish. vi.

p. 348 (1866) ; Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 50,

jolts 30g § dork, ay Everm. Bull. U.S, Nat. Mus. xIvii, 1896, p. 699.

Pecilia multilineata Le Sueur, t. ¢. p. 4.

Pacilia lineolata Girard, U.S. Mex. Bound. Sury., Fish. pac Or

pl. xxxv. figs. 9-11 (1859).

Lima pecilioides Girard, t.c. p. 70, pl. xxxviil. figs. 8-11 ;

Proc. Acad. Philad. 1859, p. 115.

Limia matamorensis Girard, Proc. Acad. Philad. 1859, p. 116.

2. Depth of body 23 to 3 in the length, length of head 33 to 4.

Diameter of eye 34 to 4 in length of head, interorbital width 14

to 2. 26 to 28 scales in a longitudinal series. Dorsal 14-16; in

adults base somewhat shorter than its distance - ‘om end of snout,

longer than its distance from caudal; rays 3 length of head.

Anal 9-10; origin below middle of dorsal. Pectoral a. little

shorter than head; pelvics reaching vent or origin of anal.

Caudal rounded. Least depth of caudal peduncle about ? length

of head. Spots on scales forming longitudinal sieinen: dorsil

and basal part of caudal with series of small spots.

3. Dorsal base, in adult, about 14 its distance from end of

snout or twice its distance from caudal; rays longer tonne head.

Anal below anterior part of dorsal ; produced rays 7 length of

head, a little longer than second pelvic ray ; amd rounded.

Spots on lower part of dorsal forming longitudinal stripes ; a

series of large spots, one on each interradial membrane, along

middle of fin; caudal spotted superiorly; with blackish lower

edge.

1012 MR. C. TATE REGAN ON

Southern United States.

Five females, 40 to 68 mm., and two males, 65 and 78 mm.,

from Pensacola and New Orleans.

3. MOLLIENISIA FORMOSA.

Limia formosa Girard, Proc. Acad. Philad. 1859, p. 115.

Mollienisia formosa, Giinth. Cat. Fish. vi. p. 349 (1866).

2. Depth of body 23 to 3 in the length, length of head 31 to 4.

Diameter of eye 3 to 32 1 in length of head, interorbital width 12

to 2. 26 to 28 scales in a longitudinal series. Dorsal 11- 14;

origin a little nearer to base of caudal than to end of snout ; base,

in adult, nearly equal to its dhisusinse from caudal, but less than its

distance from head; longest rays } to 2 2 length of head. Anal

9-10; origin below fourth ray of dorsal. Pectoral a little shorter

than head; pelvics reaching vent or or igin of anal. Caudal

rounded. Heast depth of camel peduncle 2 to ? length of head.

A spot on each scale ; dorsal usually spotted.

3. Dorsal origin nearer to end of snout than to base of caudal ;

base as long as its distance from eye; longest rays nearly ? 2 length

of head.

Atlantic slope of Mexico and Central America.

Several small specimens (¢ and @ ) up to 45 mm. from Tampico

(Jordan). Also three large females, 65 to 80 mm. in total length,

from Colon (Rachow) ; hese are structur ally similar to the bohene:

but have the coloration obscured by blackish mottling.

4. MOLLIENISIA GRACILIS, nom. nov.

Pecilia petenensis (non Moll tenisia petenensis Ginth.) Ginth.

Cat. Fish. vi. p. 342, and Trans. Zool. Soc. vi. 1868 »P- 484, pl. Ixxxv.

figs. 3, 4.

2. Depth of body 4 to 43 in the length, length of head 42

to 42. Diameter of eye 3} to 3? in length of head, inter orbital

width 2. 28 to 30 scales ina loneieudinall series. Dorsal 10- Lil

origin equidistant from preeorbital and base of caudal; anterior

branched rays 2 length of head. Anal 8-10, acutely. pointed ;

origin below fourth ray of dorsal. Pectoral as long as head.

Least depth of caudal peduncle ? length of head. Olivaceous;

scales dark-edged ; dorsal and cnndlel speckled.

3. Depth 33 to 3% in length ; depth of caudal peduncle equal

to length of head. Dorsal origin equidistant from end of snout

and base of caudal; posterior rays longest, as long as or longer

than head.

Lake Peten in Guatemala.

Two females, 120 and 150 mm. and three males, 95 to 120 mm.

in total length, types of the species.

5. MOLLrenisia SPHENOPS. (Text-fig. 173 F.)

Pecilia sphenops Cuv. & Val. Hist. Nat. Poiss. xviii. p. 130,

pl. 526 (1846); Regan, Biol. Centr.-Amer., Pisces, p. 102, pl. xiii.

(1907) (with synonymy).

CYPRINODONY FISHES. 1013

Girardinus Suse sesoaNs Steind. Dealeselau, Akad. Wien, xli. 1880,

p- 87, pl. vi. figs. 4, 5

Pecilia sulvatoris Regan, Ann. Mag. Nat. Hist. (7) xix. 1907,

p- 65; Biol. Centr.-Amer., Pisces, p. 104, pl. xiv. figs. 2, 3 (1907).

Pecilia amates Miller, Bull. Amer. Mus. xxii. 1907, p. 108.

? Platypecilus tropicus Meek, Publ. Columbian Mus., Zool. vii.

1907, p. 146.

Pecilia tenwis Meek, t. c. p. 147.

Pecilia spilonota Regan, Ann. Mag. Nat. Hist. (8) 1. 1908,

p. 460.

2. Depth of body 24 to 4 in the length, length of head 33

to 43. Diameter of eye 3 to 4 in length of head, interorbital

width 14 to 2. 25 to 30 scales in a longitudinal series. Dorsal

8-11; origin equidistant from eye or operculum and base of

caudal, Anal 8-10; origin below or a little behind that of

dorsal. Pectoral as long as or shorter than head. Least depth

of caudal peduncle equal to or less than length of head. Dark

greenish to pale olivaceous; edges of scales sometimes darker ;

often a dark spot on each scale of side of body ; sometimes narrow

vertical bars on sides; dorsal and caudal sometimes immaculate,

often with series of small spots, or with a dark basal spot which

may extend over most or all of the fin.

3. Dorsal further forward and more elevated ; spots on sides

rarely present, dark cross-bars often distinct.

Coasts and rivers from Sinaloa to Panama and from Tamau-

‘lipas to Colombia, Venezuela, and the Leeward Islands.

Numerous examples to 120 mm. in total length, from all parts

of the range of the species.

6. MOLLIENISIA ELONGATA.

Pecilia elongata Ginth. Cat. Fish. vi. p. 342, and Trans. Zool.

Soe. vi. 1869, p. 484, pl. Ixxxv. fig. 2; Regan, Biol. Centr.-Amer.,

Pisces, p. 102 (1907).

? Pecilia cuneata Garman, Mem. Mus. Comp. Zool. xix. 1895,

-p. 62.

Panama.

7. MOLLIENISIA OCCIDENTALIS.

Heterandria occidentalis Baird & Girard, Proc. Acad. Philad.

1853, p. 390.

Pecilia occidentalis Garman, Mem. Mus. Comp. Zool. xix. 1895,

p. 71; Regan, Biol. Centr.-Amer., Pisces, p. 102 (1907). ,

Pacific Coast Rivers from Arizona to Jalisco.

8. MoLLIENISIA SPILURUS.

Pecilia spilurws Gunth. Cat. Fish. vi. p. 345 (1866); Regan,

Biol. Centr.-Amer., Pisces, p. 101 (1907). —

Mexico o1 ential America.

1014 MR. C. TATE REGAN ON

26. Limra Poey, 1855.

Mem. Cuba, 1. p. 383.

Acropecilia Hilgendorf, Sitzungsb. Ges. Nat. Freund. 1889,

p. 52.

This genus is well marked off from Pacilia by the structure of

the intromittent organ (text-fig. 173 E). At some distance from

its end the first produced ray becomes abr uptly slender ; the distal

part is not serrated, and near its extremity it bears an antrorse

spine; the second and third produced rays are nearly as in

Pecilia. As in Pecilia, the intromittent organ is short, usually

shorter than the head, and its extremity is protected anteriorly

by a cutaneous hood; in fully adult males the second pelvic ray

is more or less prolonged.

Of eight species seven are from Cuba, Haiti, and Jamaica, and

one from Venezuela. I have examined adult males in all but

L. ornata.

Synopsis of the Species.

TI. Origin of dorsal fin about equidistant from middle of eye

and base of caudal; origin of anal below fifth ray of

OUOS SFT fies es eee omarpliatlar Ucn tn ugdan vad nan gameeosarobaccowues dl uouuuandae

TI. Origin of dorsal nearer to te of Peandedl re to eye.

A. Origin of anal below anterior part of dorsal.

Dorsal 8-9; length of head 33 to 4 in length of fish ............... 2. dominicensis.

Dorsal 10; length of head 32 to 4 in length of fish ............... 3. nigrofasciata.

Dorsal 10-11; length of head 3 to 3} in length of fish ............ 4. arnoldi.

B. Origin of anal below or very slightly in advance of that

of dorsal.

1. Length of head 2 length of fish (in adults of 50 mm.) ;

a lateral series of dark bars in both sexes.

Tuterorbital width less than } length of head 5. ornata.

Interorbital width 5 length of head, OLSMLONe wees eeee .. 6. caudofasciata.

2. Length of head less than 4 length of fish Gna a specimen

of 43 mm.). ¢ with 3 or 4 blackish cross-bars, g

with a blackish lateral stripe anteriorly............... 7. heterandria.

C. Origin of anal always distinctly in Ra vaaee’ of that of

dorsal ; head 4 length of fish (in adults of 50-60 mm.). 8. versicolor.

1. LIMIA VITTATA.

Pecilia vittata Guichenot in Ramon de la Sagra, Hist. Nat.

Cuba, Poiss. p. 146, pl. v. fig. 1 (1850); Giinth. Cat. Fish. vi.

p. 339 (1866); Garman, Mem. Mus. Comp. Zool. xix. 1895,

De Diss

Limia cubensis Poey, Mem. Cuba, 1. p. 388, pl. xxxi. figs. 12-13

(1855).

Limia vittata Poey, t. c. p. 389, pl. xxxi. figs. 14, 15.

? Platypecilus perugie Evermann & Clark, Proc. U.S. Nat.

Mus) xxx. 1906) ps 8515 fig:

2. Depth of body 24 to 3 in the Jength, length of head 33

to 48. Diameter of eye 3 to 34 in length of head, interorbitel

maiden 14 to 2.. 26 to 28 scales in a longitudinal series. Dorsal

CYPRINODONT FISHES, 1015

9-11; origin nearly equidistant from middle of eye and base of

caudal ; branched rays subequal, or the middle ones the longer,

3 or 3 2 length of head. Anal 10; origin below fifth ray of dorsal.

Pectoral = length of head or more ; pelvies reaching vent. Least

depth of caudal peduncle * to + length of Theme) Olivaceous ;

edges of scales darker ; often a dark lateral band or 1 to 3 series

of dark spots along the series of scales on middle of side ; ; dorsal

and caudal usually with small dark spots.

36. Dorsal origin equidistant from snout and base of caudal ;

posterior rays longest, longer than head in the adult. Intro-

mittent organ a little shorter than head; second pelvic ray

produced, inner rays ending ina knob. Least depth of caudal

peduncle nearly equal to length of head. Usually dark cross-bars

on body; spots on dorsal and caudal fins larger and blacker than

in females.

Cuba; ?San Domingo.

Several specimens from Cuba, the males 45 to 70, the females

40 to 100 mm., in total length. ;

2. LIMIA DOMINICENSIS.

Pecilia dominicensis Cuv. & Val. Hist. Nat. Poiss. xviii. p. 131,

pl. 526. fig. 1 (1846). :

Pecilia melanogaster Ginth. Cat. Fish. vi. p. 345 (1866).

Menigpoeen 8 Ge ees Evermann & Clark, Proc. U.S. Nat.

Depth of Body 2 Nes ie 3 a the length, length of head 32 to 4.

Diameter of eye 3 to 33 in length of head, interorbital width 13.

26 or 27 scales in a longitudinal series. Dorsal 8-93; origin

equidistant from occiput or operculum and base of caudal, or tip

of snout and end of caudal, first or second branched ray longest,

3 length of head. Anal 10; origin below second or third ray of

dorsal; anterior branched rays longest. Pectoral ? length of

head; pelvics reaching vent. Least depth of caudal peduncle

2 to #? length of head. Olivaceous; sometimes traces of dark

bars on upper part of sides and of a dark spot at base of caudal ;

a blackish spot on basal part of posterior 3 of dorsal fin; ripe

females with abdomen golden in front, blackish behind, the two

colours separated by a ventral line at insertion of pelvic fins.

San Domingo; ? Jamaica.

Seven specimens; two of the types of the species, ¢ and 2,

38 and 42 mm. in total length, and five females of 50 to 60 mm.,

types of P. melanogaster.

3. LIMIA NIGROFASCIATA, sp.n. (PI. CI. figs. 1, 2.)

2. Depth of body 23 to 3 in the length, length of head 33

to 4. Diameter of eye 34 to 33 in length of head, interorbital

width 14. 27 or 28 scales in a longitudinal series, Dorsal 10 ;

origin equidistant from cuidate of operculum and base of caudal ;

fin rounded, the longest rays 4 length of head. Anal 9, obtuse;

1016 MR. C. TATE REGAN ON

origin below anterior + of dorsal. Pectoral 3 length of head ;

pelvics not or barely reaching vent. Least depth of caudal

peduncle = or ? length of head. Body with 7 to 9 blackish

vertical bars ; a dark spot near base of posterior part of dorsal.

3. Depth 27 to 24 in the length. Dorsal origin equidistant

from eye or preoperculum and base of caudal; longest rays

3 length of head, or in adult nearly as long as bead,

Miragoane, Haiti.

Two females, 43 mm. in total length, an immature male of

41 mm., and an adult male of 48 mm., presented by Messrs. J.

Paul Arnold and A, Rachow.

4, LimraA ARNOLDI, sp.n, (PI. CI. fig. 5.)

2. Depth of body 3 to 3% in the length, length of head

3 to 33. Diameter of eye 3 to 34 in the length of head, inter-

orbital width 2. 26 or 27 scales in a longitudinal series. Dorsal

10-11; origin equidistant from postorbital part of head and base

of caudal ; fin rounded, the longest rays 3 length of head. Anal

9-10, rounded or obtusely pointed ; origin below anterior part of

dorsal. Pectoral 7? length of head; pelvics extending to vent.

Least depth of caudal peduncle 3 to # length of head. Body with

8 to 12 narrow dark vertical bars; dorsal sometimes spotted,

often with a small dark spot posteriorly near its base.

3. Depth of body 3 in the length. Dorsal origin equidistant

from eye and base of caudal; longest rays 2 length of head.

Intromittent organ } length of fish (to base of caudal); second

pelvic ray stout, a little produced beyond third and fourth, which

ave truncated distally. Least depth of caudal peduncle ? length

of head. No dark vertical bars, but several series of vertically

expanded spots.

Miragoane, Haiti.

Ten females, 26 to 42 mm. in total length, and three males of

28 to 38 mm., presented by Herr J. Paul Arnold.

5. Limia orNatA, sp.n. (Pl. CI. fig. 7.)

Depth of body 33 to 4 in the length, length of head 3 to 34.

Diameter of eye 34 to 4 in length of head, interorbital width

21 to 23. 28 scales in a longitudinal series. Dorsal 8-10;

origin equidistant from some part of operculum and base of

caudal; fin rounded, the longest rays 4 length of head. Anal 10;

origin below that of dorsal. Pectoral 2 or 3? length of head ;

pelvics reaching vent or origin of anal. Least depth of caudal

peduncle 2 the length of head. Head, body, and usually vertical

fins with blackish spots more or less developed ; body also with a

series of dark bars or large vertically expanded spots.

Haiti.

Five females, 31 to 55 mm. in total length, presented by Herr

J. Paul Arnold.

CYPRINODONT FISHES. 1017

6. Limia CAUDOFASCIATA, sp. n. (PI. CI. fig. 6.)

@. Depth of body 3 to 34 in the length, length of head

3 to 34. Diameter of eye 3} to 33 in length of head, inter-

orbital width 14 to 2. 26 scales in a longitudinal series. Dorsal

8-10; origin equidistant from some part of opsxomtou and base

of caudal; anterior branched rays longest, 4 length of head.

Anal 10; origin below or a little in advance of ‘Tak of dorsal.

Pectoral ? length of head; pelvics reaching vent. Least depth

of caudal peduncle 3 to # length of head. Olivaceous or

brownish ; edges of scales darker: a series of dark vertical bars

on posterior part of body ; a dark spot near base of posterior part

of dorsal; abdomen yellowish, or when tumid golden anteriorly

and blackish posteriorly, as in ZL. dominicensis.

3. Dorsal origin about equidistant from eye and base of

caudal; longest rays 3 to 2 (adult) length of head. Intromittent

organ less than } length of fish; second pelvic ray produced in

adult, nearly as long as prolonged anal rays. Least depth of

caudal peduncle 2 length of head.

Jamaica.

Several females, 28 to 42 mm. in total length, and six males of

25 to 48 mm., collected by C. A. Wray. In males of 25 to

30 mm., the anal fin has not attained the adult structure and the

second pelvic ray 1s scarcely prolonged; males of 35 to 48 mm.

have the anal fully formed, but only the largest has the pelvies

strongly produced.

7. LIMIA HETERANDRIA, sp.n. (PI. CI. figs. 3, 4.)

2. Depth of body 33 in the length, length of head 32 to 44.

Diameter of eye 3 to 31 in length ‘of head, interorbital width 2.

26 scales in a longitudinal series. Dorsal 8; origin equidistant

from some part of operculum and base of candle fin rounded,

longest rays a little more than 4 length of head. Anal 9,

pointed; origin below that of dorsal. Pectoral # length of head ;

pelvics reaching vent. Least depth of caudal peduncle 4 or #

length of head. Olivaceous; a dark longitudinal stripe from

above pectoral to level of origin of anal; a dark spot above vent ;

a black spot near base of posterior part of dorsal.

3. Dorsal origin egemchisceuat from middle of eye and base of

caudal ; longest rays 2 length of head. Intromittent organ about

2 length of fish (to ‘base “oi caudal) ; second ray produced, 3 3 as

inne as prolonged anal rays. Stripe on side of body and spot

above vent absent : body with 3 or 4 narrow dark vertical bars.

La Guayra, Venezuela.

Two females, 43 and 27 mm.,and a male of 25 mm. (adult),

presented by Herr J. Paul Arnoid.

8. Limia verstcotor. (Text-fig. 173 E.)

Pecilia donunicensis (non Cuv. & Val.) Giinth. Cat. Fish. vi,

1018 ON CYPRINODONT FISHES.

p- 346 (1866); Garman, Mem. Mus. Comp. Zool. xix. 1895,

oe Dee

Girardinus versicolor Giinth. t. ¢. p. 352.

Acropacilia tr idens Hilgendorf, Sitzungsb. Naturf. -Freunde,

1889, p. 52.

2. Depth of body 3 to 4 in the length, length of head

34 to 4. Diameter of eye 3 to 34 in leneth of head ; inter-

orbital width 12 to 2. 26 to 28 scales in a longitudinal series.

Dorsal 7-9 ; origin equidistant from some part of. operculum and

base of eam ‘middle rays longest, about 2 the length of head.

Anal 8-10; origin a little in advance of that of dor sel Pectoral

a little shorter ‘than head ; pelvics about reaching vent. Least

depth of caudal peduncle 2 402 = the length of Inn Olivaceous,

edges of scales darker; an indistinet Jateral band or a series of

short vertical bars ; sometimes irregular dark spots on the back ;

a blackish spot on base of posterior part of dorsal; caudal

sometimes with vertical bands.

3. In specimens of 30 mm. dorsal and pelvic fins as in 9, but

in those of 40 to 45 mm. dorsal more elevated, the longest rays

? length of head, second pelvic ray somewhat produced send i inner

alate rays expanded and truncated at ends.

San Domingo.

Ten females, 35 to 60 mm., including the types of the species,

and four “anoles, 30-45 mm.

EXPLANATION OF THE PLATES.

PratE XCIX.

2. Gambusia oligosticta, 9 & g.

4, Be wrayi,2? & SB.

, 6. a gracilior, 2 & g.

a es dominicensis, 9.

Pram C:

2. Pecilia picta, 2 & S.

, 4. Paciliopsis isthmensis, 2 & 3.

6. Phalloceros caudomaculatus, 2 & 3.

8. Phalloptychus januarius, 9 & 3.

PratTE CI.

. Limia nigrofasciata, 2 & g.

» heterandria, 2 & g.

» arnoldi, 2.

» caudofasciata, 6.

» ornata,

STH OVE bo

P. ZS) (er ie Gil

d.Green del. et lith.

1LBUFO HYPOMELAS. 2. AGALYCHNIS CALCARIFER.

tee arts) . USNS) IE, CHIL.

d.Green del.et. Chromo.

AGALYCHNIS SPURRELLEI.

IAS HOS) ell) (CIN.

J.Green del.et Chromo.

1. DENDROBATES AUROTZANIA

2-4.D.TINCTORIUS,var.COCTAI, 5-6.D.TINCTORIUS,vs.CHOCOENSIS.

P45: iss Pisey.

J.Green del.et lith.

CINOSTERNUM SPURRELLI.

learn oe ieols) JP i. (Wl

J.Green del.et ith

CINOSTERNUM SPURRELLI.

PP. Zeon LSP ies CMe

J.Green del.et lith.

LANOLIS BREVICEPS. 2. ANADIA VITTATA.

Ms Sa LORS. Pil. CVA.

—————

d.Green del.et lith.

1. HOMALOCRANIUM CORALLIVENTRE. 2.ELAPS MICROPS

&. LEPTOGNATHUS SPURREEEI

1913- Proc.Zool.Soc.London 1913:1019-38 ,figs.174-178

b1s.102-108

ON COLOMBIAN BATRACHIANS AND REPTILES. 1019

59. On a Collection of Batrachians and Reptiles made by

Dr. H. G. F. Spurrell, F.Z.S8., in the Choco, Colombia.

By G. A. BoutencEr, F.R.S., F.Z.S.*

{Received October, 10th, 1913: Read November 11th, 1913.]

(Plates CIT.—CVIII.7 and Text-figures 174-178.)

INDEX.

5 Page

Geographical Zoology: Choco, Colombia, Batrachians and

Reptiles... sates er Autenaetncereeo eres LOLS.

Development : bere of late omnes. Sicoatene Leeseranen LO2ZS

Ethology: Nesting-habits of Agalychnis spurr aIB.. LOA:

Variation: Dendrobates CPaerAl ie aen RL RUSS 1026

New species: Cecilia wmtermedta@ 2.2 0..c.028 senses LOZO

BF) I CUMTOGE Osecntitactiina bauer steed baaeeecesoos deo I MUr-Ut

Bufo hypomelas .. Jehiccsis watan ene neon steer O22

Agalychnis poor cE see Saanpnduce ORE!

Dendrobates paraensis (Cam Brazil) Jaisueaiee 1028

op GUROUGHOED con cornoecussenconcocencccs IKOFE)

Cinosternum spurrellt ...........1............-...-- 1030

AYRES OPRESUCGIB.:0 snocca noo cdaconcanocecconieccseoeces OBI

Anadia wittata ........... Becmecnnuoebaon cucie 40ssxs,

Homalocranium cor aUnenivel Weeeennuntn: LOSS:

JONCGIS MACPODE bacco socdooqovocass0 cen vpaeae svonenove NOS

Leptognathus spucamnclll.. ese LOSS

Dr. Spurrell, to whom the Zoological Society is indebted for so

many interesting additions to its collection of Reptiles, has now

transferred his activity as a collector from West Africa to South

America. The series of beautifully preserved Batrachians and

Reptiles brought together by him during the first few months

of his stay in the Choco, Colombia, and presented by him to the

British Museum, is one of great interest and shows how much

remains to be done in the exploration of that part of South

America. Unless otherwise stated, the specimens are from

Pena Lisa, Condoto, altitude 300 feet, Within the last few

years, series of Batrachians and Reptiles brought together by

Mr. M.G. Palmer in the same district had been acquired by the

Trustees of the British Museum and furnished the types of

several new species described by met. They showed a close

agreement with the fauna of N.W. Keuador, for a knowledge of

which we are indebted chiefly to Mx. Rosenberg §—an agreement

which is further exemplified by Dr. Spurrell’s collection.

Published by permission of the Trustees of the British Museum.

For explanation of the Plates see p. 1038.

Ann. & Mag. N. H. (8) ii. 1908, p. 515, and vii. 1911, p. 19.

Boulenger, P. Z.S. 1898, p. 107, and Ann. & Mag. N. H. (7) ix. 1902, p, 51.

wntt+ %

1020 MR. G. A. BOULENGER ON

BATRACHIA.,

APODA.

1. CxcILIA ISTHMICA Cope.

Known from the Isthmus of Darien and Western Heuador.

2. CMCILIA INTERMEDIA, sp.n. (Text-fig. 174.)

Intermediate between C. tentaculata L. and C. pachynema Gthr.

Snout as in the former, more strongly projecting than in the

latter. Outer mandibular teeth larger than in the former and

smaller than in the latter. Snout rounded, very prominent, as

long as the distance between the eyes; tentacle on the lower

surface of the snout, nearer the edge of the mouth than the

Text-fig. 174.

Cecilia intermedia.

A. Side view of head. B. Lower view of head and anterior extremity of body.

C. Lower view of posterior extremity of body.

nostril. Body cylindrical, its diameter 43 to 56 times in the

total length. 200 to 250 circular folds, mostly widely interrupted

on the back and on the belly ; from the posterior fourth of the

body short intermediate folds appear and gradually extend across

the back; the last. 60 to 80 folds close together and complete on

the back, the 15 to 30 hindermost completely encircling the body.

Uniform. blackish ; head of young yellowish or pale brown.

I have examined six specimens of this species, which was

COLOMBIAN BATRACHIANS AND REPTILES. 1021

first discovered in N.W. Ecuador by one of Mr. Rosenberg’s

collectors :-—

1. 8. Javier, N.W. Ecuador.—Total length 950 mm., dia-

meter of body 17.

2. Paramba, N.W. Ecuador, 3500 ft.—Total length 820 mm.,

diameter of body 19.

3. Pambelar, N.W. Ecuador.—Total length 690 mm., dia-

meter of body 12.

4, Pefia Lisa, Condoto, Choco (had been swallowed by an

Elaps rosenbergti 940mm. long).—Total length 690mm. ;

diameter of body 13 *.

Same locality (taken from the stomach of a Streptophorus

atratus swallowed by an Elaps corallinus).—VTotal length

280 mm.; diameter of body 6.

6. Same locality.—Total length 270 mm.; diameter of

body 5.

I avail myself of this opportunity to define a further species,

from a collection made‘in the Choco by Mr. M. G. Palmer, recently

acquired by the British Museum :—

Cecilia palmeri, sp. n. (Text-fig. 175.)—Head and dentition as

Text-fig. 175.

Cecilia palmeri.

Lettering as in text-fig. 174.

* The copulatory organ of this specimen is extruded; it measures 10 mm.

in length and terminates in a four-lobed “ glans.”

Proc. Zoot, Soc.—1913, No. LX VIII. 68

1022 MR. G; A. BOULENGER ON

in C. pachynema. Diameter of body 58 times in total length?

200 circular folds, mostly interrupted on the back and belly ;

from the posterior fourth of the body short interrupted folds

appear and gradually extend across the back; the last 95 folds

close together and complete on the back, the 16 hindermost com-

pletely encircling the body. Uniform blackish. Total length

700 mm. ; diameter of body 2s

A single specimen from Novita, Rio San Juan, 200 ft.

3. CHCILIA NIGRICANS Bler.

Described in 1902 from a single specimen from the Rio Lita,

N.W. Ecuador.

The specimen in Dr. Spurrell’s collection measures 395 mm.,

diameter of body 7.

ECAUDATA.

4, Buro HYPOMELAS, sp.n. (PI. CII. fig. 1.)

Crown without bony ridges; snout truncate, moderately pro-

minent, with angular canthus and vertical loreal region ; inter-

orbital space as br oad as the upper eyelid ; tympanum moderately

distinct, half diameter of eye. Fingers moderate, first not

extending as far .as second ; toes short, barely ihe webbed ;

subar Hoole tubercles feebly pr ominent, single ; ; two small meee

tarsal tubercles; no tarsal fold. Tlarso-metatarsal articulation

reaching the tip of the snout. Skin smooth ; parotoid gland

narrow, shorter than the head. Black, elegantly marked with

grey lines above, vermicular or ring-shaped ; a whitish streak on

each side in the lumbar region, and another on the arm ; lower

parts with small greyish-white spots.

From snout to vent 20 mm. ‘

This small toad, represented by a single specimen from the

upper waters of the Condo, altitude 1200 feet, differs from all its

American congeners in the shortness of the inner finger.

Buro Hamatiticus Cope.

Buro mARiNuS L.

