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PROCKEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY %

OF LONDON.

1912, pp. 505-913,

with 37 Puates and 63 TEx'-FIGURES.

Se Oey ee

4 ay : er a

wee Gr] MS! ;

PRINTED FOR THE SOCIETY,

‘AND SOLD AT THEIR HOUSE IN REGENT’S PARK.

LONDON:

MESSRS. LONGMANS, GREEN, AND CO,

PATERNOSTER ROW.

Dy Je teeth

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

1912,

COUNCIL.

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

Tue Earu or Auramont, F.S.A.

Sir Jonun Rosz BrapFrorp,

KaCIVEGE CF NED..4 “D:Se5

F.R.S., Vice-President.

Ricnarp H. Burne, Esq.,

M.A.

Lt.-Col. Sm R. Havenock

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

ALFRED H. Cocks, Esq., M.A.

F. G. Dawrrey Drewirt, Esq.,

M.A., M.D.

Cuartes DrumMonp, Esq.,

Treasurer.

Sir Epwarp Duranp, Bt., C.B. —

F. Du Cant Gopman, Esq.,

IDOM Ue. IDLIB SS

Sirk Epmunp G. Lopsr, Bt.,

Vice- President.

EK. G. B. MeapE-Watpo, Esq.,

Vice-President.

Prof. Epwarp A. Muncuin,

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

P. Cuatmers Mircuett, Esq.,

MeAS DSc Hon, LL.D;

E.R.S., Secretary.

W. R. Ocitvie-Grant, Esq.

ALBERT Pam, Esq.

Aprian D. W. Pottocr, Esq.

OLDFIELD THomas, Hsq., F.R.S.

AntHony H. WINGFIELD,

Ksq.

A. Smita Woopwarp, Ksqa.,

LL.D.,F.R.S., Vice-President.

Henry WoopwarD, Ksq., LL.D.,

F.R.S., Vice-President.

PRINCIPAL OFFICERS.

P. Cuatmers Mircueny, M.A., D.Sc., Hon. LL.D., F.R.S.,

Secretary.

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

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

Resident Superintendent of the Gardens.

D. Sers-Smiru, Curator of Birds and Inspector of Works.

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

Epwarp G. BouLencEr, Curator of Reptiles.

F. H. Warernouss, A.L.S8., Librarian.

JoHN Barrow, Accowntant.

W. H. Conn, Chief Clerk.

LIST OF CONTENTS.

1912, pp. 505-913.

EXHIBITIONS AND NOTICES.

' The Secrerary. Report on the Additions to the Society's

Menagerie during the months of February and March

1) ce es GRRE Gata a mS PRANAB REGS ee at ge

Mr. R. I. Pocock, F.R.S., F.L.8., F.Z.S. Exhibition of

lantern-slide of two Polar Bear cubs (Ursus maritimus).

bonniintatlnen Gama ergy ease see lt cet a

Mr. C. Tare Recan, M.A., F.Z.S. Exhibition of lantern-

slides of a Fllatfish (Platophrys podas), showing its

power of changing its colour and markings...............

Dr. R. W. SHuretpt, C.M.Z.8. Exhibition of skins of two

Virginia Opossums (Didelphis virginiana) ............... 5

Mr. D. Sera-Suiru, F.Z.S. Exhibition of lantern-slides of

the display of the male Peacock Pheasant (Polyplectron

chinquis). Also of photographs of the young Cariama

eristata hatched and reared in the Gardens. (Pl. LXITX.) £

The Secrerary. Exhibition of a living specimen of a young

female Dorsal Hyrax (Dendrohyrax dorsalis) ............

Page

5D5

556

The Secretary. Exhibition of photographs of an Elephant

Arcee aS SWANN ot ie tee a'e Viele <2 ols «lS asc ee ee ere

Mr. R. I. Pocock, F.R.S., F.L.S., F.Z.S. Exhibition of a

skin and a living specimen of a fawn variety of the

Brown Rat(Lpumys norvegicus) <1. 5.5 .ceeee se ee eee

Mr. D. Seru-Surru, F.Z.8. Exhibition of epidermal sheaths

shed from the base of the lower mandible of the King

Penguin (Aptenodytes pennanti) .............0scssaeeeeees

Dr. Francts Warp, F.Z.S.. Demonstration, illustrated by

lantern-slides and the cinematograph, of a method of

observation of fishes, birds, and mammals under

SEALS cdi ey easter ta Samm aie ccc Siaodecaelatole svn cahcitnx aiejavarelase hie rete ee nnicteeene

The Secretary. Report on the Additions to the Society’s

Menagerie during the month of April 1912...............