Buro rtypHontus L.

Dr. Spurrell, in his notes, remarks: ‘Said to be the toad from

which Indians extract arrow-poison.” See further on, p. 1028.

8. Hyna MAxima Laur.

‘“¢ Apparently strictly nocturnal. Pupil of eye at night large

and round, in daylight diamond-shaped. One caught at night on

top of partition between two rooms in bungrlow was almost

uniform chocolate-brown with a dark median streak from snout

to middle of back. By daylight it turned to a hght yellowish

brown, with very pronounced markings of warm darker brown on

COLOMBIAN BATRACHIANS AND REPTILES. 1023

the body and limbs, greyer to sepia-brown on the head; a patch

of lighter yellow across loins. Iris golden brown; lower eyelid

veiled with gold.”

It is characteristic of this species to possess a rudiment of

pollex appearing as a small tubercle in females and a larger

tubercle in males. In the male sent by Dr. Spurrell, this pollex

projects externally as a long sharp bony spine at right angles to

the metacarpal of the inner finger and piercing the skin ; whether

this is due to the desiccation of the specimen, or occasionally

happens during life as in the case of the pollex of Rana holst or

of the ends of the ribs in the Pleurodele Newt, I cannot say.

Text-fig. 176.

Hyla maxima. Inner fingers of male.

9. Hyna BpAaupint Daud.

When taken out of a dark hole these frogs. are brown above,

usually with a metallic-green streak along the upper lip. After

exposure to light the colour changes to light yellowish grey, with

small green spots on the back and bars on the hind limbs. One

of the specimens, a female, is remarkable for the longer hind

limbs, the tibio-tarsal articulation reaching beyond the tip of

the snout.

These frogs were pairing on the 3lst May.

Larve, ab different stages of development, agree in essential

characters with those of H. arborea : position of eyes, spiraculum

and vent, shape of tail, structure of the mouth and lips. Three

long series of teeth in each lip, the third of the upper lip inter-

rupted, the others continuous; beak broadly edged with black.

Total length 60 mm.; body 21; width of body 13; tail 39;

depth of tail 16.

10. AGALYCHNIS CALCARIFER Blgr. (Pi. CII. fig. 2.)

Described in 1902 from a single specimen from the Rio

Durango, N.W. Ecuador, 350 feet.

. The ‘only example obtained by Dr. Spurrell is described in his

noes as brilliant sage-green above, flanks and upper surface of

thighs (with the exception of a narrow green streak) rich orange-

yellow with black bars; hands and feet and lower parts orange-

68*

1024 MR. G. A. BOULENGER ON

yellow; iris grey, bordered with orange-yellow ; lower eyelid

transparent, edged with turquoise-blue.

11, AGALYCHNIS SPURRELLI, sp.n. (PI. CIII.)

Tongue oval, feebly emarginate behind. Vomerine teeth in

two strong, transverse or slightly oblique series on a level with

the front edge of the rather large choane. Head moderately

depressed, as long as broad or slightly broader than long; snout

rounded, not projecting beyond the lower jaw; canthus rostralis

feeble ; loreal region concave and very oblique; interorbital space

broader than the upper eyelid ; tympanum close to the eye and

two-thirds to three-fourths its diameter. Fingers much flattened

and nearly entirely webbed, the disks nearly as large as the eye

or about two-thirds its size; toes rather short, much flattened,

entirely webbed (three-fourths webbed in the young), the disks a

little smaller than those of the fingers; subarticular tubercles

very prominent. The tibio-tarsal articulation reaches between

the eye and the tip of the snout. Skin smooth, granular on the

belly and on the basal half of the lower surface of the thigh; a

regular series of granules extends along the whole length of the

lower surface of the thigh ; a narrow dermal fold along the outer

edge of the forearm, the inner edge of the tarsus, and across the

heel ; a few scattered (white) flat warts on the back, Green above,

the dorsal warts, two to nine in number and irregularly disposed,

white, edged with violet-black ; belly yellowish white; iris ruby-

red ; lower eyelid with an open meshwork of golden lines. Male

without vocal sac, with a patch of black nuptial asperities on the

upper surface of the inner finger.

From snout to vent 95 mm.

Very closely allied to the Central American A. moreleti

A. Dum., this handsome frog differs in the more extensively

webbed digits, the presence of white warts on the back, and the

absence of vocal sac in the male.

Four specimens are in the collection. The first was reported

to Dr. Spurrell to have been found at the top of a high tree that

had been felled, on March 30th. On April 2nd Dr. Spurrell took

a pair in embrace on a leaf overhanging a pool of water two feet

below ; from the sketch accompanying his notes, the amplexus is

similar to that of Hylu arborea. The pair were in the act of

breeding, and the eggs, as they were extruded, were being fixed

to the upper surface of the terminal third of the leaf, in double

rows following more or less regularly the venation, the very

prominent ribs affording a support for their attachment. On the

leaf sent with the specimens, the breeding operations of which

were suddenly interrupted, 59 eggs are attached, and 7 more

adhere to the right foot of the female, showing that she uses her

feet for the purpose of fixing the eggs.

According to a sketch by Dr. Spurrell, which is here re-

produced (text-fig. 177), another leaf on the same stalk was

already entirely beset with eggs (laid by the same female 2)

COLOMBIAN BATRACHIANS AND REPTILES. 1025

and folded over to form a nest as in Phyllomedusa hypo-

chondrialis*. The eggs resemble those of Alytes obstetricans

in size (diameter 3 to 4 mm.) and in the tough gelatinous

capsule, but differ in the upper pole being brown and in not

being strung together.

Text-fig. 177.

Agalychnis spurrelli spawning.

The leaf, which I have not been able to get identified, probably

belongs to the family Anonacez.

12. LEPTODACTYLUS PENTADACTYLUS Laur.

13. Hy opEs pALMATUs Bler.

14, HyLopks consPprciLuAtus Gthr.

15. GLOSSOSTOMA ATERRIMUM Gthr.

Originally described from Costa Rica, this Engystomatid has

since been found on the Rio Durango, N.W. Ecuador,

* Budgett, Q. J. Micr. Sci. xlii. (1899) p. 318, pl. xxviii.

1026 MR. G. A. BOULENGER ON

The specimen in Dr. Spurrell’s collection measures 55 nim, from

snout to vent. It was shining jet-black in life, with the belly

grey. The thick skin of the ventral region forms a strong fold

on each side, which extends across the pubic region.

16. Rana PALMIPES Spix.

A single female specimen, measuring 95 mm, from snout

to vent.

Tympanum three-fourths the diameter of the eye. First

finger extending well beyond second. Skin of back finely

granulate; upper surface of tibia with longitudinal ridges

formed by confluent granules. Tibio-tarsal articulation reaching

half-way between the eye and the tip of the snout. Back pale

olive-grey (appears to have been green in life), without spots ;

belly uniform white.

IT have examined smaller specimens (40 to 70 mm.) from the

same locality (Condoto) from Mr. M. G. Palmer’s collection. In

the smallest the tympanum measures barely two-thirds the

diameter of the eye, the upper parts are blackish brown and

the lower parts profusely spotted with black.

17. PHYLLOBATES PRATTI Blegr.

A single specimen, 15 mm. long from snout to vent.

This species was described in 1899 (Ann. & Mag. N. H. [7] in.

p. 274, pl. x1. fig. 3) from two specimens obtained by Mr. A. E.

Pratt at Santa Ines, north of Medellin, at an altitude of 3800 feet.

Other specimens have since been found in the Choco, at Novita,

altitude 150 feet, by Mr.M.G. Palmer. In Dr. Spurrell’s specimen,

the white line in front of the thigh is produced forward to the

upper eyelid, dividing obliquely the black lateral band.

18. DENDROBATES TINCTORIUS Schneid. (PI. CIV. figs. 2-6.)

The extraordinary colour variations to which this species is

subjected are only paralleled by those of the African frogs of the

genus Rappia, and afford the same ditticulties to the systematist.

Some of these variations are local, others are merely individual ;

great is the perplexity in having to decide which deserve to be

recognised by a special varietal name. In the collection made by

Dr. Spurrell the specimens fall under two principal categories which

1 will designate as vars. coctei and chocoensis respectively. It is

not an easy matter to fix the typical form of D. tinctorius; the

following remarks will not be superfluous, té justify the course

followed by me.

Schneider (Hist. Amph.1i. p. 175, 1799) is the author of the

name tinctorius, but the frog itself was unknown to him. He

refers us to Lacepéde (Quady, ONnalos joy DO, jal xxoabe,. 17s),

who regards the “‘ Raine a tapirer” as a variety of Hyla rubra

Laurenti, and describes it as reddish with two irr egular yellowish

white longitudinal bands; the figure is hopeless for the purpose

COLOMBIAN BATRACHIANS AND REPTILES. 1027

‘of identification. But as the name ‘“ Raine a tapirer”’ is based

on an observation recorded by Buffon (Hist. Nat. Oiseaux, vi.

p. 235, 1779) to the effect that the frog is used by the savages of

Guiana and the Amazon to dye parrots, and is small, azure blue

with longitudinal golden yellow bands, and is to be found pre-

served in the ‘‘Cabinet du Roi,” the best thing to do is to go

straight to the description of the first author after Lacepede who

had access to the specimens alluded to by Buffon, and this author

is Daudin (Hist. Rain. etc., 1802). The larger of the three speci-

mens from Guianain the Paris Museum, presumably those alluded

to by Buffon and Lacepéde, is figured in a perfectly recognisable

manner on Daudin’s pl. vii. fig. 1, and corresponds very nearly

with one from Cayenne in the British Museum, which I therefore

regard as the typical form of Dendrobates tinctorius. Upper parts

blackish brown with symmetrical white (yellow) markings forming

two bands on the back, meeting on the coccygeal region and ex-

tending, above the canthus rostralis, to the end of the snout after

expanding on the upper eyelid; a transverse bar connects these

bands across the middle of the back, and another branch extends

to the shoulder; white markings on the flanks.; belly paler brown,

with black spots. This is the var. daudini Steindachner, Verh.

zool.-bot. Ges. Wien, xiv. 1864, p. 262.

Var. coctei.__Hylaplésie de Cocteau, Dum. & Bibr. Erp. Gén.

pl. xe. fig. 1; Dendrobates tinctorius, var. B, Dum. & Bibr. op. cit.

vill. p. 654 (1841); D. tinctorius, var. cocteaui Steind. Verh.

z001.-bot. Ges. Wien, XIV. 1864, p. 260.

“ D’un brun marron ou dune teinte lie de vin, .... une tache

dun blane jaunatre sur Je museau, une autre beaucoup plus grande

et de forme ovalaire sur chaque flane, .... un large bracelet de

la méme couleur autour de chaque bras et, de chaque jambe.

Assez souvent les taches des flanes se confondent sur la région

abdominale.” Dum. & Bibr. Locality not stated.

Six specimens, all from the Choco, in the British Museum, are

referable to this variety :—

Tado, Rio San Juan, 230 ft. (M. G. Palmer). Dark reddish

brown above, with black spots, black beneath. An oval yellow

spot on the top of the head, another occupying the whole flank,

widely separated from its fellow on the other side; other large

spots are present, one on the forearm, one on the thigh, one on

the tibia, and one on the throat.

b. Same locality, same collector. Similar to the preceding, but

the spots on the head, throat, and femur absent; the large spot

on the flank extends to. the shoulder, and those on the forearm

and tibia completely encircle the limbs.

c. Upper waters of Condoto River, 1200 ft. (Spurrell). Dark

brown above, with bright vermilion- red, sharply defined spots,

black beneath. The br ight spots are as in a, except that the one

on the top of the head is situated further back; those on the

throat and forearm. are absent, but others are present on the left

1028 MR. G. A. BOULENGER ON

side of the snout and on the left foot ; the forearm is completely

encircled by the yellow spot.

d. Same locality asc. Differs from ec in the lateral spot being

produced as far as the eye on the left side, where it is confluent

with the oval spot situated on the upper surface of the head as

in @; the spot on the tibia surrounds the limb; a yellow spot on

the right foot instead of the left.

e, f. Same locality as cand d, and very similar in the markings,

which, however, were noted by Dr. Spurrell as of a vivid

yellow instead of red. e has the lateral spots very large and

narrowly separated from each other on the belly, that on the

left side extending to the temple; a spot on the right foot; fore-

arm encircled by the red spot. jf has the spot on the right foot

instead of the left.

A further Colombian specimen, from Noanama, 8. Juan

(M. G. Palmer), is not unlike the preceding, but the spots are

smaller and less regular and of a vivid magenta-red. A figure is

given to show the disposition of the markings on the black body.

Var. chocoensis. — Phyllobates chocoensis Posada Arango,

Memoire sur le poison de Rainette des sauvages du Choco (Paris,

1869); Deyrolle, Rev. et Mag. de Zool. 1872, p. 465, pl. xxvii.

The frog noticed by Posada Arango as furnishing the Choco

Indians with a deadly poison for their arrows is described as

bright yellow on the head and back and bluish black on the limbs

and lower parts. A specimen answering to this description is

preserved in the British Museum, from Juntas, Rio San Juan,

Choco, 320 ft. (M. G. Palmer).

Nine specimens, obtained by Dr. Spurrell at Petia Lisa,

Condoto, 300 feet, may be referred to this form, and show

considerable variation in coloration. A snake which started

eating one of these frogs was observed to drop it hurriedly.

‘Ground colour of upper parts a rich orange-yellow, belly dark

brown ; markings on back and limbs varying from dark reddish

orange to chocolate-brown, not sharply defined nor regularly

symmetrical ; much variation in colour and arrangement of

markings; ground colour of upper parts sometimes lemon-yellow

or brownish yellow.” ‘The specimens which most nearly approach

Posada Arango’s description have the bead and back uniform

yellow or with only one dark spot, on the middle of the back,

just behind the head; the sides are blackish brown, and the upper

surface of the limbs yellow.

The numerous varieties of Dendrobates tinctorius are much in

want of revision. Among those already described there is one

which is unquestionably entitled to specific rank and for which I

wish to propose the name Dendrobates paraensis. Disks of fingers

and toes much larger than in D. tinctorius; a small but very

distinct tubercle on the inner side of the tarsus, nearer the meta-

tarsal tubercles than the tibio-tarsal articulation. In some

COLOMBIAN BATRACHIANS AND REPTILES, 1029

specimens the upper parts, with the exception of fore limbs and

feet, are uniform greyish or yellowish white (yellow in life ?), the

rest of the animal deep black; in others the black predominates,

the white being restricted to the upper surface of the head and to

Text-fig. 178.

n B

Dendrobates tinctorius (A) and D. paraensis (B).

Upper view of hand.

more or less confluent blotches on the back ; further specimens

are intermediate between the two extremes. From snout to vent

38 mm. Hight specimens, from Para, are preserved in the

British Museum, three of which are mentioned in the Catalogue

of Batrachians, p. 143 (var. B, h, 7, £).

19, DENDROBATES AUROTANIA, sp. n. (PI. CIV. fig..1.)

Snout truncate, barely as long as the eye; loreal region

vertical; interorbital space as broad as the upper eyelid;

tympanum not very distinct, three-fifths the diameter of the eye.

Fingers slender, first extending beyond second, with rather small

disks; toes slender, disks as large as those of the fingers; sub-

articular tubercles very feeble; two scarcely prominent meta-

tarsal tubercles and a very feeble fold on the inner side of the

tarsus terminating club-shaped half-way between the metatarsal

tubercles and the tibio-tarsal articulation. The tarso-metatarsal

articulation reaches far beyond the tip of the snout, the tibio-

tarsal falling between the eye and the end of the snout; tibia

half as long as head and body. Skin smooth, of dorsal region

very porous. ‘“ Jet-black, with a metallic orange-yellow streak ”

from the end of the snout along the canthus rostralis and the

upper eyelid to the loin; ‘‘a few fine gold freckles on thighs and

arms.”

From snout to vent 30 mm.

A single specimen from Pefia Lisa, Condoto, 500 feet.

This frog approaches very closely D. trivittatus Spix, differing

only in the shorter snout and the absence of tubercles on the

back.

1030 MR. G. A. BOULENGER ON

REPTILIA,.

CHELONIA.

1. CINOSTERNUM SPURRELLI, sp.n. (Pls. CV. & CVI.)

Jaws strong; beak strongly hooked. Carapace smooth and

without keels, flattened on the vertebral region, profile descending

very abruptly behind; deep grooves between the shields;

posterior border serrated. Plastron smaller than the opening of

the shell, not emarginate posteriorly; lobes well movable, front

one longer than the fixed portion and slightly shorter than the

hind one; gular shield not half the length of the front lobe;

suture between the pectoral shields much shorter than that be-

tween the humerals; axillary and inguinal shields elongate and in

contact with each other. A patch of small horny keeled tubercles

on the back of the leg, opposed to one on the thigh. Tail ending

in a claw-like scute. Carapace blackish brown, plastron yellow

with blackish blotches; head, neck, and limbs blackish above,

whitish below; a broad oblique yellowish band on each side of

the head, from behind the eye to the neck, passing above the

. tympanum; jaws yellowish horn-colour.

This very distinct species is represented by a single male

specimen, the shell of which measures 115 mm. The curious

shape of the first vertebral shield, depicted on Pl. CV., is probably

an individual peculiarity and has therefore not been alluded to in

the description.

The position of C. spwrrelli in the system is in group II. A of

my Synopsis (Cat. Chelon. p. 38), which group, up to the present,

was unrepresented south of Mexico.

2. Nicorza wAsura Bler.

Described in 1902 from specimens obtained at Bulun and on

the Rio Durango, N.W. Keuador.

The specimen in the present collection is quite young. Shell-

length 70 mm. ‘The digits are fully webbed.

EMYDOSAURLIA.

3. CAIMAN SCLEROPS Schn.

LACERTILIA.

4. ANOLIS FASCIATUS Bley.

Only known from Guayaquil and. N.W. Eeuador (A. elegans

Blgr.) ‘*Green above, with darker green markings, yellow on

belly; crown of head and neck with faint orange markings.

Gular appendage white with six broad orange bands.”

5. ANOLIS MACULIVENTRIS Bler.

First described from N.W. Ecuador.

“ Gular appendage crimson.”

COLOMBIAN BATRACHIANS AND REPTILES. 1031

6, ANOLIS PRINCEPS Bler.

This handsome lizard, one of the largest of the genus, was

originally described from specimens from N.W. Heuador. It has

since been found in Choco, at Condoto and T'adé, by Mr. M..G.

Palmer.

“Yellowish grass-green, brighter about head and neck, with

yellow blotches round eyes; diagonal markings brownish. black ;

gular appendage white.”

7. ANOLIS BREVICEPS, sp. n. (Pl. CVII. fig. 1.)

Head once and two-thirds as long as broad, slightly shorter

than the tibia; snout obtuse; forehead and interorbital region

concave; frontal ridges short and divergent ; upper head-scales

very small, keeled; scales of the supraorbital semicircles enlarged,

separated by four or five series of scales; enlarged, strongly

keeled supraocular scales ; occipital as large as or larger than the

ear-opening, separated from the supraorbital by three or four

series of scales; canthus rostralis sharp, canthal scales three or

four ; loreal rows seven or eight; seven upper labials to below

the centre of the eye; ear-opening moderate, vertically oval.

Gular appendage very small in males, barely indicated in females ;

gular scales keeled. Body not compressed ; no dorso-nuchal fold.

Dorsal seales very small, keeled, gradually merging into the

minute granules of the sides; ventrals larger, roundish-hexagonal,

keeled. Limbs slender; the adpressed hind limb reaches the

tip of the snout, or between the eye and the tip of the snout ;

digits feebly dilated; 13 to 15 lamelle under phalanges IT and

IIT of the fourth toe. Tail cylindrical, once and a half to once

and two-thirds as long as head and body, without vertebral series

of enlarged scales. Male with enlarged postanal scales. Grey-

brown to dark brown above, with a ventral series of blackish

dots or small V-shaped markings; a dark brown, black-edged

cross-band between the eyes and another band from the end of

the snout, through the eye, to the side of the neck, very broad

and sharply defined on the temple; a more or less distinct

whitish streak may be present below this band.

Sh anes 2

Hbovellilemethiee anne nece: 108 124 mm.

ASS clapwetn tars lte ons lee datasets Ne 13

Wardithwot headless e Sein )

'EBOdy) iia. sete eeieiae seeps BySO 30

Thome lin) soa.5ccedosbeccees 21 21

dla Glielimmllo eye eee. ses 40, 40

TPilotans abo. setae: assis ede 18} 133

Sans REAR MNEs AIR 66 76

Three specimens.

Near the Central American A. godmant Blgr. Distinguished

by the shorter head, the smaller gular appendage, and the shorter

tail. =

1032 MR. G. A. BOULENGER ON

8. ANOLIS NOTOPHOLIS Blgr.

The gular appendage is red in both sexes.

9. ANOLIS MACROLEPIS Blgr.

10. Potycurus currurosus Berthold.

Two specimens, male and female.

Male green above and on the belly, cream-colour on the sides ;

a vertebral series of cordiform pale reddish-brown black-edged

spots pointing forwards, connected by a bright orange vertebral

band. Female reddish brown above and beneath, with angular

dark brown bands across the back; a few dull green blotches on

the head, and on the sacral and anal regions.

11. CoRYTHOPHANES CRISTATUS Gray.

This remarkable lizard was believed to be restricted to Central

America (specimens from British Honduras, Guatemala, Nicaragua,

and Costa Rica are preserved in the British Museum), but a

specimen from the Rio Tamana, Choco, was in the collection

made by Mr. Palmer, and two are in Dr. Spurrell’s collection.

“Shows rapid changes of colour when alive.”

12. BASsILIscUS AMERICANUS Laur.

Its habitat extends from Panama and Costa Rica to Venezuela

and Southern Colombia.

13. Bastniscus GALERITUS A. Dum.

Blackish above, dotted with yellow ; upper surface of head and

a band on each side of the belly reddish brown; throat white ;

belly and lower surface of limbs greenish white, speckled with

black.

14. ENYALIOIDES HETEROLEPIS Bocourt.

Originally described from Veragua. Numerous examples from

Colombia and N.W. Ecuador are preserved in the British

Museum.

15, DipLocLossus MonorRoPIs Wiegm.

The largest specimens are very strikingly coloured. Head

lemon-yellow with greenish tinge; back chequered brownish

black and lemon-green; throat, belly, and sides of body ver-

milion-red ; tail lemon-yellow above, with dark bars, vermilion-

red beneath.

16. AMEIVA BRIDGESII Cope.

Young black above, with seven whitish longitudinal lines ;

belly bluish grey ; tail blue.

COLOMBIAN BATRACHIANS AND REPTILES. 1033

17. Ametva Festiva Licht.

A single, half-grown specimen. Blackish above, with a pale

green vertebral band originating on the end of the snout and

prolonged on the basal half of the tail; two interrupted white

lines along each side of the body.

A specimen from Medellin, Andes of Colombia (Coll. A. KE.

Pratt), is in the British Museum.

18. AnwapiA viTTATA, sp.n. (Pl. CVII. fig. 2.)

Head depressed, with long, acutely pointed snout ; body much

elongated. Nostril in the suture between two nasals. Fronto-

nasal pentagonal, a little longer than broad, longer than the

prefrontals, which form a long suture; frontal octagonal,

narrow behind, in contact with three supraoculars; two large

parietals on each side, both in contact with the interparietal,

which is not twice as long as broad and followed by three sub-

equal occipitals; four supraoculars, first smallest, second and

third equal; four or five supraciliaries; a loreal, a freno-orbital,

and a row of infraorbitals; temples with rather small shields ;

seven upper labials, first narrowly in contact with the fronto-

nasal; six lower labials; chin-shields, one anterior and four

pairs, three pairs forming a median suture; gular scales sub-

equal, squarish, juxtaposed ; collar-scales nine, feebly enlarged.

Scales on body forming regular whorls, the dorsals quadrangular-

subhexagonal, as long as broad, the laterals a little narrower, the

ventrals quadrangular, those of the two median rows larger; 30

scales round the middle of the body, 58 from occiput to sacrum

inclusively. Preanal plates in three rows, posterior median pair

largest. Two przanal and one femoral pore on each side. Limbs

and tail as in A. ocellata. Pale brownish white above, with three

dark brown longitudinal stripes, the vertebral edged with black

and breaking up into three black lines behind the nape; lower

parts white; tail with three serrated or zigzag brown bands.

Total length (tail injured) .................. 112 mm.

JELCE Vel GM eenie te rote none Sun Ae ed ent ee Bee 10

Wel Glitlnrotmne adnate ercrtrcksS eacey isola ons 5

From end of snout to fore limb ............ 15

MMR ae Sean VECINGL craiancaiet hes cis wilt a 40

Ronen imilbpessccc ee eee ate es see 10

Tina lelinrmllneerieete set sere iate ince cece ees aces i,

A single specimen.

This species is closely allied to A. ocellata Gray.

19. ALopocLossus copit Blgr.

Originally described from Hastern Ecuador, this species has

since been found in Western Hcuador, but had not been reported

from Colombia.

1034 MR. G. A. BOULENGER ON

20. AMPHISBENA FULIGINOSA L.

Received alive by the Zoological Society.

OPHIDIA.

21. TRACHYBOA BOULENGERI Peracca.

A single young specimen of this species, recently described

by Peracca, Ann. Mus. Zool. Univ. Nap. iu. 1910, No. 12, was

referred by me to 7. gularis Peters, P. Z.S. 1898, p. 109,

pointing out the characters in which it differed from Peters’s

description and Jan’s figure. The spine-like raised scales on

the snout and above the eyes, which are so striking in larger

specimens, are merely indicated in this specimen.

The British Museum possesses two adult specimens from N.W.

Keuador, and two from the Choco are now added by Dr. Spurrell.

1 append particulars of these four specimens :—

1. 2, St. Javier, N.W. Keuador. Total length 430 mm.;

the eye on the right side.

2. 3, with claw-like anal spurs. Same locality. Total length

205 mime wall SOs Sc 30) View sor Co 20.0 atiiuhalaloiall

entering the eye.

3. &¢, with claw-like anal spurs. Petia Lisa, Choco. Total

length 230 mm.; tail 27. Sce.30; V.131; C.23. Labials

all excluded from the eye.

4. 2, same locality. Total length 270 mm.; tail 25. Se. 33;

V. 137; C. 20.- Labials all excluded from the eye.

_ The type specimen described by Peracca, the habitat of which

is unknown, has only 23 scales round the thickest part of the

body. The number of scales in 7’rachyboa is probably as variable

as it is in Atheris.

22. STREPTOPHORUS ATRATUS Hallow.

A specimen taken from the stomach of an Hlaps rosenbergii,

after having swallowed a Cecilia intermedia.

23. DRYMOBIUS DENDROPHIS Schleg.

24, PHRYNONAX FASCIATUS Peters.

25. HmRPETODRYAS CARINATUS L.

26. LEPTOPHIS BILINEATUS Gthr.

One of the three specimens is bright green above, as in the

type, the two others are of a metallic bronze on the back and

sides, the head dark green, the sides of the neck green, the lower

parts pale green.

The habitat of this species extends from the Isthmus of

Tehuantepec to N.W. Ecuador.

COLOMBIAN BATRACHIANS AND REPTILES. 1035

27. LEPTOPHIS OCCIDENTALIS Gthr.

28. LioPHIS ALBIVENTRIS Jan.

29. ATRACTUS MULTICINCTUS Jan.

In its coloration in the fresh condition (occipital blotch bright

red) this harmless snake is not without resemblance to Llaps

mipartitus.

30. ATRACTUS IRIDESCENS Peracca. :

This species was described (Boll. Mus. Torin. xi. 1896, No. 252)

from a single specimen, origin unknown. Three specimens in

Dr. Spurrell’ s collection may, I think, be referred to it.

G QD, Bs mnwi,e wall BG, We 144s Ces Bre

Oo, QQ imi, 2 wan BA, Wo s¥bs O, Beh

Wore a0 2 mia tavlelle Ven See 20:

The iridescence from which the species derives its name is

strongly marked in these specimens. Dark brown above, with two

dorsal series of transverse blackish spots, which may alternate or

form cross-bars, and may be bordered by yellow dots or short

streaks; in specimen @ the belly is white spotted with black in

front, uniform black behind; in specimen 6 it is white with a

median series of black spots and a series of smaller spots ‘on

each side.

31. PETALOGNATHUS NEBULATA IL.

32. HIMANTODES CENCHOA L.

33. LePTopIRA ALBOFUSCA Lacep.

34. OXYRHOPUS CL@LIA Daud.

A specimen two metres long.

35. HoMALOCRANIUM MELANOCEPHALUM IL.

36. HOMALOCRANIUM CORALLIVENTRE, sp. n. (Pl. CVIILI. fig. 1.)

Kye about half as long as the snout. Rostral a little broader

than deep, the portion visible from above about half as long as its

distance from the frontal; internasals about two thirds the length

of the prefrontals; frontal hexagonal, obtuse-angled in front,

acute-angled behind, nearly once and a half as long as broad,

longer than its distance from the end of the snout, shorter than

the parietals ; nostril between two nasals, the posterior of which

is in contact with the preocular; two postoculars; temporals

1+ 1; seven upper labials, third and fourth entering the eye,

seventh largest; four lower labials in contact with the anterior

chin-shields, which are a little shorter or a little longer than the

1036 MR. G. A. BOULENGER ON

posterior and in contact with the symphysial. Scales in 15 rows.