Mr. A. Buayney Perctvat, F.Z.S. Exhibition of photographs

and lantern-slides of Game Animals from British East

DAVE Tel Celene Mere ie cern gh thn ce See LiMRG os mee ER IC ole EINEM ees

Mr. D. Seru-Smirn, F.Z.S. Exhibition of two living

specimens of the Lory (Calliptilus solitarius), from

Ti deras4aacSeinGoaoes 6 soo caeceactooscodgecanarsbeantastonalsonn

Mr. G. A. Bounencrr, F.R.S., F.Z.S. Notice of a paper

entitled “Second Contribution to our Knowledge of

the Vieetles OL baer Vella wate 2 emnee ane eee ees

671

806

807

Mr. E. G. Boutencer, F.Z.S. Exhibition of a clay-ball |

containing a cocoon of the African Lung-fish (Proto-

SDLCTUSAIEMECLCTES) 2c Sashes) e eee epee eee eee eee Rey eres

Mr. E. G. B. Mrape-Watpo and Others. Discussion on the

Preservation of the Native Fauna of Great Britain ...

The Secretary. Report on the Additions to the Society’s

Menagerie during the months of May, June, July,

Aqeush, and Sepbember, 1912). leat. )-4-cneeoeeee

Mr. R. Lypexxer, F.R.S., F.Z.S. Note stating that

“ Gazella hayi” = 6 Palle JUSCLIROMSE A.B ike eemanacent

908

911

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

of a memoir on the Crustacea Isopoda collected by the

‘Porcupine’ Expedition in 1869-170 ..................

Mrs. Rosz— Hare Tuomas, F.Z.S. Exhibition of, and remarks

upon, the Kegs of Phasianus formosanus, P. versicolor,

and their F.1 and F. 2 offspring. (Text-fig. 126.) ...

Sir Epmunp G. Lover, Bt., V.P.Z.8S. Demonstration of the

capacity of the electric lantern presented by him to the

DOCIEC VE Sassen eer ee Nee tae en, 5. 150 4 Mie a ae ae,

PAPERS.

26, Observations on some Alcyonaria from Singapore, with

a brief Discussion on the Classification of the Family

Nephthyide. By Epwarp W. Suann, B.Sec., De-

monstrator and Assistant Lecturer in Zoology in the

Victoria University of Manchester. (Pls. LXI.-

ATE ROID TANS ian as shicladia nttanicft eth acs ctu ap Remade baited dao ta

27. Some early Fossil Cirripedes of the Genus Scalpellum.

By Tuomas H. Wiruers, F.G.8. (‘Text-figs. 64 & 65.)

28. Experimental Pheasant-breeding. By Rosz Hate

MONAS, HA Ses (els. Wax) eV oe eto t sees

29. A List of Moths of the Family Pyralide collected by

Felix B. Pratt and Charles B. Pratt in Dutch New

Guinea in 1909-10; with Descriptions of new Species.

By Sir Grorce H. Kenrick, F.Z.8. (Pl. LXVIII.) .

30. Ona rare Stag (Cervus wallichii) from Nepal recently

presented to the Zoological Society by His Majesty

King George. By R. I. Pocock, F.R.S., F.L.S., F.Z.8.,

Superintendent of the Gardens. (Text-figs. 66-71.)...

Page

912

913

528

546

558

31.

33.

3D.

38.

39.

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—IV. On a Species of Jnermi-

capsifer from the Hyrax, and on the Genera Zschokkeella,

Thysanotenia, and Hyracotena. By Franx KH.

Bepparp, M.A., D.Sc., F.R.S., F.Z.8., Prosector to

the eciety.- (Pext=es. 7 2-83.) eo sec cccsepenmene ee eae

. Additional Notes on the Living Specimens of the

Australian Lung-fish (Ceratodus forsteri) in the Col-

lection of the Zoological Society of London. By

Basurorp Dean. (Text-figs. 84 & 85.) ................5-

The Circulatory System of the Common Grass-Snake

(Tropidonotus natrix). By CHas. H. O’Donoeuus,

B.Sec., F.Z.8., Assistant to the Jodrell Professor of

Zoology, University College, London. (Pls. LXX.-

OOM Leama lextatins. 86-201.) cicews cnscepeaseocee nearer

. A First Account of the Courtship of the Redshank

(Totanus calidris). By Jutian 8. Huxvey, Lecturer

of Balliol (CollesesOxtordl ose. testes eee nese secre

Some Brackish-water Amphipoda from the Mouths of

the Weser and the Elbe, and from the Baltic. By

KE. W. Sexron, Marine Biological Laboratory, Ply-

mmouuthe GENS: AaXOX TIM, “ds XO XSW). 2. Seats eaenctsy ice

. Descriptions of new Fishes of the Family Loricariide in

the British Museum Collection. By C. Tare REGAN,

NE As RZS 2 CPI. ALO VC SOV eee

. On a Collection of Fishes made by Mr. A. Blayney

Percival in British East Africa to the East of Lake

Baringo. By G. A. BouLencer, F.R.S., F.Z.8. (Pls.

WEE ORO V2 ont elcies. Sctisleiets 5h test eae

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—V. On a new Genus (Dasyuro-

tenia) from the Tasmanian Devil (Dasyurus ursinus),

the type of a new Family. By Frank E. Brpparp,

M.A., D.Se., F.R.S., F.Z.S., Prosector to the Society.

(Hextafigs. G21 Oi): 0... Sa. s.0s See eeereeee neater eee

Studies in the Fossorial Wasps of the Family Scoliide,

Subfamilies Elidine and Anthoboscine. By RowLanp

E. Turner, F.Z.8.,F.E.S. (Pls. LXXXI-LXXXIIT.)

Page

576

607

656

666

677

696

40.

4].

42.

43.

44,

45.

46.

47.

48.

49.

vil

Notes on the Spanish Ibex. By Aspen Cuapman, F.ZS.

The Local Races of Burchell’s Zebra. By Major J.

Stevenson-Hamitton, C.M.Z.S., Game Warden of the

Mransvaale | (Vext-fies. NO2-V0G, ae es eco es css. esas cee

On Dipteropeltis, a new Genus of the Crustacean Order

Branchiura. By W. T. Cauman, D.Sc., F.Z.8. (Pl.

TREES, ER esse? NSE oie ee Mon cr

On two new Larval Trematodes from the Striped Snake

(Tropidonotus ordinatus sirtalis). By W1uLtam Nico.t,

M.A., D.Sec., M.D., F.Z.S., Lister Institute of Pre-

ventive Medicine, London. (Text-fig. 107.) -...........

A Note on the rare British Nudibranch Hancockia

eudactylota Gosse. By Sir Cuarues Exiot, K.C.M.G.,

Gib Zs.) (PIE eX XV |)

Ce ce i i i ie i ie i i a airy

The North Rhodesian Giraffe. By R. LypEKxxkesr,

EIS Seg He Za cb let CRONE VAI) eo Ray US od

On Antler-Growth in the Cervide, with special reference

to Llaphurus and Odocoileus (Dorcelaphus). By R. I.

Pococs, F.R.8., F.L.8., F.Z.8., Superintendent of the

Gardens and Curator of Mammals. (Text-figs. 108-

TR teeta to Na at. cts sik ciaicds ccus ing vicene digs alta ei sore legac asia Paes

Polycheeta from the Pacific Coast of North America.—

Part I. SerpuLip#, with a Revised Table of Classi-

fication of the Genus Spirorbis. By Heten L. M.

PrxeLt, B.Sc., F.Z.S., Demonstrator of Zoology and

Reid Fellow, Bedford College, University of London.