Ventrals 135-141; anal divided; subcaudals 56-59. Dark brown

above, scales edged with black; a large yellowish spot on the

upper lip behind the eye, and a yellowish occipital bar encroaching

on the posterior part of the parietal shields; gular region white,

rest of lower surfaces bright coral-red.

Total length 275 mm.

Two specimens, male (V. 135; C. 59) and female (V. 141;

C. 56).

Very closely allied to H. alticola Blgr., from Santa Rita, north

of Medellin, Colombia (9000 feet), but eye larger and frontal a

little longer in proportion to its width.

37. STENORHINA DEGENHARDTI Berthold.

38. ELAPS CORALLINUS L.

39. ELAPS ROSENBERGII Blegr.

J have examined several specimens from various localities in

N.W. Ecuador, and also one obtained by Mr. M. G. Palmer at

Tado, Choco.

40. Evaps mrcrops, sp. n. (Pl. CVIII. fig. 2.)

Kye extremely minute, barely one-fourth as long as its distance

from the mouth. Rostral broader than deep; frontal a little

broader than the supraocular, once and one-third as long as

broad, as Jong as the prefrontals, half as long as the parietals,

which are longer than their distance from the internasals; one

pre- and two postoculars ; temporals 1-+-1; seven upper labials,

third much longer than fourth, third and fourth entering the eye,

the latter very narrowly; four lower labials in contact with the

anterior chin-shields, which are as long as the posterior. Scales

in 15 rows. Ventrals 205; anal divided; subeaudals 20. Tail

ending obtusely. Black above, with 46 narrow white, black-

spotted cross-bars; these widen on the belly, which is barred

black and white; an orange band across the head, from between

the eyes to near the end of the parietal shields; tail orange-red,

with a black ring and a black terminal spot surrounding a small

white spot.

Total length 650 mm. ; tail 25.

A single female specimen.

This new Hlaps resembles strikingly #. mipartitus D. & B. in

the coloration, but is well distinguished by the much smaller eye

and by the proportions of the head-shields.

41, LeproGNATHUS SPURRELLI, sp.n. (PI. CVIII. fig. 3.)

Body very slender, strongly compressed. Hye large. Rostral

small, about once and a half as broad as deep, not visible from

above ; internasals one-third the length of the prefrontals ;

COLOMBIAN BATRACHTANS AND REPTILES. 1037

frontal as long as broad, nearly as long as its distance from the

end of the snout, much shorter than the parietals; nasal divided ;

loreal once and a half as long as deep, bordering the eye; no

preocular ; prefrontal entering the eye; two postoculars, lower

very narrow ; temporals 3 + 3; eight upper labials, fourth and

fifth entering the eye; four pairs of chin-shields, the anterior

very small and in contact with the symphysial, the second larger

and a little longer than broad. Scales in 15 rows, vertebrals

strongly enlarged, nearly as long as broad. Ventrals 208; anal

entire; subcaudals 132. Reddish brown above, with very large

blackish-brown blotches separated on the back by narrow inter-

spaces of the ground colour, these interspaces gradually widening

down the sides, where the blackish blotches are edged by small

white spots: these blotches form nearly complete rings on the

anterior part of the body, whilst further back they are gradually

more widely interrupted by the yellowish-white colour of the

belly ; lower surface of tail brown, with a few small whitish

spots; head uniform reddish brown above; lips white, with

small black spots. ‘‘ Iris golden.”

Total length 680 mm, ; tail 235.

A single male specimen.

Closely allied to Z. annulata Gthr., from Costa Rica, and

LL. temporalis Werner, from Ecuador. ;

42. LAcHuEsIS mutus L.

This is the othrops acrochordus of Garcia*. Grows to

24 metres.

43. LacuEsis atrox L.

44, LACHESIS MONTICELLII Peracca.

This Pit-Viper, easily distinguished from ZL. lanceolatus by

the shape of the rostral shield, which is much deeper than broad,

was described along with Zrachyboa boulengeri from a specimen

of unknown origin. It is interesting to find the two snakes

associated in this collection.

Two female specimens :—

a= 9A0hmmes tailelGOr seu29)-) Vie 200. 86.

bs SOO amties 4) MOO; 4, 0 a IAS eile

Lachesis punctatus Garcia, l.c. p. 31, pl. —, perhaps represents

the same species.

45. LACHESIS BRACHYSTOMA Cope.

The range of this species is now known to extend from the

Isthmus of Tehuantepec to N.W. Ecuador.

Described and figured by Garcia as Thanatophis sutus.

* Los Ofidios venenosos del Cauca (Cali, 1896), p. 23, pl. —.

Proc. Zoot. Soc.—1913, No. LXIX. 69

1038

ON COLOMBIAN BATRACHIANS AND REPTILES.

EXPLANATION OF THE PLATES.

Prater CII.

Fig. 1. Bufo hypomelas, p. 1022, enlarged, and outline natural size.

Fig.

2. Agalychnis calcarifer, p. 1023.

2a. A % Side view.

2b. Be es Open mouth.

Puate CITI.

Agalychnis sgurrelli, p. 1024. Female and eggs.

Prater CIV.

. 1. Dendrobates aurotenia, p. 1029.

2-4, 3 tinctorius, var. coctei, p. 1027.

5-6. 5 5 var. chocoensis, p. 1028.

Prats CV.

Cinosternum spurrelli, p. 1030. Upper view.

Prate CVI.

Cinosternwm spurrelli, p. 1030. Lower view.

Prate CVILI.

. 1. Anolis breviceps, p. 1031.

Wes “op % Side view of head.

Dae) as 5 Upper view of head, enlarged.

2. Anadia vittata, p. 1033.

Hols” 5 a Upper view of head, enlarged.

PA al ey - Side view of head, Pa

Poh | cn Hf Ventral and anal region, enlarged.

Bit cs 5 Lower view of head, 53

Pruate CVIII.

1. Homatlocranium coralliventre, p. 1035.

la. is BS Upper view of head, enlarged.

1b. Side view of head, 43

2. Hlaps micr ops, Dp. 1036. :

BGs gp . Upper view of head, enlarged.

2b. Side view of head, Ss

3. Leptognathus spurrelli, p. 1036.

3a. y, Bs Upper view of head, enlarged.

3b. a a Side view of head, 3

3c. is 3 Lower view of head, a}

ON THE PERONEAL MUSCLES IN BIRDS. 1039

60. The Peroneal Muscles in Birds. By P. CHALMERS

Mircpety, M.A.,D.Sce., LL.D., F.R.S.,F.Z.8., Secretary

to the Society.

[Received October 28, 1913: Read November 25, 1913.]

(Text-figures 179-190.)

INDEX.

STRUCTURE. Page

Myotoey.

Peroneal Muscles in Chauna ........- ‘ haere nate LOAD:

Systematic Description of Peroneal Muscles in Wire caeheiaesets 1043

JNTIOLOGY.

Distribution of Varieties of Peroneal Muscles in the

System .. M4 Sac qune eat ea dae Foti adnate. LOGS

Adaptation andl Deseon) Sho tunssebnareacmensbncstasesctdcesaaeten LOMO)

SysTEMATIC.

RAN AUO Na Cress 5 OAR eee LAE ee EN LOS OSMO72

(Obyayenrihel (eal: Cannaaceratecnsoce cdc osmGaaodand sted ecoemeadtodsauace ya UO a0}

In the dissection of an example of Baleniceps rex, which formed

the basis of a recent contribution to the Society’s Proceedings

(supra, pp. 644-703), I was interested to notice that one of the

very few characters in muscular anatomy which the Shoe-bill

shared with Herons, to the exclusion of Storks, was the presence

of the leg muscle known as the Peroneus brevis or profundus. On

looking up the literature relating to the peroneal muscles in birds,

I found that even Dr. Gadow’s careful descr iption of these muscles

(Gadow in Bronn’s ‘ Thier-Reich,’ Aves, pp. 180-182) was based

on a relatively small number of individuals and types, and that

his work, together with the insignificant contributions of earlier

and later writers, supplied only a vague and dubious picture of

the anatomical facts in Aves. I wished to make a survey of

these muscles as nearly as possible complete. Accordingly I have

dissected them in the birds that have died in the Society’s Collec-

tion for some months, have made use of the rich spirit material

stored in the Prosectorium (most of it dating from the time of

Garrod and Forbes), and have been allowed to examine some

specimens in the stores of the British Museum (Natural History)

and of the Museum of the Royal College of Surgeons, and

Dr. Gadow has also obliged me with material from Cambridge.

My observations therefore relate to several hundred birds, dis-

tributed, as will be seen later, fairly well over the whole Avian

system. Theoretically, no doubt, it is necessary to work out the

complete anatomical structure of an animal if one would under-

stand any portion of it, but in practice this is impossible, and in

my opinion the examination of a single structure or even portion

of a structure through the whole series by a single observer can-

not fail to be interesting and instructive. The method, moreover,

69*

1040 DR. P. CHALMERS MITCHELL ON THE

has the practical advantage that the anatomist gains, at least

temporarily, a useful dexterity in exposing the structures on

which he is engaged and is able to cover much ground in a

relatively short space of time.

After giving an account of the muscles in a generalized type, I

shali proceed to a systematic description of the conditions presented,

following the classification given by Dr. Gadow in the article

Birp, in the Eleventh Edition of the Encyclopedia Britannica,

and finally shall discuss some of the problems suggested by the

degree of coincidence between the varying conditions of these

muscles and a classification which is based on other anatomical

facts.

The Peroneal Muscles in CHAUNA CHAV ARIA (text-fig. 179).

In some respects the Screamers are generalized birds with

affinities pointing in many directions, and it happens that the

condition of the peroneal muscles in other birds can be easily

described as specializations of the Screamer condition chiefly by

loss of parts. :

Peroneus superficialis seu longus.—TVhis is the most superficial

mass of muscle seen when the skin is removed on the outer, upper

aspect of the leg from the knee downwards. Its very large fleshy

mass arises from the anterior and lateral crista of the tibia, from

the head of the fibula and partly from the fascia covering the

knee. Laterally, where it is bounded by the external head of the

gastrocnemius on the one side, and the tibialis anticus on the

other, its fasciz are closely attached to the fasciz of these muscles.

The muscular mass is roughly triangular, and from the broad

base at its origin from the upper end of the tibia and fibula,

it converges to a flat tendon which is apparent about three

quarters way down the leg and rather more on the outer than the

anterior aspect. There is a deep origin formed by a comb of

fibres projecting inwardly as a keel along nearly the whole length

of the triangular portion of the muscle and arising partly from

the whole length of the fibula and partly from the tibial surface

immediately under the fibula. When the tendon has nearly

reached the tarsal joint, it gives off a very broad and strong anchor

(text-fig. 179, Anch.) to the cartilaginous sustentaculum which is

pierced by some of the flexor tendons and joined superficially by

the gastrocnemius tendon. It is then continued outwards and

downwards across the ankle-joint, immediately under the skin,

but running through a synovial membrane which has to be cut

open to expose it fully, and ultimately, about a quarter of the

distance down the tarso-metatarsal shaft, it fuses with the flexor

tendon of the third digit (flexor primus seu perforatus IIT) (text-

fig. 179, P.L. IIL), It is supplied by a branch of the ischiadic

nerve.

The function of this muscle is to straighten the tarsus meta-

tarsus on the tibio-tarsus and then to flex the third toe. When

PERONEAL MUSCLES IN BIRDS. 1041

Text-fig. 179.

Peroneal muscles in Chauna chavaria.

Right leg, external aspect. Muscle striped: tendon dotted.

P.LON. Peroneus longus. ANCH. “Anchor”: attachment of P. longus tendon

to Sustentaculum of Flexor tendons. P.L. III. Slip of P. longus tendon passing’

across the ankle-joint to fuse with III, tendon of Flexor perforatus of the third

toe.

P.BRE,. Peroneus brevis. P.B.1. Tendon of insertion of P. brevis passing across the

tarsal jot to insertion to proximal end of the tarsus-metatarsus. P.B.2. Slip

of Peroneus brevis tendon, not usually present in birds.

Tar. Lig. Ligament crossing the tarsal joint, deep of P.L.III and superficial to

JEIB5 Ils

Gastr. External head of gastrocnemius muscle.

Flex. Flexor tendons.

1042 DR. P. CHALMERS MITCHELL ON -THE

the tendon has been dissected out of its synovial groove, it

appears too long for its purpose, but if its action be tested before

this dissection has been made, it is clear that the straightening of

the foot is carried out by the action on the sustentaculum and

that subsequently the third toe is bent by the special slip.

The arrangement is odd and very striking, and it is extremely

difficult to believe that it has been independently developed in the

large number of different groups of birds in which it occurs. In

my opinion the disposition of the muscle and its tendons shown

in Chauna can be explained best if it be regarded as the spe-

cialized derivative of a condition in which the perforated flexor

muscles and their tendons were continuous with each other and

with the peroneus ; as the foot and its digits became specialized,

the fasciz of insertion became strengthened along the lines

leading to each digit, and finally by the disappearance of the

connecting fascize the continuous sheet became broken up into

separate tendons ; as the tibio-tarsus became longer the peroneal

slip to the tendon of digit III assumed its odd course. All the

stages in the possible differentiation of continuous fascie into

highly specialized tendons still exist in the case of the alar

tendons of birds. The differentiation of the foot, however, must

have taken place before the groups of birds became separated,

and, as I shall show later, the various conditions of the peroneus

in the different groups of birds can all be explained as differ-

entiations of the condition in Chauna, in particular by the loss

of some part.

Peroneus profundus seu brevis (Tibialis posticus of R. W. Shu-

feldt, “‘ Myology of the Raven,” 1870, p. 228).—This is a much

simpler and more slender muscle arising fleshy from the fibula

and the adjacent surface of the tibia on their anterior aspects,

beginning about the level of the insertion of the biceps tendon

and extending a little beyond the distal extremity of the fibula,

where it passes into a stout rounded tendon (text-fig. 179, P- BRE).

The greater portion of the muscle is concealed by the edges of the

tibialis anticus and of the superficial peroneal which meet above

it. The tendon as it approaches the end of the tibio-tarsus is

very close to the bone and is deep of the peroneus longus tendons ;

it flattens out, runs through a synovial channel, crosses the tarsal

joint overlain by a ligament (text-fig, 179, Tar. lig.), which also

crosses the joint, and is inserted to the proximal edge of the tibio-

tarsus on its posterior, outer aspect (text-fig. 179, PAB ail) same lol

Chawna it also has a second attachment to the antero-median end

of the tarsal shaft (text-fig. 179, P.B. 2).

This muscle flexes the tarsus-metatarsus on the tibio-tarsus, and

at the same time slightly rotates the foot, depressing the axial, or

great toe side, and raising the abaxial, or fourth toe side. The

latter action varies with the exact position of the insertion and

is often very feeble, as, indeed, is the action of the muscle as a

whole,

PERONEAL MUSCLES IN BIRDS. 1043

SysTEMATIC DESCRIPTION.

I have found that the following points are the most notable :—

Peroneus longus.—Origin : “superficial,” 7. e. forming a fairly

broad external sheet at the proximal end of the tibio-tarsal shaft

and arising chiefly from the head of the tibia and fibula and from

the fascize over the knee-joint; “fascial,” from the fascie of the

tibialis anticus on the one side and from that of the external

gastrocnemius and the underlying flexors on the other ; ‘‘ deep,”

from the side of the fibular shaft and the adjacent area of the

tibia on the opposite side from the origin of the peroneus brevis.

The “ fascial ” origin is the least important morphologically and is

correlated with the strength and width of the muscle. ‘ Anchor,”

to the cartilaginous sustentaculum through which the flexor

tendons run at the tarsal joint. ‘Slip to III,” the superficial

tendon which runs from the peroneus over the tarsal joint to join

the perforated flexor tendon of the third toe.

Peroneus profundus.—Origin. Presence or absence of a bony

or fibrous bridge at the lower end of the tibio-tarsus. Tendon of

insertion.

Relative size of the two muscles.

RATITAL,

Struthiones. Struthio.-—-The peroneus longus is enormous.

Superficial origin extends over to the femoro-fibular ligament and

the patella. Fascial origin present. Deep origin represented by

a separate head from the lower end of the fibula. Anchor is

broad and flat but short. Slip to III strong and rounded.

Peroneus brevis absent, but there is a short stout ligament which

may represent either the lower portion of the muscle, or, in my

opinion, more probably the ligament which crosses the tarsal

joint in Chauna.

Rhee. LKhea americana (two examples).—P. longus as in

Struthio, except that the deep origin extends for the ereater part

of the length of the fibula. P. Tire evis absent, but tar soil hgament

present,

Casuaril. Casuarius uniappendiculatus.—P. longus enor-

mous ; superficial and fascial origins very extensive, deep origin as

in Rhea. Anchor short, very wide and partly in slips. Slip to

III stout. P. brevis absent but there is a stout tarsal ligament.

Dromeus nove-hollandie.—As in Casuarius, but P. longus not

quite so large.

Apteryges. Apterym (@ species) (three examples).—P. longus

very large; superficial and fascial origins good, but deep origin

very slight. Anchor broad and short ; slip to III a rounded

tendon. P. brevis very much reduced but present as a small flat

tendon arising from a few muscular fibres and running in the

normal position under the slip to IIT. This is notably different

from the tarsal ligament in the other “struthious birds” and is

quite certainly a rudiment of the Peroneus brevis.

1044 DR. P. CHALMERS MITCHELL ON THE

In the Ratites the P. longus is the dominant muscle and does

not differ exceptin minor details from the type in Chawna which,

for convenience, I may call the “normal” arrangement. The

Peroneus brevis is at least functionally absent, but the Apteryges

stand apart from the others in possessing an undoubted vestige

of it.

CARINAT A.

CoLYMBOMORPH.

COLYMBIFORMES.

Colymbi. I have not examined a Diver.

Podicipedes. Podiceps cristata and P. minor.—In both these

Grebes the P. longus was badly developed, with delicate super-

ficial and deep origins passing into a long rounded tendon about

half-way between the knee and the tarsal joint. In the Crested

Grebe this tendon passed straight to the sustentaculum, so that

its insertion was represented only by a narrow anchor, there

being no slip to Il]. In the Little Grebe the anchor was similar

but I was able to trace a minute but normal slip to II]. On the

other hand, in the Crested Grebe there was a minute and certainly

functionless tendon representing the P. brevis which I did not

find in the Little Grebe. |

SPHENISCIFORMES.

Sphenisci. Spheniscus demersus.—P. longus rather weak

but all origins present; anchor and slip to III both slender and

short. P. brevis was also slender but arose fleshy from the

greater length of the shaft of the fibula and tibia and was

inserted by the usual slender flat tendon running in a synovial

groove under the slip to III. Gadow mentions that it is present

but weak in Aplenodytes. Beddard’s statement that 1t does not

occur in Penguins is erroneous. ‘

PROCELLARITFORMES.

Tubinares.—Daption capensis. Procellaria (?species). Ocea-

nodroma (¢ species).—In the Cape Pigeon and the two Petrels the

P. longus was rather small with only the superficial origin, which

quickly passed into a very long, slender tendon ending in a

delicate anchor and a slender slip to III. The P. brevis had a

short but strong origin from high up the fibula ; this passed into

a strong and very long tendon with the usual flat insertion. The

P. brevis is rather stronger than the P. longus. The examples

of Petrels which I examined were part of the material of W. A.

Forbes and had been preserved in spirit for at least thirty years.

I was interested to note that they still retained a very strong

musky odour, so clinging that although I used several kinds of

soap and petrol I could not get rid of it for several days.

PERONEAL MUSCLES IN BIRDS. 1045

Tt is clear that the Colymbomorphe present no coherent

picture, the Petrels being sharply marked off from the others

by the relative dominance of the P. brevis.

PELARGOMORPH®.

CICONIIFORMES.

Steganopodes.

Phaéthontide. Phaéthon.—P. longus normal but rather small,

with superficial and deep origin, broad anchor and slip to ILL.

P. brevis normal, with long muscular origin and short tendon with

usual flattened insertion.

Sulide. Sula bassana.—P. longus normal and large but with

only superficial and slight fascial origin. Broad short anchor and

ship to III. P. brevis absent.

Phalacrocoracide. Phalacrocorax (?species).—P. longus normal

and large; origin, anchor and slip to Ill as in Sula. P. brevis

strong but very short, arising only from the lower end of the

tibial shaft, distal to the fibula, and with normal insertion by

flattened tendon.

Fregatide. Mregata.—Peroneus longus large and strong but

origin practically only deep from the whole length of fibula and

adjacent surface of tibia on side turned from the brevis. Broad

anchor and good slip to III. P. brevis minute but quite distinct

and normal, with long slender belly and tendon passing under a

fibrous bridge and then flattening out to pass to usual insertion.

Pelecanide. Pelecanws.—I have not had a recent opportunity

of examining a Pelican, but from my old notes I find that the

P. longus is large and normal with the usual anchor and slip to

III, and that the P. brevis is present but minute.

Ardez (from these I exclude Scopus and Baleniceps).

Ardea egretta, A. herodias, A. herodias lessoni, A.purpurea. Ardetta

minuta. Nycticorax gardeni, NV. violaceus (2 examples). Botaurus

stellaris, Cancroma zelodoni.—In all these the P. longus is fairly

large (text-fig. 180), but has only the superficial origin with shght

fascial origin. The anchor is rather long, broadens out, and at

least in one case (A. herodias lesson) was intwoslips. The P. brevis

is well developed usually from about three-quarters the length of

the shaft but chiefly from the fibula. The long tendon passes

through a fibrous bridge and then flattens out to the normal inser-

tion. I think there must be some mistake in Gadow’s statement

that the P. brevis is absent in Ardea. I find that its presence is

one of the differences between Herons and Storks. In Ardetia

minuta and Cancroma it is particularly strong.

Scopide. Scopus umbretta.—The P. longus is large with

superficial and slight fascial origin, broad anchor and good slip

to III. The P. brevis is either actually absent or represented by

a very small ligament.

Balenicipitide. Baleniceps rexw.—P. longus a strong broad

1046 DR. P. CHALMERS MITCHELL ON THE

muscle with only superficial and fascial origins. Anchor broad

and good slip to III. The P. brevis is strong but short, arising

only from the tibia distal to the end of the fibula; its flattened

tendon passes over to the usual insertion.

Text-fig. 180.

(he r

/ Ni)

Peroneal muscles in Nycticoraa gardeni.

Description and lettering as in Text-figure 179 (p. 1041).

Br. Bridge traversed by the tendon of the P. brevis.

Ciconie, Ciconia nigra. Dissura episcopus. Leptoptilus crumen-

iferus (2 examples). Mycteria americana. Pseudotantulus ibis

and Z'antalus loculator.—In all these Storks the conditions were

practically identical. The P. longus was large and strong with

superficial and fascial origins and no deep origin, very broad

anchor and good slip to III. The P. brevis was absent, but

running across the tarsal joint parallel with the slip to III, and

therefore approximately at 1ight angles with the normal position

of the P. brevis tendon, was a stout elastic ligament which kept

the tarsal joint flexed, so that after death the joint could not be

straightened without force until this had been cut. At first I

thought that this ligament might represent the brevis, but its

direction is different, and it does not resemble the indubitable

vestiges of the brevis which occur in some other birds.

PERONEAL MUSCLES IN BIRDS. 1047

Tbididee. Carphabis spinicollis. Platalea leucorodia. Plegadis

Jalcinellus.—In these the P. longus is normal with superficial and

fascial origins, only a few fibres of deep origin, good broad anchor

and slip to III. The P. brevis is present but very weak, arising

by a few thin fibres from the greater part of the length of the

fibula and passing into a slender but long tendon with the usual

flat insertion.

Pheenicopteri. Phanicopterus antiquorum.—In the Flamingo

the P. longus is strong with only the superficial and fascial

origins, with good wide anchor and slip to I1I. The P. brevis is

absent.

The general picture presented by the Ciconiiformes is of a

large well formed longus muscle chiefly with superficial origin,

and a brevis muscle almost invariably slender and showing a

strong tendency to disappear, but none the less with the presence

absence following the minor divisions of the assemblage,

and not being sporadic in the genera.

ANSERIFORMES.

Palamedee. Chauna chavaria. Palamedea cornuta.—Il

have already described the conditions in these (supra, p. 1040).

Anseres. Cygnus olor, C. wigricollis, dx galericulata

and 4a sponsa. Cereopsis nove-hollandie. Chloéphaga inornata.

Dendrocygna fulva. Querquedula castanea. Metopiana peposaca.

“demia nigra. Mergus serrator.—In the Swans, Geese and

Ducks the P. longus is large with a very broad superficial origin

wrapped round the front of the knee, partly covering and partly

with fascial origin from the tibialis anticus and a lesser extension

towards the gastrocnemius. The muscle narrows rapidly (text-

fig. 181) to a rounded tendon from which a very broad anchor,

frequently i in separate slips, is given off, the main tendon forming

the slip to III. The P. brevis is always present and usually

large, with a central tendon and pinnate slips of fibres from the

eveater part of the fibula and adjacent surface of the tibia. It

frequently dips under a fibrous bridge and then flattens out and

has an unusually wide insertion to the abaxial side of the end of

the tibio-tarsus. There is usually one, and not infrequently a

second separate tendon running across the tarsal joint. ‘The

variations within the group are small, all the normal parts being

present. The deep origin of the longus is usually very shght, but

as an exception it is well- marked in Dendrocygia. The anchor

is very broad and short, and usually in separate slips, but in the

ducks generally and in Dendrocygna it is rather more fan-shaped.

In @demia the brevis is very short, arising only from the tibia

below the fibula and from a small portion of the fibula; its

tendon, moreover, instead of crossing under the slip to II] runs

parallel with it, and is inserted to the axial edge of the tarsus-

metatarsus.

In the Anseriformes the peroneals are both well developed,

1048 DR. P, CHALMERS MITCHELL ON THE

the longus, however, being relatively the larger, but the brevis is

actively functional and in addition to its flexion of the tarsal

joint has a notable rotating action.

Text-fig. 181.

Th] fy,

ANCH: 748.

Peroneal muscles in Ax sponsa.

Description and lettering asin Text-figure 179.

The Per. longus has had the proximal portion removed to show the P. brevis.

FALCONIFORMES.

Catharte. Cathartes wwra(2 examples).—The P. longusis a

strong muscle, with good but rather narrow superficial origin, with

fascial origin chiefly on the side of the tibialis anticus, and with

extensive deep origin. The anchor is broad and short and the

slip to IIT is very strong. The P. brevis is very slender, arising

only from the distal two-thirds of the fibula, then passing into a

rounded tendon much weaker than the slip to III under which it

runs, and then flattens out to the normal insertion. The domin-

ance of the longus over the brevis recalls the condition in the

Ciconiiformes and differs from that present in Accipitrine birds

(excluding Serpentarius).

PERONEAL MUSCLES IN BIRDS. 1049

Accipitres. Serpentariidee.—I have had no recent opportunity

of examining a Secretary bird, but I infer from Dr. Gadow’s

statement (‘‘ Aves” in Thier-Reich) that both peroneals are present,

and that as P. longus is very large the condition is generalized and

thus ike what exists among the Ciconiiformes.

Vulturide and Falconide. Gypactus barbatus. Tinnanculus

alaudarius. Falco peregrinus (2 examples). Aquila verreauat.

Haliaétus leucogaster. Ictinia mississipiensis. Circus gouldi.—In

all these birds both peroneals are present, but the P. brevis is the

more powerful and the P. longus tends to be reduced only to its

deep origin. In the Laimmergeier there is the greatest re-

semblance with the normal arrangement. The P. longus has all:

Text-fig. 182.

P LON.

ANCH>-</

Pas A

Peroneal muscles of Falco peregrinus.

Description and lettering as in Text-figure 179.

Fib. Anterior edge of fibula.

three origins, but the superficial origin is rather narrow and

partly reduced. In all the others the superficial and fascial

origins have practically disappeared, so that the muscle, as shown

in the case of the Falcon (text-fig. 182), almost exactly balances

the P. brevis, arising from the other side of the fibula and the

adjacent area of the tibia along the greater part of the length of

1050 DR. P. CHALMERS MITCHELL ON THE

the shaft. The anchor is always present and is usually rather

long and slightly fan-shaped. The slip to III is always present

and normal. The P. brevis is very strong; it arises in the

Liimmergeier only from the lower end of the fibula and the tibia

distal of this; in J/etinia its origin begins higher up; in the

others, as shown in text-fig. 182, it arises from nearly the

whole length of the fibula below the biceps insertion, from the

adjacent surface of the tibia and from the tibia distad of the end

of the fibula. Its very strong tendon passes under a fibrous

bridge which may be calcified in old birds, then flattens out and

passes in the usual synovial sheath under the slip to III to its

normal insertion.