(LEAS) ODO HTS OO, DG) BB arrears See eee yo

The One-sided Reduction of the Ovaries and Oviducts

in the Amniota, with Remarks on Mammalian Evo-

lution. By Hans Gapow, M.A., Ph.D., F.R.S., F.Z.S.

Contributions to the Anatomy and Systematic Arrange-

wnent of the Cestoidea.—VI. On an Asexual Tapeworm

from the Rodent, /iber zibethicus, showing a new form

of Asexual Propagation, and on the supposed Sexual

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

F.Z.S., Prosector to the Society. (Text-figs. 113-121.)

Page

754

757

763

767

770

(71

ees

784

808

822

Vill

Page

50. On two new Trematode Parasites from the Indian Cobra.

By Wituram Nicott, M.A., D.Sc., M.D., F.Z.8., Lister

Institute of Preventive Medicine, London. (Text-

BS MO roc cOb opiate eases Medes tile Slee Seu Nese Ace a Re 851

D1. Statistical Note on the Worm Parasites collected from

the Animals dying in the Zoological Gardens from

December 1910 till April 1912. By Wituram Nico,

M.A., D.Se., M.D., F.Z.S., Lister Institute of Pre-

ventive Medicine: Mondon ...).:.2: aeeceseeeseenaes eee 856

52. On some new Fossil Reptiles from the Permian and

Triassic Beds of South Africa. By R. Broom, D.Sc.,

COMES =) a (bls. UEC — XCM.) 20. cee ice eer 859

53. On the Hydrocoralline Genus, Hrrina. By Professor

S. J. Hickson, F.R.S., F.Z.8., The University of

Meamchesters (Ris XC DVs IO Wale) ial .e aan eee 876

04. Descriptions of new Butterflies of the Genus Thecla

from §.E. Brazil. By E. Duxryrietp Jones, F.Z.S.,

Stee eis @ eee. CADW ES) Ne ed an a abe Mgt, eed elyeey ea a 896

50. The Bornean Bantin. By R. Lypexxer, F.RS., F.Z.S.

(Le seC HGS es Slit. Sea MANOEL Lele, BAe TE eal 902

56. Notes on the Breeding of the “ Millions” Fish (Gir-

-ardinus peciloides). By Epwarp G. BouLEncEr,

REZ. S es Curatoriotaheptiles misc): 26. tae. ssa, 906

Alphabetical List of Contributors ......0.......000eceeeceeeeeee 1x

New ‘Generie Perms 70. Stee hi ce CIR Oh ae ee XVI

Index’ of Serentific. Names 05a ee ee XVil

ALPHA BET UC A lg iiss.

CONTRIBUTORS,

With References to the several Articles contributed by each.

(1912, pp. 505-913.)

BEppDARD, Frank H., 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—IV. On a Species of Jnermi-

capsifer from the Hyrax,and on the Genera Zschokkeella,

Thysanotenia, and Hyracotenia. (Text-figs. 72-83.) ... 576

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—V. On a New Genus (Dasywro-

tenia) from the Tasmanian Devil (Dasyurus ursinus),

the type of anew Family. (Text-figs. 92-101.)............ 677

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—VI. On an Asexual Tapeworm

trom the Rodent, /iber zibethicus, showing a new form of

Asexual Propagation, and on the supposed Sexual Form.

(TSS ie tS) 13) 2a (0 AR oe ote ir ME LS 822

Page

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

Exhibition of a clay-ball containing a cocoon of the

African Lung-fish (Protopterus annectens) ........ eae 807

Notes on the Breeding of the ‘ Millions” Fish

(Gurarduniis peccrloides) tii 6. 25.5.c0. ee eee ae ee eee 906

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

On a Collection of Fishes made by Mr. A. Blayney

Percival in British East Africa to the East of Lake

JE esommalegoy, (led ise: JDO: Qi Se BDO RE Are enon ag ncunaco: 672

Notice of a paper entitled “Second Contribution to

our Knowledge of the Varieties of the Wall-Lizard” ... 807

Broom, Ropert, D.Sc., C.M.Z.S8.

On some new Fossil Reptiles from the Permian amd

Triassic Beds of South Africa. (Pls. MCX CMM.) ewe 859

Cauman, WiuuiAM T., D.Sc., F.Z.S.