It is plain that in these birds-of-prey the superficial origin of

the P. longus has been partly or completely lost and that the

P. brevis has greatly increased in relative importance, until it

may surpass the P. longus.

Pandionide, Pandion haliaétus—In the Osprey I found no

trace of the P. longus, in which Iam confirmed by Dr. Gadow.

The P. brevis was short and not very strong, arising from the

usual origin limited to the lower half of the shaft and passing

into a weak tendon, which after traversing a bony bridge,

flattened out and had the normal insertion of this muscle. As

ornithologists, arguing from other features, have differed as to

placing the Osprey with the Hagles and Vultures, or with the

Owls, it is of some interest to note that in the Owls also only the

P. brevis is present.

Apart from Pandion, it is clear that the Falconiformes

generally show a gradual reduction or specialization of the

P. longus, always retaining, however, the deep origin, the anchor

and a strong normal slip to ILI, and a gradual increase in size of

the P. brevis.

Gadow unites the Ciconiiformes, Anseriformes, and Falconi-

formes into the ‘‘ Legion” Pelargomorphe. Again excepting

Pandion, it appears that throughout the Legion, the P. longus

remains a large and important muscle always with a good anchor

and good slip to III, but that the P. brevis offers every condition

from that of perfect equality with or even superiority to the

longus in the specialized Falcons, to complete absence.

ALECTOROMORPH &.

TINAMIFORMES.

Crypturi. Rhynchotus rufescens (4 examples). Nothura

maculosa. Calodromas elegans (3 examples).—In all the Tinamus

J examined, the P. longus was well developed, with very large super-

ficial and fascial origins and long deep origin. ‘The anchor was

strong and broad and the slip to III strong. J noticed that in a

Rufous Tinamu the P. longus first straightened the tarsal joint and

then flexed sharply the second as well as the third digit. On dis-

secting out the tendons, I found that the tendon of the perforated

PERONEAL MUSCLES IN BIRDS. 1051

flexor muscle of the third digit, after being joined by the tendon

from the peroneus, gave off not only the slip to the perforated and

perforating flexor of digit ITI which occurs in birds belonging to

many different groups, but also a slip to the perforated flexor

tendon of digit II, an arrangement which I have not noted or

found recorded in any other bird.

The P. brevis was present in all the Tinamus I examined; in

Rhynchotus it was very feeble and slender, arising from the greater

part of the length of the fibula and passing into a thin tendon

which flattened out, and passing under the slip to III had the

normal insertion. In Wothura its origin was from the lower

part of the fibula, and in Calodromas it was extremely minute

and reduced to a few fibres and a tendon just visible, but

running the usual course. The definite presence of the P. brevis

in Tinamus is in contrast with its equally definite absence in

most of the Struthious birds, but resembles the condition in

Apteryx.

GALLIFORMES.

Mesitidee.—I have had no opportunity of examining Jesites,

and Milne-Kdwards does not refer to the peroneal muscles. It

would be interesting to know their disposition, as in Hurypyga

the tendon is characteristically long, and very different from the

arrangement in Galliform birds.

Turnices. Turnix dussumieri.-—The P. longus is present and

rather strong, with the three origins, a good anchor and slip to

If. The P. brevis is slender but arises from nearly three-quarters

of the shaft, from the fibula and tibia, and passes into a short

tendon which flattens out to the normal insertion.

Galli.

Megapodiide. Talegalla lathami.—The P. longus is very strong

with large superficial and good fascial origins but no deep origin.

Anchor strong but narrow, and good slip to III. The P. brevis

is also good, from the posterior three-quarters of the shaft from

fibula and tibia. Tendon is very short, passes under a fibrous

bridge and then flattens out to pass to usual insertion.

Cracide. Ortalis ruficauda. Mituatuberosa. Pauais galeata.

Penelope purpurascens.—In all these the P. longus is enormous

but chiefly from superficial and fascial origin, the deep origin being

just present. The anchor is very broad, short and strong, and the

ship to III is good. In some the tendon was ossified down to

the anchor, The P. brevis was strong, arising from the distal

three-quarters of the shaft, from the fibula, adjacent surface of

tibia and tibia distal to the end of the fibula. The tendon then

passed through a long fibrous bridge, partly calcified in Ortalis,

and flattened out to usual insertion.

Gallide. Guttera cristata, G. edowardi, and G. pucherani. Mele-

agris gallopavo. Argus giganteus, Gallus gallus (2 examples),

1052 DR. P. CHALMERS MITCHELL ON THE

Calophasis ellioti. Crossoptilon mantchuricum (2examples). Lol-

lulus roulroul. Ortyx virginianus (2 examples). Lophortyx

douglasi. Coturnix coromandelica. Francolinus infuscatus.—In

all these Galline birds the P. longus was enormous, with very

strong superficial, well-marked Suen and rather slight deep

origins. The anchor was stout and in most cases rather broad and

short, but in the Francolin was narrow and rounded. The slip to

IIT was invariably strong. The tendon of the longus was partly

ossified down to the anchor in the older birds. The P. brevis

was present in all, but in most cases rather slender, and its origin

was from the distal three- quarters of the shaft including the

fibula, adjacent surface of tibia and tibia distal to the fibula,

The rather short tendon (long in one of the Quails) usually

passed through a fibrous bridge and then flattened out to

normal insertion. In Pucheran’s Guinea-fowl, however, the

tendon did not pass across the tarsal joint, but ended on a knob

on the distal end of the tibio-tarsus in about the position

where it occasionally passes through a fibrous or calcified bridge.

The P. brevis was relatively weak in all these birds, and the

condition in the Guinea-fowl is at least complete functional

degeneration, but I have found no case of complete absence,

although [ examined Penelope and Gallus gallus, in which Dr.

Gadow found it absent.

Opisthocomi. Opisthocomus cristatus (2 examples).—The P.

longus is of moderate size with only a rather narrow superficial

origin. Good anchor and slip to III present. The P. brevis is

relatively rather strong, arising from the distal three-quarters of

the shaft in the normal fashion and with a short tendon flattening

out to the usual insertion.

In the Galliformes the P. longus remains the dominant muscle

and is always well developed, with, however, seldom much deep

origin. The P. brev is 1s present nae obviously less important.

GRUIFORMES.

Rallide. Mulica leucoptera. Gallinula chloropus and G. phani-

cura. Tribonyx mortiert. Hydrornia allent. Porphyrio porphyrio

(2 examples). Ocydromus australis. Porzana carolina. Aramides

chiricote and A. ypecaha. Hypotenidia philippensis (2 examples).

Rallus longirostris, R. maculatus, and R. celebensis.

In all these Rails (text-fig. 183) the P. longus is a strong

muscle usually with an extensive superficial and good fascial

origin and rather little deep origin. The muscular fibres converge

to a powerful tendon, frequently ossified, about the middle of the

length of the shaft. The anchor is very strong and tends to be

broken into separate slips; the slip to IIT is always present, with

the usual course. The P. brevis is alsoalways present and rather

strong, arising from the upper half of the fibula and the adjacent

surface of the tibia. It then passes into a stout tendon, some-

times ossified, and having traversed a fibrous or calcified bridge

flattens out to the usual insertion. The text-figure, drawn from

PERONEAL MUSCLES IN BIRDS. 1053

a Green-backed Porphyrio, gives the arrangement usual in the

Family.

Gruide. Gruine. Grus australasiana. Anthropoides virgo.

Balearica chrysopelargus.—In the Cranes the P. longus is rather

large with extensive superficial, fascial and deep origins, very long

tendon sometimes ossified, very wide anchor and good slip to IIT.

The P. brevis is present and arises from nearly the whole length

of the fibula but is very weak. The tendon, sometimes ossified,

has the usual insertion by a flattened extremity which passes

under both the slip to III and a very long tarsal ligament.

Text-fig. 183.

Peroneal muscles of Porphyrio porphyrio.

Description and lettering as in Text-figure 179 (p. 1041).

Br. Fibrous bridge.

Aramine. Aramus scolopaceus.—The P. longus is large with

superficial and fascial origins but without deep origin. It has a

stout anchor and good slip to III. The P. brevis is either

altogether absent or represented by a very small ligament.

Psophiine. Psophia crepitans, P. leucoptera, and P. obscura.—

The P. longus is large with chiefly superficial and fascial origins.

The anchor is broad and the slip to III is present and stout, but

peculiar in so far as after giving off a stout anchor to the per-

forated flexor of III in the region where it usually fuses with that,

it passes on and joins much more distally the tendon of the

perforated and perforating flexor of the same digit. The P. brevis

Proc. Zoou. Soc.—1913, No. LXX. 70

1054 DR. P. CHALMERS MITCHELL ON THE

is present, arising from nearly three-quarters of length of the

shaft and with usual insertion.

Dicholophide. Cariama cristata (3 examples).—The P. longus

is large with superficial and fascial and slight deep origins and a

very long tendon which gives off a very broad anchor and is con-

tinued as aslip to III. The P. brevis is represented by a minute

muscular head arising from the fibula and tibia opposite the

biceps ligament, and a very long slender tendon close to the tibia

which after crossin g¢ the tarsal joint flattens out to the usual

insertion.

Otidide. Otis tarda (2 examples).—The P. longus is very

strong with superficial, fascial, and separate deep origins. The

tendon is not longand after giving off a broad anchor is continued

as the slip to III. The P. brevis was absent.

Rhinochetide. Rhinochetus jubatus.—The P. longus was very

large with all three origins, a stout anchor and good slip to IIT.

The P. brevis was also ‘well developed with an extensive origin

along the length of the fibula and adjacent surface of the tibia

and had the usual flattened insertion ; 1t appeared to me to be

larger in the Kagu than in any of the other Gruiform birds.

Kurypygide. Hurypyga helias—The P. longus was_ well

developed with all three origins, a broad anchor and good slip to

III. The P. brevis was ale relatively strong, arising from the

upper half of the fibula and adjacent tibia and passing ‘into a long

tendon which flattened out as it approached the normal insertion.

Heliornithide. Heliornis fulica seu Podoa surinamensis (2

examples). Podica senegalensis.—The P. longus was large in all

these but with only superficial and slight fascial origins, the

deep origin being practically absent. The anchor was very broad

and strong, but the usual slip to II, although certainly present,

was delicate and could hardly have been functional. The P.

brevis was large and strong, with a rounded origin from the

proximal portion of the fibula, then a very long tendon partly

ossified which passed under a partly calcified bridge to flatten out

to the usual insertion.

Of the Gruiform birds the Rails and the Kagu seem to

present the most generalized condition of the peroneals since

both are present “and actively functional, the longus being

the dominant muscle. In the Cranes, Seriemas and “Basinzils.

the brevis shows signs of disappearing. The Sun-bitterns and the

Finfoots stand apart from the others, the former recalling the

condition most frequently found in the Limicole, and the latter,

with the tendency for the longus to be reduced, being quite

peculiar in the group.

CHARADRIIFORMES.

Limicole.

Charadriide. ZLimosa lapponica. Actitis hypoleucus. Gallinago

celestis. Machetes pugnax. Scolopax rusticula (2 examples).

PERONEAL MUSCLES IN BIRDS. 1055

Himantopus nigricollis. Recurvirostra avocetta. Hamatopus

ostralegus. Vanellus vanellus (3 examples). Charadrius pluvialis.

Rhynchea capensis.

In all these the P. longus is rather a small muscle with a broad

superficial, slight fascial, and very little deep origin from the edge

of the fibula. It narrows rapidly to a long tendon which gives

off a broad ancbor and ends in the usual slip to III. The

P. brevis is always reduced, arising only from the proximal part

of the fibula and passing into a lopg slender tendon which

passes down the leg parallel with the tendon of the longus, dips

under a fibrous bridge at the distal end of the tibio-tarsus, and

then passing under the slip to III flattens out to the usual in-

sertion. In the Avocet, twoexamples of the Lapwing, and in the

Oystercatcher, the brevis was reduced toa slender ligament which

arose from the distal end of the tibio-tarsal shaft in about the

position where it would normally pass through a fibrous bridge,

and passed across the tarsal joint to be lost in the fascize under

the shp to IIT.

Chionide. Chionis alba.—The P. longus is present with all

three origins, the deep being very feeble. The muscle passes into

a very long tendon which has the usual broad anchor and slip to

Ill. The P. brevis is also present but extremely feeble, a few

fibres from the fibula passing into a delicate tendon with the usual

insertion.

Glareolide. Glareola pratincola (2 examples).—Precisely as in

Chionis, the tendon of the feeble P. brevis being still longer and

more slender. i

Thinocorythide. ZThinocorys ? species.—The P. longus is as in

the Glareolide and Chionide, but the P. brevis is stronger, with

an extensive origin from the fibula passing into a long tendon

with usual insertion.

(Edicnemide. Wdicnemus scolopax.—The P. longus is large

and strong, with all three insertions passing into a rather shorter

tendon with broad anchor and strong slip to III. The P. brevis

is represented at most by a vestigial tendon.

Parride. Hydrophasianus chirurgus. Phyllopezus africanus.

Jacana jacana. Asarcia variabilis.—In all these long-toed, long-

legged Jacanas the P. longus is rather large with superficial,

fascial, and good deep origins. The broad band of muscle then

passes into a very long tendon which gives off a wide but short

anchor and is continued as the slip to III. The P. brevis arises

from the upper third of the fibula and adjacent tibia and gives

rise to a very long tendon which flattens out to the usual

insertion.

Lari.

Laride. Larus argentatus, L. ridibundus.—The P. longus has

only the superficial origin, and the muscle converges to a very

long tendon which gives off a broad anchor and then forms the

usual slip to IIT. The P. brevis arises only from a small

1056 DR. P. CHALMERS MITCHELL ON THE

proximal portion of the fibula and then forms a very long tendon

which runs down the leg parallel with the tendon of the longus,

dips under the slip to III, and has the usual flattened insertion.

Alcide. Alcea torda.—The P. longus is fairly large, with super-

ficial and fascial origins, then a long tendon which gives off a

broad anchor and ends as the normal slip to IIT. The P. brevis

is also rather strong but much inferior to the longus. It has an

extensive origin from the fibula and then passes into the usual

tendon with normal insertion.

Pterocles. Pterocles alchata (2 examples).—The P. longus

is broad and rather strong, with all three origins and broad

anchor and slip to IIi. The P. brevis is absent, so that in this

respect the Sand-Grouse agree with the general tendency of the

plover-like birds and differ from the game-birds.

Columbe. Starnenas cyanocephala. Leucosarcia picata

(2 examples). Phlogwnas cruentata and P. luzonica. Geophaps

plumifera. Phaps chalcoptera and P. elegans. Columbula picur.

Geopelia cuneata and G. tranquilla. Zenaidura carolinensis.

Carpophaga rufigula. Columba livia (many examples, wild and

domestic). Osmotreron bicincta (2 examples).

The Doves and Pigeons present an interesting series of modifi-

cations. A state of affairs very closely resembling what occurs

in Limicolous birds is not infrequent. In Leucosarcia, for

instance, the P. longus is very strong, with good superficial and

fascial origins and a considerable deep origin from the proximal

portion of the fibula. The muscular mass narrows to a very long

tendon, which eventually gives off a broad anchor and runs on

to form a good slip to Ill. The P. brevis similarly arises chiefly

from the pr roximal portion of the fibula and the adjacent surface

of the tibia, passes into a long tendon which after traversing a

fibrous bridge flattens out to the usual insertion. The P. longus

1s definitely the dominant muscle, but the brevis is well formed

and functional. In Columba the conditions are similar, but the

extent of the fibular or deep origin of the longus varies, and the

length of the tendon varies inversely with it. In Carpophaga

the fibular origin of the longus is still more important, and in

many of the smaller Pigeons and Doves, and in Osmotreron, the

deep origin is the more important, the superficial and fascial

origins getting weaker; the tendon is shorter, but the whole

muscle is relatively feebler. So also in the smaller Doves and

Pigeons, the P. brevis tends to degenerate, arising only from the

lower end of the fibula and the tibia distad of aie, so that the

tendon is very short and very different from the typical Limicoline

condition. On the other hand, in Osmotreron, the P. longus has

become smaller and reduced to its deep origin with a relatively

short tendon, but the P. brevis has increased in size and is

actually stronger than the longus.

In Charadriiform birds generally the P. longus is the dominant

muscle and the P. brevis tends to become weaker or to disappear.

PERONEAL MUSCLES IN BIRDS, 1057

In the case of both muscles, the normal arrangement is for the

muscular bellies to be limited to the proximal portion of the leg

and to give rise to very long tendons. The Pigeons and Doves

show thie type of the Chavadviiform group, but also indications of

moving away from it.

In the Alectoromorph group generally, the P. longus remains

as the dominant muscle and, except in the aberrant Finfoots, is

always large and powerful. ‘The P. brevis varies, sometimes

being very feeble or absent, but in some of the Columba, as: an

exception, shows signs of outstripping the P. longus. ‘The great

lengthening of the tendons of both muscles in some of the

Ralliform and most of the Charadriiform birds may easily be

regarded as in correlation with the long legs of these birds. It

is interesting to note that the Columbze show signs of a former

Jong-legged Charadriiform condition.

CoRACIOMORPH &.

CUCULIFORMES.

Cuculi.

Cuculidee. Cuculus canorus (2 examples). Guira piririqgua

(2 examples). Hierococcyx varius.—In the Guira cuckoo the

P. longus is a large muscle with good superficial, fascial, and

deep origins. Its tendon gives off a long narrow anchor and

then passes on toform the shp to Jil. In Hierococcyx the longus

is relatively smaller and the anchor is broader. In Cuculus the

deep origin is much the most important, and is chiefly from the

distal end of the shaft. In all the P. brevis is well developed,

with a strong origin from the tibia below the fibula and a varying

extension up the fibula. ‘The tendon of insertion runs the usual

course under the slip to III, and flattens out to an insertion

placed so that the muscle not only flexes the foot but has a con-

siderable power of rotation.

Musophagide. Corythaix persa (2 examples). Twracus cory-

thaix.—The P. longus is a large muscle with extensive superficial

and fascial origins and a small deep origin extending down the

fibula. The anchor is very broad, and the slip to IIT well marked.

The P. brevis is a long rounded muscle arising from about the

distal three-quarters of the length of the shaft, from the fibula

and tibia below the fibula. The tendon passes through a fibrous

bridge and then flattens out to the usual insertion.

In the Cuculi the P. longus remains the dominant muscle, but

there are signs of its being reduced to a deep origin with corre-

sponding increase of the importance of the P. brevis. The group

shows, in fact, a transition from the condition which I regard

as more generalized towards the particular type of specialization

which becomes more and more marked in other Coraciomorphines.

1058 DR. P. CHALMERS MITCHELL ON THE

Psittaci. Stringops habroptilus. Melopsittacus undulatus.

Nymphicus uveensis. Nanodes (Lathamus) discolor. Cyano-

rhamphus alpinus. Platycercus eximius. Aprosmactus cyano-

pygius. Paleornis fasciata. LHclectus pectoralis (2 examples),

E. roratus. Poeocephalus meyeri. Caica melanocephala. Pachynus

brachyurus. Chrysotis (¢ species). Myopsittacus monachus.

Conurus jendayi. Calopsittacus novee-hollandie. Cacatua sul-

phurea. Trichoglossus novee-hollandice. Loriws domicella.

Parrots carry further the specialization of the generalized type

which is already indicated in the Cuculi. In every case both

muscles are present, but there are different degrees to which the

longus is reduced and the brevis increased. In Stringops (text-

fig. 184) the P. longus is a large muscle with good superficial and

fascial origins and a deep origin extending nearly the whole

length of the shaft. The muscular belly narrows to a short

tendon which is inserted to the capsule of the flexor tendons

representing what I term the anchor in this memoir. A few

Text-fig. 184.

Peroneal muscles of Stringops habroptilus.

Description and lettering as in Text-figure 179.

Br. Fibrous bridges.

delicate strands can be made out passing in the direction of the

usual slip to III, but that structure is certainly absent func-

tionally, even if there be a trace of it morphologically. I have

found a small superficial origin of this muscle in several other

Parrots ; it was relatively large in Calopsittacus, and present in

Cyanorhamphus and Caica. Usually, however, it has been lost,

and the normal condition in Parrots is for the P. longus to be

reduced to a deep origin. ‘This may be large as in £electus,

long but slender as in Platycercus, or it may be so small as in

PERONEAL MUSCLES IN BIRDS. 1059

Chrysotis and so closely attached to the fasciz of the P. brevis

that Dr. Gadow has described it as fused with that muscle.

I found a minute but distinct slip to III in Calopsittacus nove-

hollandie, and faint traces of fibres in the direction usually

taken by that slip in Caica, Myopsittacus, and one or two

others. The P. brevis in Stringops (text-fig. 184) is a stout

rounded muscle arising from high up the shaft and passing into

a strong rounded tendon, which after traversing two fibrous

bridges flattens out to the usual insertion. In most cases,

however, 1t is much stronger than the P. longus, arising from

the whole length of the fibula below the biceps insertion, from

the adjacent area of the tibia and from the tibia distad of the

Text-fig. 185.

Peroneal muscles of Platycercus eximius.

Description and lettering as in Text-figure 179. s

Br. Fibrous bridge.

fibula. Its strong tendon usually passes under a fibrous bridge

and has the normal flat imsertion to the proximal end of the

tarsus-metatarsus shaft. Platycercus (text-fig. 185) shows the

most usual condition of this muscle in Parrots. In Trichoglossus

and some others the P. brevis is even larger, arising from the

head of the fibula proximad of the biceps insertion.

Parrots thus show conditions of the peroneals which link them

with the arrangements found in the groups I have already

discussed, but as a group are moving away from the normal type

in the Coraciomorphine direction.

The Cuculiformes retain marked indications of their former

1060 DR. P. CHALMERS MITCHELL ON THE

possession of the generalized condition of the peroneals, but have

moved away from that condition, the Parrots further than the

Cuckoos and Plantain-eaters.

CoRACIIFORMES.

Coracie:

Coraciidee. Coracias garrula, C. indica (2 examples). Hurystomus

orientalis.—The P. longus is rather weak, but retains a small

superficial, fascial, and slight deep origins. The anchor is very

short, and the slip to III is extremely slender but present, at

least occasionally. The P. brevis is stronger than the longus ;

it arises from about the distal three-quarters of the shaft, from

the fibula and tibia, passes under a fibrous bridge and is inserted

by the normal flat tendon.

Momotide. Momotus lessoni and M. subrufescens. Aspatha

gularis.—In these the P. longus is well developed with all three

origins. It has a short anchor and is continued as a good slip

to II]. The P. brevis is large and its tendon is just stronger

than that of the longus. It arises from the proximal half of

the fibula with the adjacent surface of the tibia and gives rise

toa rather long, rounded tendon which flattens out at the normal

insertion.

Alcedinide. Alcedo ispida (2 examples), A. asiatica and A.

bengalensis. Ceryle aleyon, C. americana, C. inda, and C. maxima.

Ceyx -rufidorsa. Cittura cyanotis and C. sanghirensis. Dacelo

gigantea (3 examples). Halcyon pileata and H. rufa. Sauwropatis

chloris, S. sancta, S. sordida, and S. vagans.—As I have already

pointed out in an account of the Anatomy of the Kingfishers

(‘ Ibis, 1901, p. 97), the P. longus is present in Kingfishers but

is plainly degenerating, possibly in association with the degene-

ration of the fibula. It is best-marked in Dacelo; certainly I

cannot confirm Beddard’s statement (‘Structure and Classification

of Birds,’ p. 199) that it is absent in that bird, and he makes no

mention of its presence or absence in other Kingfishers. It has

a superficial origin reduced to a narrow tendon from the external

corner of the tibial crest, joined by a few fibres from the tibia

aleng the region of the fibula representing the normal deep

origin. It is ‘inserted to the capsule of the flexor muscles, this

being the usual anchor, but there is no trace of a slip to ITI.

This is the most common condition, but in a few, notably

Ceryle, Halcyon, and Ceyx, the muscle is reduced to a simple

tendon with only a few muscular fibres, the insertion being the

anchor, The P. brevis is always present and strong, arising

from the area of the tibia usually covered by the lower end of

the fibula and passing into a stout tendon which flattens out to

the usual insertion.

Meropide. Merops apiaster, M. philippensis—The P. longus

is present but is very much reduced, having only a narrow super-

ficial and a few fibres of deep origin. A very short anchor is

present, and the shp to II although present is very slender.

PERONEAL MUSCLES IN BIRDS. 1061

The P. brevis is much stronger, with a good muscular origin from

the distal three-quarters of the shaft; its tendon passes under

a fibrous bridge and flattens out to the usual insertion.

Upupidee.

Upupine. Upupa epops (2 examples).—The P. longus is absent.

The P. brevis is large, from the distal three-quarters of the shaft,

including both fibula and tibia. The tendon flattens out to the

normal insertion.

Bucerotinze. Bucorvus (? species). Rhytidoceros undulatus.

Lophoceros erythrorhynchus.—I\n the Hornbills the P. longus is

absent, although I find in my notes that there is a degenerate

set of tendinous fibres which might possibly represent a vestige

of it. The P. brevis is very strong, arising from the lower

half of the shaft, from the fibula and its fibrous continuation,

and from the adjacent surface of the tibia. The strong tendon

flattens out to form the usual insertion. In Secorvus the

P. brevis was equally strong but rather shorter.

Striges. Athene noctua (3 examples). Bubo lacteus (2 ex-

amples), B. maculosus, B. maximus (2 examples). Asio otus.

Text-fig. 186.

Peroneal muscles of Strix flammea.

Right leg, outer view. Muscle striped; tendon dotted.

P. BR. Peroneus brevis. P. B.1. Tendon of P. brevis.

Gastr. External head of Gastrocnemius.

Flex. Flexor muscle of foot.

Tib. Ant. Tibialis anticus.

Sl. Sling of tibialis anticus.

1062 DR. P. CHALMERS MITCHELL ON THE

Strie flammea. Speotyto cunicularia.—There is no trace of the

P. longus in any of the Owls. The P. brevis is enormous, but

usually arises only from the tibia below the fibula. In Strix

Jlammea (text-fig. 186) it is relatively longer, arising from all

the fibula distad of the insertion of the biceps, from the adjacent

tibial surface, and from the tibia distad of the end of the fibula.

In Athene it is equally long. The stout tendon flattens out after

passing through a fibrous bridge (absent in Strix) and is inserted

in the normal fashion to the proximal end of the tarsus-metatarsus.

Its chief action is to rotate the foot on the tibial shaft so as to

depress the great toe side and elevate the fourth digit side. It

comes into action after the enormous tibialis anticus has flexed

the foot.

Caprimulgi.

Steatornithide. Steatornis caripensis.—Vhe P. longus is absent.

The P. brevis is large and strong, with origin from high up the

shaft from both fibula and tibia and fairly long tendon which

flattens out to usual insertion.

Podargide. Podargus cuviert. Nyctidromus albicollis. df go-

theles nove-hollandie.—In Podargus the P. longus has a broad

superficial origin with no fascial or deep origin. It gives off a

broad anchor and is continued as a slender slip to III. The

P. brevis is stronger than the longus. It arises from fibula and

tibia just below the insertion of the biceps, and its rounded

muscular belly passes into a stout tendon which flattens out to

the usual insertion. In Wyctidromus the P. iongus is practically

the same as in Podargus, but the P. brevis is absent. In #go-

theles nove-hollandie the P. longus is absent, and the P. brevis

is well developed, as in Steatornis.

Caprimulgide. Caprimulgus europeus.—The P. longus is large

with superficial, fascial, and deep origins. It has a broad anchor

and a good slip to II]. The P. brevis is absent.

I have followed the usual arrangement of the genera of Capri-

mulgi that I have examined, and certainly I do not propose to

rearrange them simply on the evidence of the peroneal muscles.

It is plain, however, that so far as these muscles are concerned,

Podargus has remained in the primitive condition, with both

muscles present and normal; Steatornis and #gotheles have

specialized in the same direction as the Owls, by losing the

P. longus; Caprimulgus and Nyctidromus present a condition

very aberrant amongst Coraciiform birds, and by retaining the

P. longus and discarding the P. brevis recall the condition which

frequently occurs in the great assemblages which I have already

passed in review.

Cy pselli.

Cypselide. Cypselus apus.—The P. longus is absent. The

P. brevis is large and strong, from the proximal end of the shaft

including the head of the fibula and adjoining area of the tibia

down three-quarters of the shaft, then a very strong tendon

PERONEAL MUSCLES IN BIRDS. 1063

passing over the bridge of the tibialis anticus to usual flat

insertion.

Trochilidee. Amazilia felicie.—The P. longus was absent and

the P. brevis exactly as in the Swift.

Colu. Colius capensis.—The P. longus was present in a ves-

tigial condition, with only superficial origin and a*slender tendon

ending in the anchor, with no slip to III. The P. brevis was

very large and strong, arising from nearly the whole length of

the shaft, involving both fibula and tibia. Its strong but rather

short tendon flattened out to the usual insertion.

Trogones. Zrogon puella and 7’. atricollis—The P. longus

was present but not so large as the P. brevis; its origin is chiefly

deep, and it has a broad anchor and fair slip to III. The

P. brevis is very large and strong, arising from the greater part

of the length of the shaft and ending in a short but stout tendon

which flattens out to the usual insertion.

Pie.

Galbulide. Galbula albirostris, G. rufiventris. Urogalba para-

disea.—My material for examining these was not good, con-

sisting of partly dissected specimens which had been in spirit for

many years. ‘The P. longus was certainly very small, but there

was a distinct tendon running to form an anchor, and in one

case a trace of the slip to Ifl. The P. brevis was larger than

the longus and had a stout tendon flattening out to the usual

insertion. I should be glad, however, to examine fresh specimens.

Capitonide. Megalema virens. Cyanops flavifrons. — The

P. longus was a large muscle with a wide superficial, and short

fascial and deep origins. The triangular muscular belly rapidly

narrowed to a stout tendon, which, however, was not so strong as

the tendon of the brevis, gave off a long and rather narrow

anchor and was continued as a good slip to III. The P. brevis

arose from the fibula beginning just below the biceps tendon,

and from a considerable part of the tibia, and passed into a

strong tendon which flattened out to the usual insertion. (Text-

fig. 187.)

Rhamphastidee. Aulacorhamphus prasinus. Pteroglossus in-

scriptus. Lhamphastos discolor (2 examples). Selenidera macu-

lirostris (2 examples).—-In all these Toucans (text-fig. 188) the

P. longus was of fair size but with chiefly fascial and long deep

origins. The short tendon gave off a narrow anchor and was

continued as a slender slip to III. The P. brevis was a stronger

muscle, its stout tendon forming the axis of a muscular mass

arising from the tibia and fibula for three-quarters the length of

the shaft. The tendon then passed through a strong fibrous

bridge and had the normal flat insertion.

Picidee. Dendrocopus major. Brachypternus aurantiacus. Co-

laptes mexicanoides. Gecinus vittatus. Hypoxanthus rivolit.—In

the Woodpeckers I found the P. longus always present but rather

weak and with chiefly superficial origin. Its tendon was very

thin and flat and formed the usual anchor, and gave off a thin

1064 DR. P. CHALMERS MITCHELL ON THE

Text-fig. 187.

psy /

iti Ih Hy

Vth Vy Hiff)

mn by

Peroneal muscles of Megalema virens.

Right leg, outer view. Muscle striped; tendon dotted.

P. LON. Peroneus longus. ANCH. Anchor to Sustentaculum. PL. III. Slip

to Flexor of III toe. P. BRE. Peroneus brevis.

Text-fig, 188.

Peroneal muscles of Rhamphastos discolor.

Description and lettering as in Text-figure 186 (p. 1061).

P. LON. Peroneus longus. ANCH. Anchor. P.L. III. Slip to III.

PERONEAL MUSCLES IN BIRDS. 1065

strand which in a fresh example of Dendrocopus formed the usual

slip to IIT, but which in the other Woodpeckers, of which I had

only very old spirit specimens, I could not trace completely to

their insertion. The P. brevis was strong, arising from the distal

three-quarters of the shaft, and ending in a short tendon which

flattened out to the usual insertion.

The Coraciiform birds form an interesting and difficult series,

but it is clear that the general tendency in the group is for the

P. brevis to increase and for the P. longus to be reduced. The

series, however, is not very coherent. A certain number have

remained almost in the generalized condition with the P. longus

possibly slightly reduced in its origin and equal to or weaker

than the P. brevis, but showing the anchor and slip to ITI, and

with the P. brevis a strong but not excessively strong muscle.

These central types are the Coracize, Momotide, Meropide,

Podargus amongst the Caprimulgi, and the Trogones, Capitonide,

and Rhamphastidee. In others the P. longus is still further reduced

and has lost the slip to III, whilst the P. brevis has relatively

still further increased. These are the Alcedinide (Dr. Gadow,

however, states that Pelargopsis has proceeded still further to

the complete loss of the P. longus), the Coli, the Galbulide

(probably), and most of the Picide. In yet another set the

specialization has proceeded to the complete loss of the P. longus,

and the P. brevis is always very strong. These are the Buce-

rotide, Upupide, Striges, and, amongst the Caprimulgi, Steatornis

and AMgotheles, the Cypselide and the Trochilide. Finally, in

Nyctidromus and Caprimulgus, although the P. longus is partly

reduced, the P. brevis is absent.

PASSERIFORMES.

Passeres Anisomyode.

Subclamatores. Hurylemus ochromelas. Cymbirhynchus

macrorhynchus.

Clamatores. Picolaptes affinis. Chasmorhynchus nudicollis.

Tyrannus melancholicus. Myiarchus tyrannulus. Pitangus

sulfuratus. Pitta strepitans.

Passeres Diacromyode.

Suboscines. Menura superba.

Oscines :—

Corvide, Cracticus destructor. Creadion carunculatus. Cyano-

corax luxuosus.

Paradiseidee. Parotia lawesii. Mluradus melanocephalus.

Sturnide. Hnodes erythrophrys. Mimo dumonti. <Acrido-

theres fuscus.

Icteridx. Jcterus jamaicai. ‘Ostinops decumanus.

Ploceide. Hstrelda phenicotis.

Tanagride. Tanagra sayaca. Rhamphocelus brasilius.

Cerebide. Coreba cyanea.

Meliphagide. Hntomyza cyanotus. Acanthorhynchus (2

species).

1066 DR. P. CHALMERS MITCHELL ON THE

Nectariniide. <Avrachnechthra zeylonica.

Troglodytide. Campylorhynchus wnicolor.

Hirundinide. Hirundo rustica.

Laniide. Lanius excubitor.

Artamidee. Artamus lewcogaster.

Dicruride. Dicrurus (? species).

Oriolide. Oriolus galbula.

Parride. Liothrix luteus.

Turdide. Zurdus tristis. Merula tamaulipensis. Geocichla

citrina. Mimus orpheus.

The Passeriform birds that I examined were well distri-

buted over the divisions into which systematists have attempted

to divide this group, and presented a fair sample of the manifold

types of habit and size that occur in the group, a sample that I

take to be fairly representative, as I found extremely little

divergence. The type which recurs throughout the group with a

uniformity that is almost tiresome, is well shown in the figure of

Parotia lawesit (text-fig. 189). Both peroneals are present, well

Text-fig. 189.

Peroneal muscles of Pavotia lawesii.

Description and lettering as in Text-fig. 188 (p. 1064).

developed and functional. The longus has a broad superficial,

good fascial, and fairly long deep origins; the muscular mass is

an elongated triangle which ends in a short tendon which almost

at once forks, the shorter and stouter fork forming a long narrow

anchor, the other forming the “slip” to III, This ‘‘V ”-shaped

forking is very different in appearance from the broad and short

anchor which is the more common type in other groups, but it

occurs also in other birds and can be regarded only as typically

not characteristically passerine. The P. ‘brevis arises from the

PERONEAL MUSCLES IN BIRDS. 1067

fibula below the insertion of the biceps tendon and from the

adjacent surface of the tibia; the fusiform belly gives rise to a

short tendon which frequently, but not invariably, traverses a

fibrous bridge and then flattens out to the normal insertion to

the proximal end of the tarsus-metatarsus.

The minor differences which are to be found, but too irre-

gularly to be correlated with size, habit, or systematic position,

relate to the relative sizes of the two muscles. Sometimes the

P. longus is rather feebler with little deep origin and a relatively

narrow superficial origin; the P. brevis, on the other hand,

becoming very thick and strong. The muscular bellies of both

muscles may be shorter and their tendons longer.

In only two of the Passeriform birds examined did I find

notable differences. In Menwra superba, the inclusion of which

amongst the Passeriformes I doubt on other grounds, the anchor

of the P. longus was much shorter and rather wider than usually

occurs In the group, and the P. brevis was relatively considerably

weaker. In Hirundo rustica the P. longus had the usual broad

superficial origin but very little deep or fascial origin, and the

slip to IIIT was absent. The P. brevis was of the usual Passerine

type and relatively weaker than in the Swifts.

The Coraciomorphine birds present many different phases of

the degeneration of the longus and of the increasing importance

of the brevis, and in this respect are in marked contrast with the

other Legions into which Dr. Gadow has arranged the Orders of

birds.

SUMMARY AND CONCLUSIONS.

The Peroneus longus muscle shows every gradation from

elaborate structure and apparently important function to com-

plete absence, and the facts fall into a coherent picture if we

suppose that we have to deal with loss of parts originally present.

The presence and absence of origin from the fasciz of the neigh-

houring muscles are the most irregular and may well be interpreted

as in direct relation to function. If the muscle be large and

important, it overgrows its bounds and comes in organic contact

with the adjacent fasciz; if it is smaller and less important it

remains isolated. When the muscle as a whole appears to be in

process of degeneration, this may proceed in two ways. Occasion-

ally, but rarely, the deep origin begins to disappear first, and the

superficial origin gradually narrows until it becomes only a thin

superficial tendon. More often the process starts with the

degeneration of the superficial origin, and this is often accom-

panied by strengthening and lengthening of the deep origin,

until finally nothing but the deep origin is left. This next

gradually shortens until the whole muscle disappears. Of the

tendons of insertion, the slip to ILI goes first and the anchor

persists. There are many cases in which the slip to III has been

lost, the anchor remaining as the only insertion, but I have found

no instance in which the anchor has disappeared leaving only the

1068 DR. P, CHALMERS MITCHELL ON THE

slip to III. The pull on the anchor by the contraction of the

muscle not only straightens the tarsal joint, but steadies the

sustentaculum and therefore assists the action of all the flexors

of the toes; the slip to III acts merely as an accessory to the

proper flexor of the third toe.

I find the Peroneus brevis very difficult to understand. Its

rotating action is often extremely slight, and in its action in

flexing the tarsal joint it appears to do little more than assist

the much more powerful tibialis anticus, the mechanical arrange-

ment of which is more favourable. So far from it being sur-

prising to find that it has degenerated or completely disappeared

in so many groups, it is remarkable that it should have been

preserved at all. In the cases in which it has become stronger

and has surpassed the longus, it seems to me that its power of

rotation is greater, and in a number of cases, particularly in

Passerines, its contraction appears to have the effect of partly

flexing the toes and the great toe, as if its action would assist in

perching. I am afraid, however, that an equally elaborate

investigation of all the muscles of the lower leg would be necessary

before the varying conditions of the peroneals could be inter-

preted in terms of function.

On the information at my disposal I find it extremely difficult

to associate the conditions of the peroneal muscles with differences

in habit that point directly to functional adaptation. On the

other hand, with a few, a very few, exceptions to which I shall

refer later, there is a close conformity between the condition of

the peroneals and what appear to be the most securely founded

systematic divisions. Birds seem to have this or that type of

peroneal muscle, not because they are arboreal or terrestrial,

swimmers or waders, scratchers, predatory or vegetarian, but

because it is the type occurring in this or that systematic division.

Whatever be their habit they seem to make shift with the type

of peroneal which occurs in their group. All the four Legions

into which Dr. Gadow groups the Orders of Carinate birds, and

most of the orders themselves, show certain members with what

I have assumed in this paper to be the generalized, possibly the

more ancestral condition of the peroneals, and all show signs of

moving in a definite direction away from this generalized con-

dition. In the Ratites the longus is dominant but the group has

moved away from the central type by extreme reduction or total

loss of the brevis. Of the Colymbomorphe the Penguins show

the central type, the Grebes have a weak longus with the slip

to III occasionally absent, but the brevis is always rudimentary

or absent. In the Petrels the longus has always at least a super-

ficial origin, the anchor and slip to IIT, but the brevis has rela-

tively increased in size. In the Pelargomorphe (except Pandion),

the longus remains the dominant muscle and always has both the

anchor and the slip to III, and the brevis, except in the Falconi-

formes, tends to disappear. Some Steganopods, Herons and

Tbises, Screamers, and Ducks and Geese remain in the central

position. Other Steganopods and Storks and Flamingos have

PERONEAL MUSCLES IN BIRDS. 1069

lost the brevis altogether. The Falconiformes show an interesting

series. Serpentarius isin quite the central position; in Catharte

the superficial origin of the longus is slightly reduced, but the

muscle remains the dominant of the pair. In the Falconide

(omitting Pandion) the superficial origin and the longus generally

tend to be reduced, but the anchor and slip to III always persist

and are strong and functional; the brevis increases greatly and

may equal or surpass the longus. In so far the Falconiformes

present a parallel with the Owls, but are to be distinguished from

that group by the retention of the longus with its. anchor and

slip. In the Alectoromorph Legion the great majority remain in

the central condition. The longus is invariably present, the

superficial origin is almost invariably the stronger, the anchor is

always present and the slip to ITI absent only in the Heliorni-

thide. The brevis is almost invariably the weaker muscle and

shows a strong tendency to disappear, e. g. among the Tinamus

it may be excessively feeble, it 1s feeble or degenerate in some

Galliform birds, absent or reduced in some Gruide and some

Charadriide. The Columbe are specially interesting; the longus

is always present with anchor and slip to III, but there seems to

be a tendency for it to be reduced and for the brevis to increase,

especially in the smaller and most Passerine-like Pigeons,

In the Coraciomorphine Legion the characteristic tendency is

for the reduction of the longus at the expense of the brevis. The

Orders, however, show marked differences in the extent to which

this process has oecurred. Of the Cuculiformes, the Cuckoos and

Plantain-eaters remain almost in the central condition, and,

although the brevis is always well developed, it is surpassed by

the longus. The Parrots, like the Pigeons, show within the

group all stages from an almost central condition where the longus

is complete and surpasses the brevis, through stages in which the

longus is still well developed, although it has lost the slip to III,

to the final stage in which there is anne no trace of the loner

remaining. The Coraciiformes have moved furthest from the

central condition. A very few, the Motmots, some of the King-

fishers, Podargus among the Caprimulgi, and Trogons and Barbets

show almost the central condition, but even amongst them the

longus is usually very little superior to the brevis, although it

retains its parts. In most Coraciiformes the longus is at least

feeble, has usually lost its slip to III and is often entirely absent.

The Passeriformes, on the other hand, are remarkably constant

and very near the central condition, although the brevis may

equal or nearly surpass the longus, and in one case (H irundo)

the slip to III has been lost. In the diagram reproduced as

text-fig. 190, I have endeavoured to show in a diagrammatic

fashion the general trend of the modification of the peroneale in

the groups of birds. The central space represents the central or

primitive condition, the area to the left the degeneration of the

brevis, that to the right the degeneration of the longus. The

placing of the named enclosures represents roughly the position

Proc, Zoou, Soc.—1913, No. LX XI. ie

1070 DR. P. CHALMERS MITCHELL ON THE

of the different groups with regard to the central avian condition.

It is to be noted that nearly every group seems to have made

experiments in both directions, but as we read from above down-

wards in the diagram, we pass from movement towards the left to

movement towards the right. And it may also be noticed that,

in a general way, what are usually regarded as the groups ex-

hibiting the lower modifications of avian structure are towards

the top of the diagram, those representing the higher types

towards the bottom. If I had felt justified in expanding this

diagram, by placing the names of the minor groups in their

proper orientation with regard to the peroneals, it would have

been found in the same fashion, that inside each group, on the

whole, those which are wenerally regarded as the higher types

were towards the right hand side of the spaces, those representing

the lower types towards the left hand. In other words, I think

JI may say that the higher types of avian modification are associated

with a tendency for the degeneration of the Peroneus longus and

an increase of the Peroneus brevis.

Adaptation, direct or associated, may be the fundamental

explanation of the facts that I have tried to set out, but it is

difficult to follow, and it is easy to see that kinship appears to be

a more important factor. The Eagles and Vultures have many

adaptive resemblances with the Owls, but the former contain

members directly linking them with the primitive condition and

have never moved far from it, and the latter show the extreme

modification of the Coraciiform group. Swifts and Swallows have

many points in common, but the former, in the condition of the

peroneals, are extreme Coraciiform birds, the latter very slightly

modified from the true Passerine condition. So also Humming-

birds are extreme Coracuformes in this respect, and Sun-birds

are true Passerines. The family tradition appears even in many

of the minor changes; all the Storks have lost the brevis and

the Herons have retained it.

There remains to say a word as to the few birds which do not

lie comfortably, so far as the peroneals are concerned, in the

positions usually assigned them. In a memoir dealing with a

similar large series of facts in bird anatomy (“On the Intestinal

Tract of Birds,” Trans. Linn. Soc., Zool. viii. p. 173), I pointed out

what seems a logical necessity (frequently, i overlooked

by those who use anatomical characters for systematic purposes),

that if we have reason to believe a particular character to have

been ancestral, we cannot assume that animals now without it

are more nearly allied than those that have retained it. There

is no @ priort ground for assuming that it may not have been lost

twice or several times independently. It follows therefore that

the loss of the longus muscle, or of any important part of it, or

of the brevis muscle, is no valid clue to systematic position.

Tt is equally clear that the common retention of the ancestral

condition is no ground for placing the descendants of a particular

ancestral stock together, if relative affinity and not convenience

is to be the basis of classification,

PERONEAL MUSCLES IN BIRDS, 1071

A glance at the table (text-fig. 190) will show why anatomists

on the hunt for convenient systematic characters would take

Text-fig. 190.

<—— Peroneus longus greater ; ———>

than P. brevis, with super-

P. brevis ficial and deep origins, P. longus

disappearing anchor and slip to Ill. disappearing

P. brevis functional.

Colymbomorphee

Pelargomorphe

Alectoromorpha2

Cuculiformes

Coraciiformes

Passeriformes

Diagram of Peroneal museles in Birds.

The central space represents the condition regarded as primitive in this memoir,

the P. longus being fully developed, with superficial and deep origins, anchor

and slip to III, and stronger than the P. brevis, which, however, is present and

functional. The various enclosures represent the groups of birds indicated by

the lettering. Coincidence with the central space indicates that the primitive

condition of the peroneals is present; displacement towards the left indicates

reduction of the P. brevis, displacement to the right reduction of the P. longus.

1072 ON THE PERONEAL MUSCLES IN BIRDS.

little interest in the peroneals. Some of the Colymbomorphs,

most of the Pelargomorphs, most of the Alectoromorphs, most

of the Cuculiformes, a few Coraciiformes, and most of the

Passeriformes would have to be placed together. We know from

other reasons that the association would be absurd, but it is not

more illogical than to place Ornithorhynchus and H chidna in the

same Oren because they have retained a reptilian legacy that

was probably once the common property of all mammals. The

Ratites, some of the Colymbomorphs, some of the Pelargomorphs,

and a few of the Alectoromorphs would have to be placed to-

gether because of the absence of the brevis. Some of the

Colymbomor phs, and a few Pelargomorphs, a few Alectoromorphs,

a few Cuculiformes, most of the Coraciiformes, would similarly

have to be placed together because of their loss of the longus. A

rather tedious piece of work may be of some use, if only it serve

to direct attention to the confusion that must arise if characters

be not carefully analysed before they are employed for systematic

purposes, and to the information which apparently unruly facts

may afford, if they are plotted out over a sufficiently compre-

hensive field.

On the other hand, although the common retention and common

loss of ancestral character are no proof of affinity, I myself, like

most anatomists who have taken an interest in trying to correlate

their facts, have been impressed by the fashion in which allied

animals march along the path ef modification in the same

direction. I have found this to be the case in Pigeons (Journ.

Linn. Soe., Zool. xxvii. p. 210), in Kingfishers (‘ Ibis,’ 1901, p. 97),

in Gruiform birds (P. Z.8. 1901, p. 629), and in Limicoline birds

(P. Z.8. 1905, p. 155), and the general results which I have been

stating in this communication point in the same direction. It

appears to me therefore that the existence of notable unconformity

at least suggests that the position assigned in the System calls

for enquiry.

The first notable instance is Pandion, which in the matter of

the peroneals differs from all the Eagles and Vultures and con-

forms with the Owls. I cannot find that this point in its

structure has been recorded previously, but many systematists

have seen in Pandion a link between the nocturnal and diurnal

birds of prey, and others have actually placed it amongst the

Owls. The question calls for re-examination.

T am less impressed with the cases of Vyctidromus and Capri-

mulyus. So far as the longus is concerned they agree closely with

Podargus, but although the latter, like all other Coraciomorphine

birds, has a well-developed brevis, the two former have no brevis.

IT have already pointed out that at the best the action of the

brevis is difficult to understand, and appears to be frequently

superfluous. I am therefore of the opinion that its loss in these

isolated cases, however curious, is insignificant.

1HqdAUE VaaLlLadoONy We

: e eon

cepeemer tet _ Ee

“SID lel “SIG “S Z al

ON BRYDE’S WHALE. 1073

61. On the External Characters and Biology of Bryde’s

Whale (Salenoptera brydet*), a new Rorqual from

the Coast of South Africa. By SOrsan OLsEn,

Zoological Laboratory, Christiania University f.

[Received June 27, 1913: Read November 25, 1913. ]

(Plates CIX.—-CXIII.4)

INDEX. Page

Balanoptera brydet 0.0 ieee eeetee reese es LOTS

Distributions seater cnt ean ene ene OOD,

Introductory remarks.

Before the Norwegian whaling trade started in South Africa in

1909, opportunities for studying whales from that part of the

world seldom occurred, but when at rare intervals one of the

bigger whales was stranded on the 8. African coast, if it was not

at once eaten by sharks it was, as a rule, quickly destr oyed by the

warm climate and heavy sea breaking on the beach.

The country is very extensive, the communication along the

sea-shore bad, and journeys difficult and expensive. If, there-

fore, one of the few scientists of the country received a report of

a stranded whale, it was as a rule impossible for him to reach

the spot before it was too late. So when we take all these

difficulties into consideration it is not surprising that the bigger

whales from this part of the world have remained unknown until

quite recently.

Mr. W. L. Sclater has only the following to say about the

enus Balenoptera in the standard work ‘ Fauna of South Africa’

(1901) :—* Fin-whales are not uncommon off the coast of 8. Africa,

but so far as I know, no specimen has ever been secured for a

museum, or been examined by a competent authority.” After

this he gives the characteristics of the four northern fin-whales,

in order to facilitate comparison with the Cape species of the

same genus.

The South African whaling industry was founded by Consul

Johan Bryde, from Sandefjord, Norway, who in 1909 erected the

first factory in Durban and another in the following year in

Saldanha Bay on the west coast.

When the Norwegian whalers returned home after having

finished their first season in Saldanha Bay, they mentioned a

whale which had been found outside the latter place, and which

was very similar to the ‘“ seihval ” (Balenoptera borealis).

As the ‘“seihval” (Rudolphi’s whale) had been only known

%* (I have consulted the issue of the Norwegian newspaper ‘Tidens Tegn’ dated

November 12th, 1912, in which this name was first published by the author.

Technically the species dates from the description in that Journal, but no details

are given there which are not fully explained here.—Ep1ror. |

ar “Communicated by the SECRETARY.

+ For explanation of the Plates see pp. 1089, 1090.

1074 MR. GRJAN OLSEN ON

until then as inhabiting the eastern parts of the North Atlantic,

it was very desirable from a scientific point of view to get further

information about the species from Saldanha Bay, and also about

other South African whales, by studying them on the spot.

Mr. Johan Bryde, who has often displayed his interest in and

generosity towards science, then offered me the necessary pecuniary

assistance, and thus enabled me to undertake a journey to South

Africa to study the above questions and some other scientific

matters there. During this expedition, which lasted nearly a

year, I visited both the east and west coasts and was able to

make many interesting observations.

The ‘‘seihval” from Saldanha Bay proved to be very different

from Rudolphi’s whale (B. borealis), and is described in this paper

as a new species, named after Johan Bryde, to whose generosity

IT am so much indebted. I have also to thank Capt, Andr.

Ingebrigtsen, Capt. P. J. T. Larsen, and Mr. Lars Iversen for

their kind help.

BALENOPTERA BRYDEI Olsen.

‘Tidens Tegn,’ November 12, 1912. (A Norwegian newspaper.)

Synopsis.

The average total length is 13 and the maximum nearly

15 metres. The shape is very elongated; the greatest height

13-14 per cent. of the total length. The distance from the tip of

the snout to the angle of the mouth is about 20 per cent. of the

total length. The flippers are slender and pointed, 8-10°6 per

cent. of the total length. The dorsal fin is small; its vertical

height 2-2-4 per cent. of the total length. The distance from

the tip of the snout to the anterior margin of the dorsal fin is

69-70 per cent. of the total length, occasionally shorter; the

minimum is 65 per cent. The distance from notch of flukes to

vent is about 27°2 per cent. of the total length; the number of

ventral furrows 42-54; they extend backwards to the umbilicus,

about 1°2 metres behind the tip of the flippers. The number of

baleen-plates (without the anterior rudimentary baleen) 250-280 ;

their greatest length nearly 0°50 metre. The bristles of the

baleen are very thick, long and stiff, not curling ; their colour is

erey. ‘The anterior baleen is asa rule more or less white; the

other greyish-black. The upper side of the flipper is bluish-

black, the under side grey. The colour of the body is bluish-

black above, with oblong light-coloured spots; the throat and

an area up to about 0°65 metre below the flippers on each side

are dark bluish-grey; the other parts of the under side are

white, more or less yellowish, often with a grey band across the

belly in front of the umbilicus.

General Characteristics and Measurements.

Balenoptera brydei is a comparatively small species, with an

average total length of about 13 metres, or occasionally a little

EGE

1913

ESF

BAL AGNOP TE RAY BRY DEI:

Es Zo & ISIS, Pl, CX.

BALAINOP TERA BRYDEM

9, lla. BALATNOPTERA BOREALIS.

Ls S, IGS, Pl, Cit,

IO, Wis, 13. IIR WIDISI.

14:

BALATINOPTERA BRY]

I 4. 3. Ils, Pil

DEI.

» Cox,

BRYDE’S WHALE. 1075

more. The longest specimen measured by me, and one of the

largest captured “by the Norwegian whalers in South Africa, was

a female from Durban measuring 14°95 metres. The female

seems to attain a slightly larger size than the male.

During my stay at the whaling stations in Durban and at

Saldanha’ Bay I was able to examine twelve specimens in ail ;

and the following table gives the measurements and some other

characteristics of three which showed comparatively great

differences.

Measurements of Balenoptera brydei (in metres).

a WE II Sho Geakllss ye ley ane

per | per | | per |

| | cent cent.) cent.

| Total length .. wse-+e+2-| 12°35 | 100 |] 13:07 | 100 | 14:95 | 100)

Tip of snout to anterior end of eye ene ae | 2741 |19°5 |) 2°40 )183)| 29 19-4

| s os angle of mouth .... ..| 2°50 202) 2°48 |18°9)) 2-9 |19°4

nS i anterior end of dorsal fin .| 8°55 |69°2) 85 (650) 105 '70:0

a S blowholes ..........:.. .| 185 |15:0 || 2°05 |15°6|, 2°36 |15°6

flippers (anterior margin) . 4:0 |32°3 || 40 |30°6]) 49 |32°7

| From notch of flukes to vent.. 3°38 |27°3 | 3°56 |272| 45 (301

a5 3 s hinder side of penis | |

OW WRUNG, co5cc0 ce: 4°43 1350 | 4°61 35°24 48. |32°1)}

Length of the flippers (from axilla) ......... 10 | 80] 1:39 |10:6|; 1:23 | 8:3}

Greatest breadth of the flippers............... | 0:33 | 2°6|| 086 | 2°7||- 0-40 | 2:6}

Breadth of the flippers at base ...............) ... || 028 | 21); 0°35 | 2:3

Vertical height of the dorsal fin ............ 0:30 | 2°4|) 0°30 | 2:°2)| 0:31 | 2:0

Miami eben ofeyer grat watercourse re vase GAO | cco |] OL Woo. |) CLO

Length of the eye-opening .....................} 0°75 |

' Length of the blowholes.. I oes Hh coo, ff OPED ip nce fh, | OS |

| Distance between the anterior ends of the|

blowholes 0:02 | 0:04

Distance between the posterio ior ends of the

Dow Oleserssee sees cacceteens sateen ide soe. I) OPER cos! I OPUS

VETER N OH (WOE AYROS Joo cco sacaHo cou sspacaces006b0 OD | oon |) Ole |) oon ii} Owes

Length of the penis or vulva.................. WEY ooo, |i LOS | | 018

rameter Of pemis|abeoase mes eer eee 0°20

Number of baleen-plates m each jaw (the |

rudimentary baleen not included) ...... jca. 260) ... jca. 250) ... |jca. 280)

Greatest length of the baleen-plates (with-

MUUG LORTSHAIOS). Gao sovodoseandssasocas soo asated ene O38a |e 0°32 | ... || 0°49

Greatest breadth of the longest baleen (at II

| thes base)ertiee see tere ee manawa se: 018}... | O14 {| 0:23

Greatest thickness of the longest baleen ...| 0°05 | ... a ca.0°03

| Length of the anterior (rudimentary) |

LOEW IEEIN | cee aanoy Sebban senate SoodapocuEt a ccaGaene OMS | Vee | Onk2

Length of the longest bristles (on the

Loneestabaleen)) meee aetna eae 0;20 |... I OWl4 |... (02

Number of furrows between the flippers...) 48 | ... 54 , 54

Height of the body at the flippers............ ca. 1°8 |14°7

Distance between the flippers on the ventral

RIGID -tedeaoeoctiag oan sen azeeceRan een aeee eae | mes son || 2RO

lt if

The shape of B. brydei, as of the other species of the genus

Balenoptera, varies somewhat, but as a rule it is remarkably

elongated (PI. CIX.), often even more so than in the Fin-whale

1076 MR. ORJAN OLSEN ON

(B. physalus). The greatest height of the body (at the anterior

end of the flippers) is usually about 14 per cent, of the total

length, occasionally less, and only in a single case (a male of

Saldanha Bay. 12.9.12) is it as much as 147 per cent. The

greatest breadth is almost the same as the height, or a httle less.

In five specimens measured by me in Saldanha Bay the pro-

portion of the greatest breadth (at the flippers) compared with

the total length was as follows :—12°8, 12:9, 14, 14:2 per cent.,

which gives an average breadth of 13-4 per cent. Ina single

EOI OF very thin male—the breadth was only 11°8 per

cent. The body is$therefore of a far less powerful build than in

B. borealis and B. acutorostrata, and resembles more that of the

Fin-whale (B. physalus). The hinder part of the body (from the

genital opening to the tail) is comparatively less compressed than

in B. borealis, but decreases gradually and slowly towards the

rear up to the root of the tail-flukes. \'The caudal part forms a

thin ridge above and below, which is at its sharpest along the

dorsal line (Pl. CXI. fig. 7). In order to facilitate the com-

parison between B brydei and the other species of the genus

Balenoptera, 1 have added a table showing the most remarkable

differences. J have not considered it necessary to mention the

Blue whale (4. musculus), as its size, dorsal fin, colour, and

many other characteristics make any confusion with B. brydei

impossible.

TABLE

showing the most notable differences between Balenoptera

brydei and the other species of the same genus (except

the Blue whale).

B. acuto- B. B. B.

rostrata. borealis. brydei. physalus.

Total length (maximum) ............. 10°3m, 163m. 15m. 24m.

Per cent. of total length.

From tip of snout to blowhole ...... 13°5 lbp 15°3 tise

ys . % angle of mouth 13-17 178 19°4-20:2 ea. 20

5 5 3 MOY DVRS sos cos oo oy 29°8 30°6--32°7 33

i dorsal fin ...... La 61-68 65-70 69-73

Height of body at the flippers ...... 20 18 14 sth

Height of the dorsal fin ......... 4-5'6 3°3-4:7 2-2°4 oe

Length of the flippers (from axilla) ibe 87-93 8-10'6 §°9-11:3

Greatest breadth of the flippers com- aslto35 aslto

pared to their length ............... a or 3°6 2°2 oes

Number of ventral furrows ......... ar iss 42-54 60-100

Number of baleen-plates oH ithout oy x oF 5

the rudimentary baleen)........ he AS) ABZBD | AHO 200

Greatest length of baleen in | adult “ ? ; ~ ;

specimens (without bristles) ...... ‘ DeaDiee 6 Oe ee Dee ia SRO a

Distance from notch of

flukes to anus.

Ventral furrows............

Colours, PO eae Re

Bristles of baleen .........

Food and habits

BRYDE'S WHALE.

B. borealis.

31:6 per cent. (of the total

length).

Extend to the middle of

the belly one foot behind

tip of flippers.

The throat always white:

the underside behind anus

bluish black, like the

back.

Unusually fine, short, curl-

ing and wool-like, white.

Always small crustaceans,

chiefly Calanide. Never

takes fish,

Blows as a rule only once

or twice during each visit

to the surface.

Migratory.

1077

B. brydei.

27°2 per cent. (of the total

length).

Extend to the umbilicus,

3°5 feet behind tip of

flippers.

The throat dark bluish-

grey: the underside

behind anus zhite,

sometimes yellowish or

whitish grey.

Very thick, long and stiff,

not curling, colour grey.

Fish. When it occasion-

ally takes crustaceans,

they are Euphausiide,

like the Blue whale.

Blows 5-6 times during

each visit to the surface

(ike the Fin- and Blue

whales).

Stationary.

The distance from the tip of the snout to the angle of the

mouth is, as we have seen, 19°4—20°2 per cent. of the total

length, about the sameas on the fin-whale, whereas in Rudolphi’s

whale it is only about 17°8 per cent. (according to R. Collett).

The tips of the snout and of the lower jaw are also far more

acute than in B. borealis, more resembling those of B. physalus.

The flippers are very slender and pointed, generally com-

paratively short, but varying in length from 8-10°6 per cent.

of the total length, and they are thus sometimes remarkably

longer than has ever been observed in B. borealis. ‘Their greatest

breadth (at the middle) compared to the length is as 1 to 2°2.

They differ in this respect very much from those of &. borealis,

where the proportions are as 1 to 3°5 or 3°6, and they are more

like the flippers of B. physalus. There is often a cut of about

an inch in breadth and depth in their posterior margin, near the

axilla. P

The dorsal fin is of very moderate size, and its vertical height

only about 2°2—2°4 per cent. of the totallength. Itisthus smaller

than is usual in the fin-whale, and very different from the large

dorsal of B. borealis and B. acutorostrata. Its length from the an-

terior to the posterior end is only slightly greater than its vertical

height. The front margin is convex, with a slightly curved apex,

and the hinder margin is deeply concave and often has a small

cut in it near the base. The point of the fin is very thin and

sharp. ‘The dorsal fin is placed noticeably further back than is

the case in Rudolphi’s whale—often where it would be on the

fin-whale. In twelve adult specimens examined by me the

distance from the tip of the snout to the anterior end of the

dorsal fin was in most cases 69-70 per cent. of the total length,

and in one instance only 63 per cent. In B. borealis the dorsal

1078 MR. ORJAN OLSEN ON

is usually placed anterior to the beginning of the last third of the

body (according to R. Collett).

The tail-flukes ave broad and comparatively somewhat shorter

than in B. borealis. The length of each fluke is from 10-11 per

cent. of the total length, or about the same as in the fin-whale. In

amale specimen, of which the total length was 13:07 metres, their

thickness at the base was about 0°20 metre and 0-12-0'14 metre

halfway between the base and the tip.

On each side of the tail, above and below, there is a thin ridge

which gradually becomes lower towards the rear, and disappears

between the flukes, about O°-10 metre from the end of the

body.

The eyes are comparatively larger than in the fin-whale, and

placed just above the angle of the mouth, in the same place as in

the fin-whale, but further back than in Bb. borealis.

The distance from the tip of the snout to the blowholes is about

15°3 per cent. of the total length. In adult specimens the length

of the blowholes is from 0°32-0-40 metre. They are placed

on a broad ridge on the head in two long furrows, which converge

very much towards the front. Between them there is a shorter

central furrow.

The ventral furrows run along the whole length of the lower

jaw and extend to the umbilicus, about 1:2 metres behind the

tip of the flippers (as in the fin- and blue whales). In B. borealis

they end about 0°30 metre beyond the tip of the flippers

(R. Collett). Above the long furrows there are 5-8 shorter ones

with a length of 0°30—-0°60 metre, and these terminate in the

axilla. In front of the flippers there are also a few short furrows

which extend forward into the angle of the mouth. On the belly,

in the area below the angle of the mouth, some shorter furrows

are as a rule wedged in amongst the others. Just in front of the

umbilicus, where the long furrows terminate, a number of quite

short furrows are also to be seen. The number of long furrows

varies from 42-54.

The anus is noticeably further to the rear than in B. borealis,

and the distance from the anus to the notch of the flukes is as a

rule about 27:2 per cent. of the total length.

The penis is rather big and with the vulva is placed ina generally

dark-coloured furrow 1-1°5 metres long, about two-thirds of

which run anterior to the genital opening, and about one-third

behind.

A varying number of whitish-grey oblong spots, about 7-8 cms.

in length and about 3 ems.in breadth, are distributed over the

whole body. Along the centre of their long axis there runs

usually a dark line, from which fine radii of similar lines run in all

directions. These patches probably show places once attacked by

parasites. In addition to these whitish spots a number of fresh

wounds, with a length of as much as 10 cms. and 3-4 cms. deep,

can be seen in most cases. They are marks caused by parasites—

generally a Pennella.

BRYDE’S WHALE. 1079

The majority of specimens of Bryde’s whale are very thin, and

their stratum of fat has an average thickness of only 4—5 ems.

In a single instance—that of an old and probably diseased female

specimen from Durban—the thickness was only about 3°5 cms.

Some big specimens, especially females with foetus, may be quite

fat and their stratum of fat may attain a thickness of as much

as 7 ems., at any rate on the dorsal side. Specimens as fat as

this may yield 15 or occasionally nearly 20 barrels of oil, while

the common yield of this species is only 6-8 barrels.

Baleen.

During my stay in 8. Africa I was able to examine the baleen

of twelve adult specimens, and also a large quantity of whale-

bone which lay heaped up outside Mr. Bryde’s whaling station in

Saldanha Bay.

The baleen of B. brydei is very distinctive of this species, and

compared with the size of the whale, very small (see P]. CXII.

fig. 11 B). The longest baleen-plates measured by me, and from

a female specimen of nearly 15 metres in length, were only

0-49 metre long (the bristles not included), while baleen from a

Rudolphis whale of the same size attains a length of about

0-70 metre. The baleen differs remarkably in shape from that

of B. borealis; whereas the baleen-plates of 5. borealis are very

long and slender, those of Bb. brydet are comparatively very broad

and curve inwards along the inner margin. The comparative

proportions of the breadth of the largest baleen-plates at their

base and their length were in four specimens of B. brydei:—

43, 43-1, 46-9, and 47 per cent., which gives an average of 45 per

cent. In B. borealis the same proportion is about 27:2 per cent.

(see Pl. CXIT. fig. 11 4). The baleen of B. brydei seems also to be

comparatively somewhat thicker and more strongly built than in

B. borealis. The number of plates in twelve specimens examined

by me was, as a rule, about 260 in each jaw, though the number

varied between 250 and 280. But if the rudimentary plates are

included, the number may be about 350. ‘The number of baleen-

plates in B. brydei is remarkably small ; for Rudolphi’s whale the

number (without the rudimentary baleen) is from 320 to 340,

and in the fin-whale from 360 to 400.

The rudimentary plates along the tip of the upper Jaw are not

compressed, and attain a length of 0-10-0°15 metre; they are

numerous and difficult to count. The first compressed plate

is seen about 0:25-0:30 metre behind the tip of the snout.

There is also some rudimentary baleen above the angle of the

mouth.

The bristles (Pl. CXII. fig. 10) are longer than in B. borealis

and thick and stiff, not curling, and on the whole of very strong

construction—even comparatively stronger than in the fin-whale.

Their average thickness is about 1 mm.; a little more at the

distal end of the baleen, and perhaps a little less at the base.

1080 MR. ORJIAN OLSEN ON

B. brydei differs very much in this respect from B. borealis,

and it would hardly be possible with its imperfect straining

apparatus to keep back such small crustaceans as the Calanide,

which form the chief food of B. borealis.

The colour of the baleen in the anterior part of the jaws,

and about 0°70 metre backwards from the tip of the snout, is

as a rule more or less white, sometimes perfectly white, but

more frequently with grey stripes; further back it is greyish

black, and after death perfectly black. The bristles are grey,

whitish grey or yellowish in the anterior part of the mouth.

The colour of the baleen is on the whole rather similar to that of

the fin-whale, and the whalers told me that in some cases they

had even seen the white colour asymmetrically placed, as in the

fin-whale. The baleen might in one jaw be white over a com-

paratively large area, and in the other jaw fairly uniformly dark-

coloured. A male caught in Saldanha Bay, March 5, 1913, had

only dark-coloured baleen. In this respect too B. brydei differs

considerably from B. borealis, in which the whalebone is only

rarely white-mottled and as a rule uniformly black with white

bristles.

In connection with the description of the whalebone, it is

worth mentioning that some time after I had given the first

preliminary report on 5. brydei in a Norwegian newspaper,

Mr. Bryde wrote to me saying that he could now understand the

reason for a law-suit in which he had been implicated some time

before. He had sold a consignment of baleen from his factories

inS. Africa which his agents there considered to be “ seihval ”

(B. borealis). The buyers, however, complained and returned

the whalebone, saying that 1t was not from #&. borealis but from

another species of whale. The result was a law-suit in which

the authorities who examined the whalebone gave judgment

in favour of the buyers, and Mr. Bryde was obliged to pay a

large fine and take back the whole consignment.

The Norwegian whalers in South Africa said too that this whale

(B. brydet) was not the proper “ seihval”; but as it was most

like the “seihval” in size and colour, they generally called it

that.

Hairy covering.

In all the specimens examined by me (except in the case of

an old and probably diseased female specimen from Durban, on

which I could see no hairs at all), I found two rows of hairs on

the tip of the lower jaw, with twelve hairs in each row, thus:

Mouth.

BRYDE’S WHALE. 1081

The distance between these rows in adult specimens was about

18 mm., or a little more than between the successive hairs

in each row. Outside the upper end of these rows were two

other hairs on each side, placed at the same distance from the

long rows as between the rows themselves. I noticed no hairs on

the upper jaw.

The number of hairs is thus 28 in all. On two foetuses, with

a length of 3°78 and 0:935 metres, the hairy covering was as well

developed as in adult specimens. The length of the hairs was

from 5-10 mm.

Colowr.

The upper side of the body—above the flippers—is bluish black,

and in colour much resembles the fin-whale. After death the

colour on this part becomes almost entirely black. On the sides

of the body the colour becomes gradually lighter, and there is no

sharply detined border between the dark colour of the upper

side and the white belly. The flippers are of the same colour as

the back above, or sometimes darker; below they are grey,

occasionally whitish grey. The under side of the head, and a

band extending to about 0°70 metre below the flippers and

backwards to their tip are bluish grey. The centre of the belly

is a more or less yellowish white. In the dark area below the

flippers some isolated white spots may sometimes be seen ; similar

white patches are occasionally distributed along the centre of the

under side of the lower jaw, and more rarely a small white line

runs out from the white area on the belly to within one metre or

less behind the tip of the lower jaw. On the border between the

dark colour of the throat and the white belly there are often a

number of bluish grey patches or stripes, and these are more

thickly sprinkled towards the front, and then gradually form a

more uniform dark colour in front of the angle of the mouth.

Between the tip of the flippers and the umbilicus—at the posterior

end of the long furrows—the white colour of the under side is

generally interrupted by a bluish grey belt running across the

stomach. This belt, which has a breadth varying between 0-70

and 1°5 metres, is often formed by a large number of smaller

and larger patches or stripes, not of uniform colour; in other

cases it 1s very little developed.

Considerable variations occur in the breadth, size, and shape of

the white area on the belly.

From the genital opening to the tip of the tail-flukes the

whole under side is dirty white, sometimes yellowish or whitish

ervey. This white area begins above the genital opening at the

middle line of the sides of the body, and in some specimens is

further back and becomes gradually narrower.

From the tip of the upper jaw, near the roots of the baleen,

sometimes a white stripe runs backwards, and the length of

this is about 0°5 metre and the breadth 5-7 ems.

1082 MR. GRIAN OLSEN ON

The dark patehes which are sometimes found on the under side

are generally situated in the long furrows, or in the furrow at the

genital opening.

Occurrence and Capture.

Balenoptera brydei seems to be common everywhere along the

south coast of Africa. It is especially so in Saldanha Bay,

though also in Mossel Bay and near the Cape of Good Hope this

species is captured in rather large numbers, and is often observed

from whaling steamers going along the $8. African coast.

In Darban &. brydet is comparatively rare, but 1s occasionally

to be seen at all seasons of the year. The total number captured

from three whaling stations on the Natal Bluff during the season

1912 was only sixteen, of which one (captured early in 1 November)

was only about 8 metres long, and not fully developed. In 1911

six specimens were captured from Myr. Bryde’s station in Durban.

Of these two were taken on June 5th, one on the 14th, and one

on the 15th of the same month, one on October 10th, and one on

November 7th. In 1912 the following were caught from the

same station:—one on June 4th, one on July 18th, one on

Aug. 19th, and one on Nov. loth. Some specimens were seen

but not pursued because, as the whalers told me, they were

very shy and difficult to catch, and at the same time their value

was comparatively small. At Mr. Bryde’s whaling station

near Inhambane, Portuguese EH. Africa (23° 30'S. 1.), B. brydet

was very seldom seen, and only a single specimen of it was

captured. Further to the north at Bazaruto Is. (21° 45'8.1.),

where the floating factory ‘‘ Mangoro” was stationed for two

months, B. brydet was never seen. Bryde’s whale therefore

seems to be a southern species, avoiding the warm waters of the

tropics, and it is therefore rare on the east coast where the hot

Mozambique current runs southward.

Almost all the specimens of 2B. brydet captured in Saldanha

Bay (33° 8.1.), were found between this bay and Capetown, and

the whalers told me that this species was much more rarely seen

north of Saldanha. This fact 1s quite in accordance with the

experiences of Mr. Green, the manager of Storm Bull’s whaling

station in Walfisch Bay. He considered the “‘seihval” to be very

rare along the coast of German S8.W. Africa, and he had not

observed this species at all in Walfisch Bay.

I was somewhat surprised therefore when Capt. Andr. Inge-

brigtsen, who had been stationed with a floating factory at

Porto Alexandre, Benguela (15° 45'S. 1.) for a couple of years,

told me that he had observed rather a large number of ‘ seihval ”

there. The same statement was made by Capt. L. Fredriksen,

who had seen many specimens of this whale outside Lobito Bay,

and single ones as far north as 7° 8.1. But when we remember

that a cold antarctic current runs along the west coast of Africa

BRYDE’S WHALE. 1083

far to the north, it is not so surprising that B. brydei may be able

to live outside the tropical parts of the coast. The conditions of

existence, however, for this species cannot be very good there,

because the 3 or 4 specimens captured (during nearly three years)

outside Pt. Alexandre were all unusually thin and did not yield

more than 2-3 barrels of oil, or sometimes even less. All the speci-

mens seen at this place were in very bad condition, with numerous

wounds from parasites over the whole body, and so thin that the

whalers did not care to pursue them. They were never seen

accompanied by young ones outside Pt. Alexandre, and they were

all lying almost motionless and did not make any efforts to escape

from the steamer. Unfortunately not a single specimen has been

examined from this part of the African coast, and very few were

captured, because the whalers always preferred to take the more

valuable “humpback” whales, which were to be found in the same

locality. It has therefore not yet been proved whether the

‘“‘seihval” from the coast of tropical W. Africa really is Baleno-

ptera brydet.

Capt. Andr. Ingebrigtsen also told me that the ‘“seihval ” out-

side Pt. Alexandre appeared in these waters every year about the

middle of June and disappeared again about September or

October.

Tf this whale follows the same route as the ‘“ humpback”

generally does—along the coast—we might expect to see migrating

specimens pass Saldanha Bay, and to find a greater number of

them there in the spring, October-November, but this is not the

case. It is probable therefore that they choose another route,

perhaps further out to sea, where the current is stronger and the

water cooler.

It is not impossible that some of the ‘“sei”-whales from

Portuguese W. Africa may have been Rudolphi’s whale (B. bore-

alis), a typical specimen of which was captured outside Saldanha

Bay in November 1912. This is the first S. African specimen of

B. borealis which has been reported further south than off Cape

Blanco (20° 45' N. 1.). It was easily recognized as the proper

““seihval” by the Norwegian whalers; and Capt. P. J. Larsen

kindly presented me with a quantity of the baleen, which was

quite distinctive of B. borealis (Pl. XCII. fig. 9).

As I have mentioned above, Saldanha Bay is the locality where

the greatest number of 5. brydei are captured. The whaling

trade was started there in 1910, and only twenty-four specimens

of the ‘“ seihval” were captured in the first year, and all of

them in the spring. The first three specimens were caught on

March 9th and 29th, and on April Ist. Five were captured

from the 20th to the 30th of April, four from the Ist to the

10th of May, six from the 10th to the 20th of May, and six from

the 20th to the 30th of May. After that time the “‘ humpback ”

was found at this spot in large numbers.

The following list shows the number of ‘ seihval” captured at

1084 MR. GRIAN OLSEN ON

Mr. Bryde’s station in Saldanha Bay during the season of 1911

(with two whaling steamers), and in the season of 1912 (with three

whaling steamers).

April 10-20

April 20-30

May 1-10

May 10-20

May 20-30

June 1-10

June 10-20

June 20-30

July 1-10

July 10-20

July 20-30

Sept. 10-20

Sept. 20-30

Oct. 1-10

Oct. 10-20

Oct. 20-30

Nov. 1-10

Nov. 10-20

Nov. 20-30

Dec. 1-10

Dec, 10-20

—

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The table shows that the majority of 6. brydei are captured in

the autumn (April and May), and at this time this species also

seems to be the most numerous outside Saldanha Bay, as well as

Durban. But we see also that 1t is captured in greater or less

numbers during the whole season, and it is seen outside Saldanha

Bay all the year round.

The difference between the real number of B. brydei in the

autumn and at other times of the year is in fact not so great as

the capture seems to indicate, because the whalers always prefer

to take the more valuable “ humpback” whale when this species

appears on the spot in May. In 1912 an unusually small number

of ‘““humpbacks” was seen, and the trade depended on the

« seihval” to a greater extent than before.

In 1913 the first specimen was caught on March 5, and from

BRYDE'S WHALE. 1085

that date until I left Saldanha Bay a fortnight later, the total

eatch of 2. brydei consisted of eleven specimens. Almost all of

these specimens were found unusually far (40-70 miles) from

the sea-shore, surrounded by large numbers of crustaceans, Eu-

phausiide. As a rule the whales are to be found quite close

to the coast, and it is probably the richness of crustaceans

further out to sea that has tempted them to go out there. It

is worth mentioning that these crustaceans were Euphausiide,

which the Blue whale prefers, and not Calanide, the chief food

of Rudolphi’s whale.

Food.

The food of B. brydei consists chiefly of fish, apparently

-usually a variety of herring which is often found in large

quantities (many hectolitres) in its stomach. This, for instance,

was the case with a male examined by me in Saldanha Bay on

September 12th, 1912. It sometimes takes a species of mackerel

one foot or more in length, and in Durban more than a hectolitre

of this fish has been found in its stomach.

B. brydet is very voracious on the whole—more so than any

other species of its genus. As an illustration of this, Capt. L.

Fredriksen told me that he had many times seen it hunting

among large crowds of small sharks, and that he had found

sharks of a length of more than two feet in its stomach. <A case

from Saldanha Bay which was related by Captain Christoffersen,

and mentioned by many others, was most astonishing: they had

found there no less than 15 large penguins (Spheniscus demersus)

and “malagass” (Sula capensis) in its stomach. These birds, the

moment the whale reached the surface of the water, had probably

dived down into its open mouth endeavouring to catch fish in

that abundant hunting-ground, and had thus themselves been

involuntarily captured by the whale. In similar cases observed

among B. borealis, the birds have always been spat out of the

mouth again, and it seems hardly possible that such large birds

as those mentioned could be swallowed by this species at all.

L. brydet is not dependent for food on the occurrence of

crustaceans in the sea and so does not migrate, but is generally

seen very close to the coast pursuing fish.

Like the fin-whale, B. brydei cecasionally takes crustaceans and

then of a larger kind, the Kuphausiide.

Liology, ete.

In its biology B. brydei is most like B. acutorostrata and

BL. physalus, and when the Norwegian whalers started their trade

in South Africa they were doubtful as to whether they should

consider this species to be a fin-whale or a “sei”-whale (B.

borealis). In Durban many of them preferred to call it a

Proc. Zoou. Soc.—1913, No. LX XII. 72

1086 MR. ORJAN OLSEN ON

‘“sildehval” (herring whale), because it hunted herrings. But as

the colour and size were most like that of the “seihval,” they

generally called it so, although they knew that it was not the

proper ‘‘seihval.”

In Saldanha Bay, where the largest numbers were captured, it

was often called “bastard” or ‘“seihval bastard,” and considered

to be a hybrid between the fin-whale and the ‘seihval,” because

it had the baleen of the first species, but some other characteristics

of the latter. Besides this ‘“ bastard,” the whalers in Saldanha

Bay told me about another variety of “ seihval” with exceedingly

dark-coloured baleen. It was of somewhat smaller size than the

“bastard,” quicker in its movements, dived deeper down in the

sea, and was more difficult to catch. It was seen in approxi-

mately the same numbers all the year round, and in the same

localities as the “bastard,” but seemed to occur in greater

numbers northward of Saldanha Bay than the latter, as proved

by Capt. Christoffersen, who had captured it many times

outside Paternoster Bay, When the whaling steamers went to

Capetown for coaling, these two varieties were often seen in

greater numbers together, and the whalers picked out the

“bastard” as more valuable and easier to catch.

Unfortunately, during my stay at Saldanha Bay, not a single

specimen of this smaller variety was captured, and it is therefore

difficult to say anything about them, but I consider it probable

that they are only younger specimens of B. brydei.

B. brydet is as a rule to be found very close to the coast,

generally in large numbers, and it is comparatively rarely found

further out to sea. The whalers in Saldanha Bay often found it

pursuing fish only a few hundred yards from the beach between

Robben or Dassen Island and the continent. In Durban it was

also seen close to the coast (occasionally as far out as 15 miles

from shore), but never in company with the small ‘ minkehval”

(probably B. acutorostrata, ov a subspecies closely akin) nor with

other whales. As mentioned above it has occasionally been found

60-70 miles from the coast feeding upon Kuphauside.

It is most like the fin-whale in its manner of swimming, and

can easily be distinguished from the small “‘minkehval.” When

coming to the surface to breathe, it may be seen under the water

for some time before it appears on the surface. During each

such visit it blows four or five times (like the fin- and blue

whales, whereas Rudolphi’s whale blows only once or twice)

before diving down into deep water again. Between these four

or five blows it does not go too deep down to be seen from the

ship and followed. It blows far more strongly than does

B. acutorostrata, and its breath smells worse than that of the

other whales. In Durban it was observed to remain for an

unusually long time under water between each visit to the

surface. It is not seen to take crustaceans in the same peculiar

way as Rudolphi’s whale (B. borealis). When shot it often swims

BRYDE’S WHALE. 1087

round in circles very quickly and gets the line wound round it,

and is on the whole difhicult to manage. Large females are less

ditiicult to shoot than the others, as they are not so quick in

their movements.

Gravid females do not seem to be followed by their males as is

the case with the humpback whale. For instance, two females

captured in Saldanha Bay, March 12, 1913, and with feetuses

measuring 0°935 and 3°78 metres, were swimming about quite

alone. Some males captured at this time were also solitary.

They are gravid at very different times, and females were found

with foetuses of different sizes both in Saldanha Bay and Durban,

at the same time as others were followed by young with a length

of about 6-7 metres.

It may be mentioned as a curious fact that almost all the

“sei ”-whales captured in Saldanha Bay from March 5-13, 1913,

went southwards when the whaling steamers began to hunt

them.

The flesh of B. brydet contains less oil than does that of other

species of its genus, with the exception perhaps of B. borealis.

When fresh and served like beef-steak it tastes quite good, as I

had an opportunity of proving when at Saldanha Bay.

The Fetus.

About an equal number of each sex of B. brydei seem to be

eaught. Many of the females were gravid, and, as mentioned

above, with fetuses of very different sizes :—ée.g., in two gravid

females examined by me in Saldanha Bay, March 12, “1913

the fetuses had a length of 0°935 and 3°78 metres (PI. CX LIT.

figs. 12, 13). The mammee in both these specimens were very

well developed, and, -when they were dragged out of the water,

about a half-gallon of mucus ran out of their genital openings,

as has sometimes been observed shortly after the birth of the

young in other whales. The largest foetus was full-grown, and

fell out when the men began to remove the stratum of fat from

the whale.

The following table shows some measurements of the two

foetuses.

In the larger foetus the penis was placed in a furrow which

ran backwards from the umbilicus for about 12 cm. The

baleen had just appeared and was thick and soft, with a length

of about 3 cm. The bristles had an average length of 6—

8 mm. and were generally only to be seen at the base of the

baleen. The anterior and posterior baleen only appeared as

small ridges. The dorsal fin was bent down to the back on the

left side, both the apex and the tip being curved to an unusual

extent, and the hinder margin deeply concave. The tail-flukes

were strongly bent together towards the median line of the body.

The colour above was a uniform bluish black, and whitish grey

(ie

1088 MR. ORJAN OLSEN ON

| Male. Female.

per cent. “per cent.

| Total length .. a |) 3°78 100 || 0:935 | 100

| Tip of snout to anterior end of eye .... | 063 1EiGH OMS AS ae)

e es angle of mouth | 0°70 18°5 | Ong ee |

ae Fe anterior end of dorsal fin . 4 2°47 65°33 || -0°655 70:0

a » blowholes .. seocasuosae| OE mies) | Os} 13:9

o flippers (anterior margin). 1:2 31:7 || +030 32:0

From notch of flukes to vent ...... ee esl ee OAs 27°5 | 0°255 O72 |

vulva or penis ...| 1°35 Sry |, ORS, Peril |

Leneth of flipper .. sib sgaasdaongagaa scale Oy if a IOES}) I) OOe 66 |

| Greatest breadth of flipper Per eee GOLD Meson Oe: 32 |

| Breadth of flipper at its base...... ue teddies wna 27 || 0042 4-4 |

| Vertical height of dorsal fin ..................; 013° | 34° |! 0:026 iy

Length of dorsal fin at its base............. le hat ae O38 4:0

| WRaieaeIEe OP GO uo sudenscassadvegaucocasooieonan) OOH — If cu, | 0:023 |

HIG EEAN One ChVSAOHANUE?. cgassncsvarcasouecsseoal| , lece | his coal yy | ORO) |

| Length of blowholes ................. pamecaselht s OPLON Ne ORO. | |

| Distance between the anterior end of | | |

| blowholeste ea O23 earn O00

| 3 5% posterior end of |

blowholes ....... 0°073 Sa 0°025

| eres ape HITE Moa ease onnsoanromtecesvacoocea| OO2 iI) 0°007

| Length of penis or TUINGED. de sgncoueeeomcoosbeelh OO Keseene 0015

Diameter of penis at its base........ i) OXORs |

Number of furrows between the flippers... 48 i 42

| Distance from penis te the umbilicus ......) 0°38

From tip of lower jaw to anterior margin

of umbilicus ...... 5904s REE: Fae a eee ae O51 54:5

Diameter of the umbilicus cc t OOOu ale pe 0:03

Breadth of tail-flukes at their base. sceooonsel), O82 7} |

Distance between the tips of flukes ....... OAR |) 3: 0:20 OMB |

Distance from notch of flukes to tip of |

flukes .. sbdedeosnl!| OFSts)

Breadth of the body ‘at the flippers See BO tN fe OA elais

Height of the body at base of flippers ...... oe NY i OPN TPR

Be 33 at tip of flippers .... 630 13°8

: : ati iMeleyesmen.. sence At 13°2 |

3 rs at the dorsal fin ...... eae EWR ORO |

5 55 at the tail, just in | |

| front of flukes ...... Bedale vit barn BEG |

everywhere on the under side. The change from the dark to

the light colour along the sides was ames ne and no definite

outline could be seen. There was a more ereyish area just

behind the eyes on each side.

No baleen was to be seen on the smaller fcetus, and the whole

body was of a homogeneous, reddish brown colour.

Abnormal Specimens.

A large female specimen with a total length of 14:95 metres,

captur wel in Durban, November 15th, 1912, was very like the

fin-whale in colour, Rowethe flippers were white below and the

colour asymmetrical’ as it is in the fin-whale. The lower jaw and

the distal part of the upper jaw were white on the right side, while

those parts on the left side were bluish black. ‘On ‘ae, right

side there was also a greater number of white baleen-plates ‘shar

BRYDE’S WHALE. 1089

on the Jeft. The furrows were bluish black in a belt across the

under side of the head and backwards to the flippers, but only a

comparatively small part of the throat on the left side was

uniformly dark-coloured. No hairs were to be seen on the tip

of the lower jaw. | This specimen was unusually slender and thin

and probably was very old and diseased. The stratum of fat was

only about 3°5 mm. thick; and there were numerous wounds

distributed over the whole body (Pl. CXIII. fig. 14). These

wounds were more or less inflamed and partly filled up with

mortifying fat, and in some of them a Pennella was found. This

specimen was lying quite motionless on the surface when it was

observed from the whaling steamer, and was thus very easily

taken.

The whalers from Durban told me that on one occasion

they had caught a whale of this species, which on the whole

under side of the body was of a colour much resembling that of

the blue whale (4. musculus); but when it was taken it proved

to be a “ herring ” whale (B. brydei).

Parasites.

Parasites are comparatively rare in this species ; an old female

(Durban, November 15th, 1912) had, as mentioned above, dis-

tributed over the whole body a large number of oval-shaped

wounds, with a length of as much as 10 cm., and a depth of

2-4 em. (PI. CXIII. fig. 14). Almost ali of these wounds were

partly filled with mortifying fat and had been caused by a large

Pennella, specimens of which were found in some of them. . A

smaller number of these wounds was found on this species at

Saldanha Bay, but they were very numerous in the few old

and apparently diseased specimens taken at Porto Alexandre.

A species of Myxinoid sometimes makes similar wounds in

Bryde’s whale, but I did not obtain specimens of them because

they always leave the whale when it is dragged out of the water.

I do not know whether they are to be found on the whale when

alive, or only after its death.

EXPLANATION OF THE PLATES.

Phare CIX.

Balenoptera brydei.

Fig. 1. Side view of male.

2. Lower surface of male.

Prat CX,

Balenoptera brydei.

3. Male specimen, showing the dorsal side and anterior end of the body.

Saldanha Bay, 12.9.12.

4, Same specimen as fig. 3, from the ventral side.

Female specimen, from the dorsal side. Saldanha Bay, 12.3.13.

Cee

ise)

1090 THE SECRETARY ON ADDITIONS TO. THE MENAGERIE.

Prare CXI.

Balenoptera brydei.

Fig.

Head of a large, but very thin and apparently sick, female specimen, with

numerous wounds from parasites (Pennella). Durban, 15.11.12.

. Caudal portion, with the dorsal fin, of a male specimen.

. Female specimen, from the ventral side. Saldanha Bay, 12.3.13.

mnt

Pratre CXII.

Fig. 9. Bristles of baleen from Balenoptera borealis, captured outside Saldanha

Bay, —.11.12.

10. Bristles of baleen from B. brydei, captured outside Saldanha Bay,

—.11.12.

11. Baleen from adult specimens of (A) B. borealis, and (B) B. brydei.

Prate CXIII.

Balenoptera brydei.

Fig. 12. Foetus, 0°935 m. in length. Saldanha Bay, 12.3.13.

13. Foetus, 3°78 m. in length. Saldanha Bay, 12.3.13.

14. Portion of body of same specimen as fig. 6, showing wounds from parasites

(Pennella). Durban, 15.11.12.

EXHIBITIONS AND NOTICES.

October 28, 1913.

Prof. KE. A. Mincuty, M.A., F.R.S., F.Z.8. Vice-President,

in the Chair.

THe Secrerary read the following report on the additions that

had been made to the Society’s Menagerie during the months of

May, June, July, August, and September, 1913 :—

May.

The registered additions to the Society’s Menagerie during the

month of May were 268 in number. Of these, 133 were acquired

by presentation, 52 by purchase, 31 were received on deposit, 13 in

exchange, and 39 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 185,

Amongst the additions special attention may be directed to :—

2 White-bearded Gnus (Connochetes albojubatus), from

British East Africa, new to the Collection, received in exchange

on May 19th.

2 Chimpanzees (Anthropopithecus troglodytes), 1 Anubis Baboon

(Papio anubis), from West Africa, and several small Monkeys,

presented by W. O. Danckwerts, Esq., K.C., F.Z.S., on May 2nd,

28th, and 31st.

1 Crowned Duiker (Sylvicapra coronata), presented by Capt.

C. H. Armitage, C.M.G., D.8.0., F.Z.S., on May 4th.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 1091

1 Poppig’s Woolly Monkey (Lagothrix poppigi) and 1 Ocelot

(Felis pardalis), from Manaos, presented by Albert Reymond,

Esq., on May 28th and 20th.

2 Naked-tailed Mice (Uromys bruijnii), new to the Collection,

and 2 Sclater’s Cassowaries (Casuarius sclateri), from 8. Dutch

New Guinea, presented by A. F. R. Wollaston, Esq., and C. Boden

Kloss, Esq., on May 3st.

2 Himalayan Yellow-backed Sun-birds (#thopyga seherie) and

1 Nepalese Bullfinch (Pyrrhula nepalensis), from the Himalayas,

new to the Collection, purchased on May 29th.

2 Swainson’s Blue Jays (Aphelocoma sordida), from Mexico,

new to the Collection, purchased on May 26th.

1 Chestnut-faced Barn-Owl (Strix castanops), from Tasmania,

new to the Collection, purchased on May 10th.

4 Mexican Blue Herons (Ardea herodias lessont) and 2 Northern

Boatbills (Cancroma zeledoni), from Mexico, new to the Collection,

received in exchange on May 28th.

1 Gentoo Penguin (Pygosceles papua), from the Falkland

Islands, presented by the Rt. Hon. L. Vernon Harcourt, M.P.,

F.Z.S., on May 6th.

3 Smooth-backed Zonures (Psewdocordylus microlepidotus), from

S. Africa, new to the Collection, deposited on May 7th.

1 East African Scorpion (Lepreus tricolor), trom Lake Nyanza,

new to the Collection, presented by C. B. Blencowe, Esq., on

May 20th.

JUNE.

The registered additions to the Society’s Menagerie during

the month of June were 295 in number. Of these 144 were

acquired by presentation, 35 by purchase, 56 were received on

deposit, 15 in exchange, and 45 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 189.

Amongst the additions special attention may be directed to :—

1 Pudu Deer (Pudu pudu), from Chili, and 2 Patagonian

Cavies (Dolichotis magellanicus), from Patagonia, presented by

Albert Pam, Esq., F.Z.8., on June 2nd.

3 Pumas (felis concolor), born in the Menagerie on June 13th.

2 Canadian Beavers (Castor canadensis), from Canada, received

in exchange on June 5th.

1 Savanna Sparrow (Passerculus savanna), from North-East

America, new to the Collection, purchased on June 18th.

1 Ceylon Mynah (Acridotheres melanosternus), new to the

Collection, presented by Dr. P. H. Bahr, F.Z.S., on June 29th.

1 Golden-fronted Woodpecker (Melanerpes flavifrons), from

Brazil, new to the Collection, purchased on June Oth.

1 Condor (Sarcorhamphus gryphus), from Chili, presented by

Albert Pam, Esq., F.Z.S., on June 2nd.

1092 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

2 Crested Screamers (Chauna cristata), bred in the Menagerie

on June 19th:

2 Sun Bitterns (Hurypyga helias), from South America,

purchased on June 16th.

1 Kagu (Rhinochetus jubatus), from New Caledonia, presented

by the Marquess of Tavistock, F.Z.8., on June 27th.

2 Spiny-tailed Skinks (Zgernia depressa), from Australia, new

to the Collection, purchased on June 10th.

A collection of Snakes from Sierra Leone including 3 Sooty

Snakes (Boodon fuliginosus), new to the Collection, presented by

Guy Aylmer, Hsq., F.Z.S., on June 18th.

A collection of Snakes from India including 1 Forsten’s Tree-

Snake (Dipsas jforstenii), new to the Collection, received in

exchange on June 2nd.

2 Gopher Frogs (Rana capito), from North America, new to

the Collection, received in exchange on June 2nd.

JULY.

The registered additions to the Society's Menagerie during the

month of July were 279 in number. Of these 119 were acquired

by presentation, 46 by purchase, 21 were received on deposit, 37

in exchange, and 56 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 250.

Among the additions special attention may be directed to :—

1 Orang-Utan (Simia satyrus), from Borneo, presented by

Marcus 8. Abrahams, Esq., ¥.Z.S., on July 24th.

1 Small-clawed Otter (Lutra leptonyx), from India, presented

by H. R. Taylor, Esq., F.Z.8., on July 25th.

1 Long-tailed Mouse (Orizomys longicaudata), 2 HKlegant

Opossums (Marmosa elegans), from Valparaiso, new to the

Collection, presented by Walter Goodfellow, Esq., F.Z.S., on

July 8th.

4 Capybaras (Hydrocherus hydrocherus), from Fray Bentos,

Uruguay, presented by the Liebig’s Extract of Meat Co., Ltd.,

on July Sth.

1 Grysbok (Nototragus melanotis), from Constantia, Cape

Colony, presented by W. Lester, Esq., on July 29th.

2 Australian Flower-peckers (Dicewm hirundinaceum), from

Australia, new to the Collection, presented by G. A. Heumann,

Esq., on July 5th.

1 Shoebill (Saleniceps rev), from the Sudan, presented by

Lt.-Gen. Sir Francis R. Wingate, G.C.V.O., K.C.M.G., K.C.B.,

D.S.O., H.M.Z.8., on July 15th.

3 Seed-Snipe (Thinocorys rumicivorus), from Chili, new to the

Collection, received in exchange on July 24th.

2 Spinose Fence-Lizards (Sceloporus spinosus), 1 Graham’s

Snake (Zamenis grahanz), from N. America, new to the Collection,

purchased on July 22nd.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 1083

1 Bird-eating Spider (Citharoscelus kochit) 2, 1 Bird-eating

Spider (Paraphysa manicata) 3, from Valparaiso, new to the

Jollection, presented by Walter Goodfellow, Hsq., F.Z.S., on

July 8th.

A Collection of Millipedes (Archispirostreptus nitidus), from

Trinidad, new to the Coliection, purchased on July 11th.

AUGUST.

The registered additions to the Society’s Menagerie during the

month of August were 201 in number. Of these 122 were

acquired by presentation, 15 by purchase, 14 were received on

deposit, 7 in exchange, and 45 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 231.

Among the additions special attention may be directed to :—

1 Fettered Cat (Felis ocreata) and 1 Cheetah (Cynwiwrus

jubatus), presented by Dr. R. KE. Drake-Brockman, F.Z.8.,

1 Caracal (felis caracal) and 1 Cheetah (Cynelurus jubatus),

presented by Capt. J. L. Berne, and 2 Cheetahs (Cynelurus

jubatus), presented by Arnold Hodson, Hsq., from Berbera, on

* August 8th.

2 Indian Gazelles (Gazella bennetti), from Karachi, presented by

A. C. Cooke, F.Z.5., on August 11th.

1 Barred Warbler (Sylvia nisoria), from Kurope, new to the

Collection, presented by R. Pearce Page, Esq., F.Z.8., on August

Sth.

6 West-African Quail-Finches (Ortygcespiza atricollis), from

Gambia, new to the Collection, presented by Dr. E. Hopkinson,

D.S.0O., F.Z.S., on August Ist.

1 Spot-billed Toucanet (Selenidera maculirostris), bred in the

Menagerie on August 11th.

2 Bornean Fireback Pheasants (Lophwra nobilis), bred in the

Menagerie on August 17th.

A collection of rare Reptiles from Sierra Leone, including a

Black-collared Cobra (Waia nigricollis), a Blanding’s Snake

(Dipsadomorphus blandingii), and 3 Green. Night Snakes

(Dipsadoboa unicclor), all new to the Collection, presented by

Guy Aylmer, Esq., F.Z.S., on August 2nd.

1 Hamadryad (Waia bungarus), from India, received in

exchange on August Ist.

6 Pigmy Rattlesnakes (Sistrurws miliarius), born in the

Menagerie on August 19th.

4 Miiller’s Xenopus (Xenopus miillert), from West Africa,

new to the Collection, purchased on August 7th.

1 Electric Cat-Fish (Malapterurus electricus), from Tropical

Africa, new to the Collection, purchased on*August 28th.

1 Scorpion (Bathus hottentotta), from Sierra Leone, new to the

Collection, presented by Guy Aylmer, Hsq., F.Z.S.,0n August 2nd.

1094 ON OPOSSUMS AND DEGUS.

SEPTEMBER,

The registered additions to the Society’s Menagerie during the

month of September were 200 in number. Of these 109 were

acquired by presentation, 21 by purchase, 7 were received on

deposit, 26 in exchange, and 37 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 177.

Amongst the additions special attention may be directed to :—

10 Mouflon (Ovis musimon), from Sardinia, presented by

H.G. The Duke of Bedford, K.G., F.R.8., Pres.Z.8., on September

16th.

1 Tigrine Cat (Felis tigrina) and 1 Three-toed Sloth (Bradypus

tridactylus), from Colombia, presented by W. D. McDougall. Esq.,

on September 15th.

2 Variegated Agoutis (Dasyprocta variegata), 4 Woolly Opos-

suns (Philander laniger), and 1 Sooty Amphisbeena (Amphishena

Juliginosa), from Colombia, presented by Dr. H. G. F. Spurrell,

E.Z.5., on September 13th.

1 Azara’s Opossum (Didelphys azarw), 1 Burmeister’s Cariama

(Cariama burmeistert), and 1 Western Boa (Loa occidentalis),

from Cordova, presented by Wilfred A. Smithers, Esq.,C.M.Z.S., _

on September 16th.

2 Abyssinian Weavers (Cinnamopteryx rubiginosa), new to the

Collection, received in exchange on September 7th.

1 Finch-like Cow-bird (Molothrus fringillarius), from Kastern

Brazil, new to the Collection, presented by the Marquess of

Tavistock, F.Z.8., on September 24th.

1 Brown-headed Parrot (Pwocephalus fuscicapillus), from East

Africa, presented by Lady D. Prain on September 24th.

2 Mikado Pheasants (Calophasis mikado), from Mount Arizan,

Formosa, presented by Mrs. E. J. Johnstone on September 6th,

The Secretary exhibited a preparation of the alimentary tract

of a young Elliot’s Pheasant (Phasianus elliott), dissected so as to

show the gut-pattern in relation to the mesentery, blood-vessels,

and visceral nerves. The preparation was mounted as a trans-

pavent object in glycerine jelly, and showed how the various

morphological conditions of the intestinal tract to which he had

called attention (Trans. Linn. Soc., Zool. vil. p. 173) might be

displayed as permanent museum objects.

Young Woolly Opossums and Degus.

Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals,

exhibited a female and three young specimens of the Woolly

Opossum (Philander laniger), from Buenaventura in Colombia,

presented by Dr. H. G. F. Spurrell, F.Z.S., and drew attention

ON HEEL-PADS IN YOUNG TOUCANET, 1095

to the rapid growth of the young. When Dr. Spurrell captured

the mother, the little ones were still in the pouch. Six weeks

afterwards, when they reached the Gardens, they had left the

pouch for good, were about one-fourth grown, and were just

beginning to feed on their own account, though mainly dependent

upon their mother’s milk. In another six weeks—that is to say,

three months after capture—they were three-fourths their

mother’s. size, and apparently quite capable of fending for

themselves.

Mr. Pococx also exhibited two young examples of a Degu

(Octodon degus), born from a pair from Valparaiso, presented by

Mr. Walter Goodfellow, F.Z.S. The mother had a litter of four,

but died from inability to give birth to a fifth. The little ones

were fed by hand, and although two of them quickly died from

pneumonia, the remaining two throve and promised to do well.

Like the young of all Hystricomorphous rodents, the Degus are

of large size at birth and open their eyes within twenty-four

hours, and are in every respect very precocious as compared with

young rats. Particular attention was directed to the lateral

position of the teats in the Degu, a condition which is repeated

in the Viscacha and Chinchilla and the Coypu. This condition

is probably an adaptation to the large size of the young, which are

enabled to suck lying alongside the nother, so that they do not

interfere with each other. In the Coypu, as first described by

Owen, the teats are higher up the side than in the Chinchilla and

the Degu, an arrangement which permits the young of that

species, it has been alleged, to suck while swimming alongside

their mother in the water.

Heel-Pads in young Toucanet.

Mr. D. Sera-Smiru. F.Z.8., Curator of Birds, exhibited a

spirit-specimen of a young Toucanet (Selenidera maculirostris)

(text-fig. 191) which had been hatched in the Society’s Gardens

in July, and called special attention to the well-developed serrated

pads on the back of the ankle-joint or heel. This pad had been

previously observed in the nestlings of Woodpeckers, Wrynecks,

and Barbets, and it was only to be expected that it would be

present also in the Toucans.

The use of this serrated pad, which disappears soon after the

young bird leaves the nest, is doubtless to enable it to climb up

the side of the hollow cavity in a tree in which it is hatched—

these pads forming, as it were, a second set of claws. It is note-

worthy that no such pads are found in the Parrots, which nest

in similar situations to those chosen by Barbets and Toucans, but

which are provided with a hooked beak which aids them in

climbing.

1096 ON MELANOCELDUS JOMNSONII.

Text-fig. 191.

Nestling of Spot-billed Toucanet (Selenidera maculirostris) showing feather-tracts

and serrated pads (a) on metatarsal joint. A. Enlarged view of serrated

pad.

A Deep-Sea Angler-Pish, MELANOCETUS JOHNSONII.

Mr. C. Tate Ruean, M.A., F.Z.S., exhibited a specimen of a

remarkable fish, Melanocetus johnsonit Giinth., that had been

presented to the British Museum by the Hon. Walter Rothschild,

F.R.S., and made the following remarks :—

“This eurlous fish has an enormous mouth armed with slender

ON A YOUNG MATAMATA TERRAPIN. 1097

pointed depressible teeth and an extraordinarily distensible

stomach ; it belongs to the order Pediculati and is a bathy pelagic

species, uniformly blackish in colour; like the Angler (Lophius

piscatorius) it has the first ray of the spinous dorsal fin situated

on the snout and modified into a line and bait; in this ease the

bait is a luminous bulb.

«“The species is known from six specimens from the North

Atlantic, three dredged at considerable depths by the ‘ Travailleur ’

‘Talisman,’ ‘ Valdivia’ and ‘ Michael Sars’ expeditions, and

three in the British Museum collection. The last are from 3 to

3? inches long (from snout to end of caudal fin) and include the

type described by Gunther (Proc. Zool. Soc. 1864, p. 301, pl. xxv.)

from Madeira; this had in its stomacha partly digested Scopeloid

fish (Lampanyetus crocodilus Risso) about 7 inches long. The

second specimen was taken at the surface in 31° N. and 37° W.,

and had in its stomach an almost perfect ZL. crocodilus nearly

8 inches long; this was captured by Captain Charles C . Dixon,

master of the bark ‘Hlginshire, who also secured the fish

exhibited to-night at 7 a.m. on August 2/th, 1910, in 40° 19' N.

and 42° 25' W. By a strange coincidence this specimen also

contains a LZ. crocodilus, and judging by the anterior part of the

head, which is still visible, one about 8 inches long; it is curled

round into a complete circle in the stomach of the Metinoceme

‘‘Captain Dixon writes that the J/elanocetus was stiil alive

when captured and that from his observations he believes that it

could make the ‘lure’ luminous or not at will. The fact that

it was caught early in the morning is interesting, for it is known

that at night many fishes that dwell at a depth of 300 fathoms

or more may approach much nearer the surface; here the large

Lampanyctus may have been seized and its struggles may have

brought the captor still further out of its depth and rendered it

helpless.”

November 11, 1913.

Dr. S. F. Harmer, M.A., F.B.S., F.ZS.,

in the Chair.

Mr. E. G. Boutrnesr, F.Z.8., Curator of Reptiles, exhibited

a quite young specimen of the Matamarn Terrapin (Chelys fim-

br iata), the shell of which measured but 5 em. in length. This

specimen, the smallest yet recorded, differed from the existing

descriptions of the young of this species in the under surfaces

being of a bright magenta-red instead of pale yellow. Mr. Bou-

lenger suggested that as all the young previously described were

more than double the ‘size of the specimen exhibited, red might

be the normal coloration of the under surfaces as a juvenile

livery.

Text-fig. 192.

1098 ON A HYBRID BLAGK-WINGED PEAFOWL.

Hybrid Birds.

Mr. D. Sers-Srru, F.Z.S., Curator of Birds, exhibited a

remarkable hybrid (text-fig. 192) between a Black-winged Peacock

(Pavo nigripennis) and a Domestic Hen (said to be a cross-bred

game and Leghorn), which had been bred and presented to the

Society by Mr. R. P. Wheadon of minster. This bird was

hatched in 1911 and is believed to be only the second example

Photo. by D. Seth-Smith.

th its mother, a brown-and-white domestic hen.

ged Peafowl, wi

-win

Hybrid Black

ON DORSAL GLANDS IN ARMADILLOS. 1099

of this hybrid ever produced, although hybrids between peafowl

and guinea-fowl had been recorded. It was mostly white in

colour, splashed, especially on the neck, with reddish brown, a

colour derived from the domestic hen, its mother. It ehonwed

clearly the characters of both parents.

Mr. Seth-Smith also showed photographs, by means of lantern-

slides, of a male hybrid Pheasant between Calophasis nikado

and C. ellioti and of a male Jungle-fowl between Gallus varius

and G'. gallus.

The former had been bred in Sussex by Mrs. E. J. Johnstone,

and kindly presented to the Society. It resembled a very dark-

coloured Elliot’s Pheasant, with the head and neck, breast, and

abdomen black. The tail was black, suffused with chestnut and

crossed with broad white bands.

The hybrid Gallus, which was one of several bred in the

Gardens, was very intermediate between the two parent species.

The comb was shaped like that of G. varius, but was shghtly

serrated, while the central wattle of G. variws was present

though reduced in size, as well as the two side wattles of

G. gallus. The neck hackles, which in G. varius are short and

rounded, were in the hybrid moderately long as in G. gallus,

but rounded at the ends, while the colour of these hackles

as well as of the tail, was violet-purple. This bird appeared to

be identical with that named Gallus eneus by Temminck.

Dorsal Glands in Armadillos.

(‘Text-figures 193-195.)

Mr. R. I. Pocock, F.B.S., F.Z.S., Curator of Mammals, ex-

hibited some specimens of Armadillos of the genus Hauphractus*

(formerly Dasypus) to show the presence of a row of glands upon

the pelvic portion of the carapace, and remarked :—‘“ In the

middle line of the pelvie shield there is an unpaired series of two,

three, or four sears which are ill-defined in the living animal but

tolerably conspicuous in dried skins and clearly marked when the

scales are removed from the osseous scutes. ‘Their conspicuousness

in museum specimens cannot have escaped the notice of syste-

matic workers, but apparently their significance has never been

investigated.

os My attention was originally drawn to them by noticing that

in newly born Hairy Armadillos (#. villosws) there are two or

three small holes, one on the fourth, a second on the fifth, and

sometimes a third on the sixth row of scutes of the shield in

question. At this early stage the integument 1s unossified and can

easily be cut with a razor. <A section through the holes in

question shows that they are the orifices of shallow pits directed

* According to Thomas, P. Z.S. 1911, p. 141, Dasypus must be transferred to the

Peba Armadillos, replacing J GeO

1100 MR. R. I. POCOCK ON

downwards ant forwards and dipping into the dermis which is

especially thickened and apparently highly glandular beneath

Text-fig. 193.

A. Longitudinal section of the three glands in the newly born

Hairy Armadillo (Huphractus villosus).

Ep., epidermis ; D, glandular thickened dermis; O, orifice of pit.

B. Pelvic carapace of the same showing the glandular orifices

on the 4th, 5th and 6th belts.

DORSAL GLANDS IN ARMADILLOS. 1101

them, the topmost layer of the thickening being marked with a

few spots of black pigment. The glandular nature of the pits

was readily ascertained in the case of living adult armadillos of

Text-fig. 194.

Photograph of part of the pelvic carapace of an adult living Hairy Armadillo

showing the three glands on the 4th, 5th, and 6th belts. (The gland-bearing

scute of the 6th belt was fractured and the pits being filled with secretion

showed up more clearly than when dry.)

this species by squeezing the scutes bearing the scars and pressing

from them drops of fluid with the same objectionable odour as

Proc. Zoou, Soc.—1913, No. LX XIII. 73

1102 DORSAL GLANDS IN ARMADILLOS.

that which characterises these animals. Since the glands, judging

from external appearances, are equally developed in adults of both

sexes and appear at birth, there is no reason for thinking their

function is in any way sexual. Their smell may be sufficiently

nauseous to protect armadillos from enemies or it may act only

as a guide enabling these animals to track each other and keep

together.

Text-fig. 195.

Drawing of the central portion of the pelvic carapace of the Six-banded Armadillo

(Euphractus sexcinctus) with four glandular pits, suggesting the origin of

the pits from the enlargement and concrescence of the bristle-pores.

‘‘ When the dorsal carapace of an armadillo is stripped, cleaned

and dried, the position of the glands is marked on the under side

of the pelvic shield by a row of bony prominences. Each pro-

minence represents a gland and is in reality a hollow bony pit

resulting from the ossification of the dermal layer immediately

beneath the gland. This at all events is the case in a carapace of

Euphractus sexcinctus kindly lent to me by Mr. R. EK. Holding. In

this specimen there are seven free rings behind the scapular

shield, but the seventh is not so free as those that precede it.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 1103

Following this are nine rings fused together and constituting the

pelvic shield. The median glandular pits are observable upon the

third, fourth, fifth and sixth of these ; and the scutes which carry

them are distinguished by their shape from those of the rest of

the pelvic shield. The glandular pit upon the sixth ring is much

smaller and shallower than the rest and is especially interesting

as showing the origin of the specialized glandular areas. The

scutes of the carapace in this speciesare covered with pores, many

of which are quite small; but a varying number, generally about

ten and lying in two grooves, are much larger and arranged in two

longitudinal lines meeting anteriorly and forming a somewhat

horseshoe-shaped pattern. That these large pores perforate

the bony scutes may be seen by holding the carapace against

the light.

‘“‘ According to de Meijere* these pores are bristle-pores and

indicate lines of fusion between small and originally separated

scutes. However that may be, there is no doubt, I think, that

the glands above described result from the concentration and fusion

of these rows of pores. The evidence for this is particularly strong

in the case of the posterior of the four glands in Huphractus

sexcinctus where the pores may be seen to be concentrated in the

shallow central depression of the scute that bears them.”

November 25, 1913.

Prof, E. W. MacBrips, M.A., D.Sc., F.R.S., Vice-President,

in the Chair.

THe SEcrETARY read the following report on the additions

made to the Society’s Menagerie during the month of October

1913 :—

The number of registered additions to the Society’s Menagerie

during the month of October last was 214. Of these 104 were

acquired by presentation, 42 by purchase, 16 were received on

deposit, 38 in exchange, and 14 were born in the Gardens.

The number of departures during the same period, by deaths

and removals, was 237.

’ Amongst the additions special attention may be directed to :—

3 Entellus Langurs (Presbytis entellus), from North India, and

1 Wanderoo Macaque (Macacus silenus), from Malabar, presented

by W. O. Danckwerts, Esq., K.C., F.Z:S., on October 14th and

24th.

1 Sclater’s Dog (Canis sclateri), from the Amazons, purchased

on October 2nd.

1 Mexican Tree-Porcupine (Coendu mexicanus) and 1 Peba

* De Meijere, Morphol. Jahrb. xxi. pp. 316, 318 (1894).

C3"

1104 MR. R. H. BURNE ON F@TAL SKELETONS.

Armadillo (Tatusia novem-cincta), from Mexico, purchased on

October 8th and 30th.

1 Dorsal Hyrax (Dendrohyrax dorsalis), from Sierra Leone,

presented by Mrs. A. M. Bettington on October 7th.

1 Black-backed Hangnest (/cterws mesomelas), from Central

America, | Loo-Choo Jay (Lalocitta lidthii), from the Loo-Choo

Islands, and 1 Purple-breasted Parrot (Zriclaria cyanogastra),

from 8.E. Brazil, all new to the Collection, received in exchange

on October 27th.

2 American Harriers (Circus hudsonius) and 1 Mississippi Kite

(Icterus mississippiensis), from North America, both new to the

Collection, received in exchange on October 7th.

3 Pileated Guans (Penelope pileata), 3 Araucuan Guans (Ortalis

araucuan), and 2 Rufous-necked Rails (Aramides chiricote), from

Para, all new to the Collection, presented by the Goeldi Museum

on October 20th.

2 Brazilian Tortoises (Platemys platycephala), new to the

Collection, purchased on October 15th.

Prof. H. Maxweti Lerroy, M.A., F.Z.S., Honorory Curator of

Insects, briefly described the arrangement of the New Insect

House, the progress made with the fitting-up of the interior, and

the installation of and method of displaying the exhibits.

Mr. R. H. Burne, M.A., F.Z.S., showed some specimens of

foetal skeletons prepared by the Beale-Schultze method. The

fetus after being dehydrated is soaked in 1-3 per cent. caustic

potash, and finally mounted in glycerine. By this procedure the

soft parts become almost completely transparent, leaving any

calcareous deposit opaque.

The specimens shown were fetuses of the Fowl, Horse, and

Man. In the Fow!] attention was drawn to the three separate

metatarsals that at an older stage fuse to form the single tarso-

metatarsus. In the same way in the Horse the ossifying shafts

of the ulna, fibula, and splint-bones could be clearly distinguished.

A specimen of the skeleton of a Star-fish (Astertas rubens)

prepared by the same method was also exhibited.

No. 124.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

October 28th, 1913.

Prof. E, A, Miycuin, M.A., F,R.8., F.Z.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed,

The Sscretary read a Report on the Additions made to the

Society's Menagerie during the months May to September,

ONS: ;

The Secretary exhibited a preparation of the alimentary tract

of a young Elliot’s Pheasant (Phastanus elliott), dissected so as to

show the gut-pattern in relation to the mesentery, blood-vessels,

and visceral nerves. The preparation was mounted as a trans-

parent object in glycerine jelly, and showed how the various

morphological conditions of the intestinal tract to which he had

called attention (Trans. Linn. Soc., Zool. vill. p, 178) might be

displayed as permanent museum objects.

Mr. R. I. Pocock, F.R.S., F.Z.8., Curator of Mammals,

exhibited a female and three young specimens of the Woolly

Opossum (Philander laniger), from Buenaventura in Colombia,

presented by Dr. H. G. F. Spurrell, F.Z.S., and drew attention

to the rapid growth of the young. When Dr, Spurrell cap-

tured the mother, the little ones were still in the pouch.

Six weeks afterwards, when they reached the Gardens, they

had left the pouch for good, were about one-fourth grown,

and were just beginning to feed on their own account, though

* This Abstract is published by the Society at its offices, Zoological Gardens

Regent’s Park, N.W., on the Tuesday following the date of Meeting to whieh

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post -free for

the sum of Six Shillings per annum, payable in advance,

mainly dependent upon their mother’s milk. In another six

weeks—that is to say, three months after capture—they were

three-fourths their mother’s size, and apparently quite capable of

fending for themselves,

Mr. Pocock also exhibited two young examples of a Degu

(Octodon degus), born from a pair from Valparaiso, presented by

Mr. Walter Goodfellow, F.Z.S. The mother had a litter of four,

but died from inability to give birth to a fifth. The little ones

were fed by hand, and although two of them quickly died from

pneumonia, the remaining two throve and promised to do well.

Like the young of all Hystricomorphous rodents, the Degus are

of large size at birth and open their eyes within twenty-four hours,

and are in every respect very precocious as compared with young

vats. Particular attention was directed to the lateral position of

the teats in the Degu, a condition which is repeated in the Viscacha

and Chinchilla and the Coypu. This condition is probably an

adaptation to the large size of the young, which are enabled to

suck lying alongside the mother, so that they do not interfere

with each other. In the Coypu, as first described by Owen, the

teats ave higher up the side than in the Chinchilla and the Degu,

an arrangement which permits the young of that species, it has

been alleged, to suck while swimming alongside their mother in

the water.

Mr. D. Seru-Smiru, F.Z.S., Curator of Birds, exhibited a

spirit-specimen of a young Toucanet (Selenidera maculirostris)

which had been hatched in the Society’s Gardens in July, and

called special attention to the well-developed serrated pads on

the back of the ankle-joint or heel. ‘This pad had been pre-

viously observed in the nestlings of Woodpeckers, Wrynecks,

and Barbets, and it was only to be expected that it would be

present also in the Toucans.

The use of this serrated pad, which disappears soon after the

young bird leaves the nest, is doubtless to enable it to climb up

the side of the hollow cavity in a tree in which it is hatched—

these pads forming, as it were, a second set of claws. It is note-

worthy that no such pads are found in the Parrots, which nest

in similar situations to those chosen by Barbets and Toucans, but

which are provided with a hooked beak which aids them in

climbing.

Mr. C. Tare Recan, M.A., F.Z.8., exhibited a specimen of

Melanocetus johnsonii Giinther, a curious fish having an enormous

mouth armed with slender, pointed, depressible teeth and an

extraordinarily distensible stomach. It has the first ray of the

spinous dorsal fin situated on the snout and modified into a

line and bait, the latter being a luminous bulb,

Dr. F. E. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the

Society, read a paper on the anatomy and systematic arrangement

of the Cestoidea, in which he described a new genus and species

of Tapeworms from the Double-striped Thicknee (Wdicnenwus

bistriatus).

Dr. F. A. Baruer, M.A., F.R.S., F.Z.8., read a paper entitled

“The Fossil Crinoids referred to Hy ypocrinus Beyrich.” The

two specimens of Zypocrinus schneidert Beyr. described by Beyrich

and Rothpletz respectively are redescribed and refigured. The

structure of the genus is shown to agree with that of the Devonian

family Gasterocomide, the content of which is discussed ; but it

is suggested that in this case and in that of ‘ Lecythiocrinus”

adamst the distinctive features may have been independently

acquired.

The holotype of Hypocrinus piriformis Rothpletz is redescribed

and refigured, and proved to be no Hypocrinus. It is thought to

be a highly modified descendant of the Taxocrinide, by way of

such a genus as Cydonocrinus. The left posterior radial appears

to have borne a large arm, but the other arms are more or less

atrophied, and the right posterior radial has almost disappeared.

A paper on “ Batrachiderpeton lineatum Hancock & Atthey,

a Coal-Measure Stegocephalian,” communicated by Prof. J. P.

Hill, D.Sc., F.R.S., F.Z.S., was read by Mr. D. M.S. Watson,

M.Se. It contained the description of the skull, lower jaw, and

pectoral girdle of this species, based on a series of specimens in

the Newcastle Museum, derived from the Low Main Seam of

Newsham Colliery.

A paper, communicated by Dr. C. W. Andrews, F.R.S., F.Z.5.,

was received from Mr. R. W. Patmer, M.Sc., entitled ‘‘ The

Brain and Brain-Case of a Fossil Ungulate of the Genus

Anoplotherium,” in which a cranium from the Phosphorites

of Quercy, together with an exceptionally perfect and well-

marked brain-cast obtained from it, were described from material

in the British Museum collections.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, November 11th, 1913, at half-past Hight

o'clock p.u., when the following communications will be made :-—

Exnrerrions AnD Noricrs.

Sree seattle ctor een

W. T. Cauman, D.Se.; F.Z.S.

On Freshwater Decapod Crustacea (Families Potamonide

and Palemonide) collected in Madagascar by the Hon. Paul

A. Methuen.

On a Collection of Reptiles and Batrachians made by

Dr. H G. F. Spuvrell, F.Z.S., in the Colombian Choco.

C. Tare Rea@an, M.A., F.Z.S.

A Revision of the Cyprinodont Fishes of the Subfamily

Poeciliinee.

Prof. W. N. Parxer, Ph.D,, F.Z:8.

Sponges in Waterworks.

Prof. J. Puayrain McMurricu, C.M.Z.S.

On Two new Actinians from the Coast of British Columbia.

The following papers have been received :—

SORTAN OLSEN.

On the External Characters and Biology of Bryde’s Whale,

a new Rorqual from the Coast of South Africa.

Miss M. V. Lepour, M.Sc.

A new Species of Trematodes of the Genus Lechriorehis from

the Dark Green Snake (Zamenis gemonensis).

T. AH. Wivuers, E.G.

Ciripedes from the Cenomanian Chalk Marl of Cambridge.

F. F. Laipnaw, M.A., F.Z.S.

Contributions to a Study of the Dragon-fly Fauna of Borneo.

—Part II. The Gomphine and Chlorogomphine.

Surgeon Josern C. Taompgon, U.S.N.

Further Contributions to the Anatomy of the Ophidia,

2 Cir eile) ilravotoneteg Mieke, Lees END US ZAI

The Peroneal Muscles in Birds.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL SoctEery oF Lonpon,

REGENT’s Park, Lonpon, N.W.

November Ath, 1913.

ate Rann 3 7m k

eg ROD PERRO y boas

re ?

t , ie

No. 125.

ABSTRACT OF THE PROCEEDINGS ~

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

November 11th, 1913.

Dr. 8. F. Harmer, M.A., F.R.S., F.Z.S., in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

Mr. R. I. Pococr, F.R.S., F.Z.S., Curator of Mammals, ex-

hibited a newly-born example of the Hairy Armadillo (Luphractus

villosus) and the dried shell of a Six-banded Armadillo (Z. sea-

cinctus), Showing the presence of a series of from two to four

small orifices in the middle line of the pelvic shield; and explained

that the orifices are the apertures of shallow invaginations of the

epidermis dipping down into the dermis, which is specially

thickened and glandular immediately beneath them, In the

living animal drops of malodorous fluid can be squeezed from the

orifices of these glands.

Mr. D. Seru-Smitu, F.Z.S., Curator of Birds, exhibited a re-

markable hybrid between a Black-winged Peacock and a domestic

_hen (cross-bred game and Leghorn) which had been bred by

Mr. R. P. Wheadon of Ilminster in 1911. This bird was be-

lieved to be only the second example of this hybrid ever produced,

although a hybrid between a peafowl and a guinea-fowl had been

recorded (Hocker, J. f. O. 1870, p. 152).

The hybrid was mostly white in colour, splashed, especially on

the neck, with reddish brown, a colour derived from the domestic

hen, its mother. It showed clearly the characters of both

parents.

Mr. Seth-Smith also showed photographs of a hybrid Pheasant

between Calophasis mikado and C’ elliott, and of a Jungle-fowl

between Gallus varius and G, gallus. The latter bird, which had

been bred in the Society’s Gardens, corresponded to the description

of the bird described by Temminck as Gallus eneus.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance,

Mr. E. G. Bounencer, F.Z.8., Curator of Reptiles, exhibited

a quite young specimen of the Matamata Terrapin (Chelys jim-

briata), the shell of which measured but 5 cm. in length. This

specimen, the smallest yet recorded, differed from the existing

descriptions of the young of this species in the under surfaces

being of a bright magenta-red instead of pale yellow. Mr. Bou-

lenger suggested that as all the young previously described were

more than double the size of the specimen exhibited, red might

be the normal coloration of the under surfaces as a juvenile

livery.

Dr. W. T. Catman, F.Z.S., read a paper on Freshwater Decapod

Crustacea (families Potamonide and Palemonide) collected in

Madagascar by the Hon. Paul A. Methuen, in which one new

species of Potamon and five varietal forms of P. madagascariense

were described. It is suggested that the River-Crabs of Mada-

gascar may have had an autochthonous origin from some form

resembling P, madagascariense. No clear affinities can be traced

with the Potamonide of Africa or of Peninsular India, but it is

pointed out that in the present state of knowledge the River-

Crabs appear to be a hazardous subject for zoogeographical

speculation.

Among the Palemonide the most interesting is the species

’ described by Hilgendorf as “‘ Bithynis ? hildebrandiz,” which is

figured for the first time. Evidence is brought forward to show

that the species has no close affinity with the Bithynis gaudichaudi

of Chili and Peru, and that the single character which led to its

being referred to Lithynis is a matter of individual variation.

Mr. G. A. Boutenegsr, F.R.S., F.Z.8., gave an account of a

collection of Reptiles and Batrachians made by Dr. Spurrell,

F.Z.8., in the Colombian Choco and presented by him to the

British Museum. The series of specimens was of great interest,

and contained several new species.

Mr. C, Tarr Rucan, M,A., F.Z.S., read a paper entitled

“A Revision of the Cyprinodont Fishes of the Subfamily

Peeciliinz.” A number of new genera were defined and several

new species were described ; the structure of the intromittent

organ was found to be of great systematic importance,

A paper received from Prof. W. N. Parker, Ph.D., F.Z8.,

gave an account of some investigations made on a growth of

Spongilla lacustris in the Cardiff Waterworks system, and described

the methods adopted to eradicate the sponge from the infected

areas,

Prof. J. Puayrarr McMurricu, C.M.Z.S., communicated a

paper in which he described two new species of Actinians from

the coast of British Columbia. These specimens probably repre-

sented stages of a single species, and belonged to a group hitherto

not recorded from the west coast of America.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, November 25th, 1913, at half-past Eight

o’clock P.m., when the following Conmcniic ations will be made :—

EXHIBITIONS AND NOTICES.

RIAN OLSEN.

On the External Characters and Biology of Bryde’ s Whale,

a new Rorqual from the Coast of South Africa.

Miss M. V. Lesour, M.Sc.

A new Species of Trematodes of the Genus Lechriorchis from

the Dark Green Snake (Zamenis gemonensis).

T. H. Wrruers, F.G.S8.

Cirripedes from the Cenomanian Chalk Marl of Cambridge

P. Cu CHALMERS Mircnent, M.A. IDs 1h LL.D., emia R.S., PAS.

ernie Per oneal Maneales = in ae

The following papers have been received :—

FE. F. Larpuaw, M.A., F.Z.S.

Contributions to a Study of the Dragon-fly Fauna of Borneo.

—Part II. The Gomphine and Chlorogomphine.

Surgeon JosrpH C. THompson, US.N

Further Connbutione to ey ree of the Ophidia.

EU Ue eee

Sen on an Bieri dovolaped Specimen of Hehinus

esculentus.

Gre EN BouLENGER, E.R.S., B.ZS.

‘An Real ist ee the Reptiles and Batrachians collected

by the British Ornithologists’ Union Expedition and the

Wollaston Expedition in Dutch New Guinea.

C.F. U. Mzex, MSc. LS. F.ZS8.

“The Possible Comecaon between Spindle-length and Cell-

volume.

W ILLIAM Nicout, M.A., 7D) Sc., ., M -D., 8 ZS.

Ty eines peed Ba om Gittins dying i in the Zoological

Society’s Gardens during 1911-1912.

mic ae 7 ce ae a oe Arrangement

of the Cestoidea.— XII. Further Observations upon the! Genus

Urocystidium Beddard.

Communications intended for the Scientific Meetings shoul

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZooLoGIcAL Society oF Lonpon,

Recent’s Park, Lonpoy, N.W.

November 18th, 1913.

No. 126.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

November 25th, 1913.

Prof. E. W. MacBripe, M.A., D.Sc., F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Spcrerary read a Report on the Additions made to the

Society’s Menagerie during the month of October, 1913.

Mr. R. H. Burne, M.A., F.Z.S., showed some specimens of feetal

skeletons prepared by the Beale-Schultze method. The fetus

after being dehydrated is soaked in 1-3 per cent. caustic potash

and finally mounted in glycerine. By this procedure the soft

parts become almost completely transparent, leaving any calcareous

deposit opaque.

The specimens shown were feetuses of the Fowl, Horse, and

Man. In the Fowl attention was drawn to the three separate

metatarsals that at an older stage fuse to form the single tarso-

metatarsus. In the same way in the Horse, the ossifying shafts

of the ulna, fibula, and splint-bones could be clearly distinguished.

A specimen of the skeleton of a Starfish (Aséerias rubens),

prepared by the same method, was also exhibited.

Prof. H. Maxwetu Lerroy, F.Z.8., Hon. Curator of the Insect

House, briefly described the arrangement of this new building,

the progress made with the fitting-up of the interior, and the

installation of and method of displaying the exhibits.

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent’s Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications ; but it may be obtained on the

day of publication at the price of Srpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance.

A paper received from Mr. Orsan OLseN, and communicated

by the Secretary, contained the description of a new Rorqual from

the Coast of South Africa and gave a detailed account of its

external characters, biology, and distribution.

Miss Marie V. Lrsour, M.Sc., presented a paper, communi-

cated by the Secretary, in which she described a new species of

Trematodes of the genus Lechriorchis, found in the body-cavity

of a Dark Green Snake (Zamenis gemonensis) that had died in

the Society’s Gardens.

Mr. T. H. Wirners, F.G.S., contributed a paper, communi-

cated by Dr. W. T. Calman, F.Z.8., based upon a large series of

Cirripede remains from the Cenomanian Chalk Marl in the neigh-

bourhood of Cambridge. The greater number of the specimens

are referred to two species of the family Pollicipedide, and add

materially to our knowledge of the phylogeny of the pedunculated

Cirvipedes. Both forms are remarkable for their advanced form

of seutum, in which the umbo is sub-central, and show that the

transition of the scutal umbo from an apical to a sub-central

position was acquired independently by unrelated forms in distinct

lines of development.

The Srcrerary, Dr. P. Coatmers Mircueti, M.A., F.R.S.,

F.Z.8., read a communication on ‘‘The Peroneal Muscles in

Birds.” Dr. Mitchell had dissected these muscles in over 300

birds, and believed that he was able to give a nearly exhaustive

account of the varieties of form presented by these structures.

The paper described the peroneal muscles in Chawna chavaria,

and gave a systematic account of the conditions in the different

Avian groups which could all be represented as derivatives of

the Chauna condition by loss of certain portions and increased

development of other portions. It was diiticult to interpret the

changes in terms of function, family-relationship appearing to be

more important in this respect than habit. The paper discussed

the very few cases of want of conformity between the peroneal

nuscles and the place assigned in the best-founded systems of

classification. It also dealt with the use of such anatomical

characters in systematic classification, and showed that there was

a definite relation between specialized conditions of the peroneal

muscles and specialized conditions of the Avian structure

generally.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, February 3rd, 1914, at half-past Hight

o'clock p.m., when the following communications will be made :—

EXHIBITIONS AND Noricnus.

G. A. Boutencsr, F.R.S., F.Z.8.

An Annotated List of the Reptiles and Batrachians collected

by the British Ornithologists’ Union Expedition and the

Wollaston Expedition in Dutch New Guinea.

Se ae ae cae aa ea Arrangement

of the Cestoidea.—XII. Further Observations upon the Genus

Urocystidium Beddard.

H.G. Prmver, F.R.S., P.Z.5

ae mi a a Se occurred in the Zoological

Gardens during 1913.

The following papers have been received :—

FF, Larpraw, MA. F.ZS8. —

Contributions to a Study of the Dragon-fly Fauna of Borneo.

—Part II. The Gomphine and Chlorogomphine.

Surgeon Josep C. Tuompson, U.S.N.

Further Contributions to the Anatomy of the Ophidia.

H. C. Cuapwick, A.L S.

Note on an imperfectly - developed Specimen of Hehinus

esculentus.

CT Ue Mipies Wleee UNly ein eats.

Tike a eeae Ge peoween! Spindle- -length and Cell-

volume.

Wiiuram Nicotu, M.A., D.Sc., M.D., F.Z.8.

Bie eae ele

Tyematode Parasites from Animals dying in the Zoological

Society’s Gardens during 1911-1912.

F. J. Mraerrt, M.Se

The Structure and Life-History of a Tapeworm (Lchthyotenia

Jilicollis Rud.) parasitic in the Stickleback.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpon,

Rueent’s Parx, Lonpon, N.W.

December 2nd, 1913.

ol.

53.

A New rematode of the Genus Lechriorchis from the Dark Green Snake (Zamenis

gemonensis). By Marin V. Lupour, M.Sc., Assistant Lecturer and Demonstrator in

Zaolory, Weeds University. (Pl MCLE) site ge te ccs See eee ecw nese rs

54, Cirripedes from the Cenomanian Chalk Marl of Cambridge. By Tuomas H. Wirusrs,

BE erses eee (Chal NCO ENV ee RG NCO) Vis) avs etc ictal arene talon «is ciaaee“e'lalwns sista sistaiphers ott oletpiera'e ed

55, Batrachiderpeton lineatum Hancock & Atthey, a Coal-Measure Stegocephalian. By

D. M. 8S. Warson, M.Sc., Lecturer on Vertebrate Palaontology in University College,

Hondon. (Pls. XCVI. & XCVII. and Text-figs. 162-107.) ...... 2.0.02 ...s cece

56, On two New Actinians from the Coast of British Columbia. By J. Pusyrair

IMeniCorpnoe Oni Aish = Gab ONALRE icine ad dade an onScMeonooConobocenboOooc Onn

57. Sponges in Waterworks. By W. N. Parxer, Ph.D., F.Z.8., Professor of Zoology,

Unrweneiiy- Collage, (Chnthtttcouscoccaocndovduscser on duos acennde syslesaranie whose tenets

58. A Revision of the Cyprinodont Fishes of the Subfamily Peciliine By C. Tats

Ruean, M.A., F.Z.S. (Pls. XCIX.-OI. and Text-figs. 168-173.) ........++-.00-00s

59. On a Collection of Batrachians and Reptiles made by Dr. H. G. F. Spurrell, F.Z.S., in

the Choco, Colombia. By G. A. Bouruyerr, F.KS., F.ZS8. (Pls. C1L-CVII.

SUMTER POR ali AiG 5) aie we eicce eee n cinch, oo ale olniote apices 6 a's wha cos siee siete aig rich

40. The Peroneal Muscles in Birds. By P. Cuaumrers Mircng.y, M.A., D.&c., LL.D.,

F.RBS., F.Z.S8., Secretary to the Society. (Text-figs. 179-190.) ..........e-sseeee

61. On the External Characters and Biology of Bryde’s Whale (Balenopitera brydei), a

new Rorqual from the Coast of South Africa. By Mrsan Oxsuy, Zoological

Laboratory, Christiania University. (Pls. CIX.-CXIIL.).......... 000s. ereens

CMGI AED Sagiaqoks orien acne aS onan uonciintipn eaten Ae seldtacio Seis Emery rer Ame eo

Mast OMe OUNCME MELO LNCETS: aan, <ioh eval of crs ciescie s!«)-ele/oidcatlei-oinim sl cleats vin moles ic aie ln'e Gorka :

brats OF ClonesiiS so on cu dbconn ange 6000 dene UOEOULhe Doub an OO CUMeD SD UOOeES Co Dn de Soe

Alphabetical List of Contributors .......... -.. ab cis dg OAR Odo SHO AGO Cue soo Lo oon

EN@wa Generic Donn sisyseicrteec sera) e oteinc toss sicreatiar a toile: « Bia ciel eb eared iol ovate steislaraieicle: shelsiaeie

iaglae @E SieianinitG INEINES Ses hoo con ogc don dd 50 KOUn Guat Op oo ues HAO sopoodod oo :

Biirrtve aro ce elas teen proms Gers s: lace scaoe vei olstoe a ereitiare eicletaieiiar ct aneketauas suatsravansavn elapse eilelencvate lesson tonslinne

Papers (continued).

The Fossil Crinoids referred to Hypocrinus Beyrich. By F. A. Barnur, M.A., D.£c.,

bors Hee See (Eee sand: NeXtn eS OOM.) cei e's ie wore civisis eaimie wie cele oa als

. On Freshwater Decapod Crustacea (Families Potamonids and Palemonidz) collected

in Madagascar by the Hon. Paul A. Methuen. By W. T. Cauman, D.8c., F.Z.8.

Gi seeNUC Mero oN Cll eancl Metin trom slallly Wractenyoleletorti ere c's e vies siertierace ols e10) ole e/ekelecinmis ols

Pago

894

914

933

937

949

963

973

977

1019

1039

1073

LIST? OF PLATES,

1913, Part LV. (pp. 861-1104).

Plate Page

XC. 1-6. Hypocrinus schneideri. oe “FL? purcfornst,.ea82 & Oot

XOI. Potamon methuent....... tro tttntereeetereeeeereee | gry

XOIL -Palemon hildebrandtii 22, }

ICME Lechrvonchis weniiisiteteetateyeia ciebevelaliate)ereraiele sipensten tote eee 933

KCEV.. -Zengmatolepas mocklert wea. +» scls.)ea es oc ote wee ste aera “937°

XCV. Calantica (Titanolepas) tuberculata ....++....<+.-- Fiete es \

Se } BORO ELIGAS OPRT D.C 50 HAG FO DOO iC oie.) Adee

XOVIII. 1-4. Peachia quinquecapitata. 5-7. Bicidiwm equoree.

SABA panasicicwmerawere «ati oe ole slo eee oreo 963.

XOIX. 1,2. Gambusia oligosticta., 3,4. G. wrayi. 5,6. G. gra-\

eoliors. “x (Gr, GhOMTNGHUSES * SaobOS oc anne accocnand0s8 os |

C. 1, 2. Pecilia picta. 3, 4. Peciliopsis isthmensis. 5, 6. | 977

Phalloceros caudomaculatus. 7, 8. Phalloptychus januarius. f

CI. 1,2. Limia nigrofasciata. 3, 4. L. heterandria. 5. L. |

arnoldi. 6. L. caudofasciata. 7. L. ornata............/)

CII. 1. Bufo hypomelas. 2. Agalychnis calcarifer ........+++» yl

CIO Agalychnis\spur rena nacelle aha idler tainty = = ateleietcla hina |

CIV. 1. Dendrobates awrotenia. 2-4. D. tinctorius, var, coctéi.

5, 6. D. tinctorius, var. Chocensts .s..-...-seereecees

ean \ OiNOstenmartySPUnnellt Wa carere teietelegsietetntiets etek ee ee eee riOle

CVIL. 1. Anolis breviceps. 2. Anadia vittata ......06.02..00005

CVIII. 1. Homalocranium coralliventre. 2. Hlaps microps.

3. Leptognathus spurrellt .. i. -0secess veers sccccs ose

CIX. )

@X. | Balenoptera brydet 72... 2. ee ce www ces ce ce AMODOS O08

CXI. | 1073:

CXII.. 9, 11a. Balenoptera borealis. 10,110. B. brydet ........ |

(Ch:GUU ny IRA AMMAR OVO» argosuotseAaontobs obodcb on casmqecce y)

NOTICE.

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,

so that the complete reference is now P. Z. 8. 1913, P- ... The Distribution

is as follows:—

Part I. seared in March.

pie ld Remains June.

pry EB - September.

a SEL ee December.

‘Proceedings,’ 1913, Part III. (pp. 3389-860), were published on

September Ist, 1913.

The Abstracts of the ‘Proceedings,’ Nos. 124-126, are

contained in this Part.