On Dipteropeltis, a new Genus of the Crustacean Order

Branchiura., 9 aX XT Via) ae eee 2 ha ae nee ad 763

CuapMAN, ABEL, F.Z.S.

Notes onjthe Spanish Wibex: g.c.accu. ceeneecee ele eee ee 754

Dean, Basurorp, Ph.D.

Additional Notes on the Living Specimens of the

Australian Lung-fish (Ceratodus forsteri) in the Col-

lection of the Zoological Society of London. (Text-

Hig OA BO.) 2. dana aerckigtinal | eae ee ee 607

Eutot, Sir Cuartes, K.C.M.G., C.B., F.Z.S.

A Note on the rare British Nudibranch Huancockia

eudactylota, Gosse,.. (Pl UX X XV.) wsssescpececssderesesenees 770

Gapow, Hans, M.A., Ph.D., F.RB.S., F.Z.S.

The One-sided Reduction of the Ovaries and Oviducts

in the Amniota, with Remarks on Mammalian Evolution.

Hamitron, Major J. Srevenson-. See SreveNson-HAmMIL-

TON, J.

Hickson, Professor Sypney J.. D.Sc., F.R.S., F.Z.8., The

University of Manchester.

On the Hydrocoralline Genus, Hrrina. (Pls. XCIV.—

BCVA Oy tie be casas Cacat sae eeeeeret eC eS ONS, aca yley BA GLA

Houxtey, Junian S., Lecturer of Balliol College, Oxford.

First Account of the Courtship of the Redshank

(LAGU IRIBIS TETTID G50) | ie A setts, art oe eee ieee om

JonEs, E, DUKINFIELD, F.Z.8., F.E.S.

Descriptions of new Butterflies of the Genus Thecla

from 8.K. Brazil. (Pl. PA OE US Taea aah eae ae bal: Are Paes

Kenrick, Sir Grorce H., F.Z.S.

A List of Moths of the Family Pyralide collected by

Felix B. Pratt and Charles B. Pratt in Dutch New

Guinea in 1909-10; with Descriptions of new Species.

CETTE SeaNATELT ie cee eA aU Se gE ANE 8 UE

Lover, Sir Epmunp G., Bart., V.P.Z.S.

Demonstration of the capacity of the electric lantern

presented, by, him, to the Society -!s 2: see hesc 4 fessseee eee

LypEKKER, Ricuarp, F.R.S., F.Z.S.

The North Rhodesian Giraffe. (Pl. LXXXVL.) ......

The Bornean Bantin. (Text-figs. AO od crunante es

Note stating that “ Gazella hayi” = Gazella fusci-

DRL TINE a xcs, SMe eRe Pre M MEN et snc AE veoh a Rats Bid

Page

808

876

647

896

546

913

xii

Mrape-Watpo, Epmunp G. B., V.P.Z.S., and Others.

Discussion on the Preservation of the Native Fauna

OMCnreabelNGatn =.) ov save oo tet Oe eee ee a oe

Mircuexi, P. Cuatmers, M.A., D.Sc., Hon. LL.D., F.RB.S.,

F.Z.8., Secretary to the Society.

Report on the Additions to the Society’s Menagerie

during the months of February and March 1912 .........

Exhibition of a living specimen of a young female

Dorsal Hyrax (Dendrohyraa dorsalis) .........0.0.cceeeeeee ee

Exhibition of photographs of an Elephant Kraal in

SIGIOIY +) une ania a Secc aaa aR eRRAS ee aan eee ara ees ccm aoaols aan

Report on the Additions to the Society’s Menagerie

dumng ne momph of Avorn MOND Meo: i. ie. aie uth alee

Report on the Additions to the Society's Menagerie

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

Sie IN| NETS, NEO eae onc agosacde ain: cof see Sodass3Indan- sabes

Nicont, Wittram, M.A., D.Sc, M.D., F.Z.8., Lister

Institute of Preventive Medicine, London.

On two new Larval Trematodes from the Striped

Snake (Zropidonotus ordinatus sirtalis). (Text-fig. 107.).

On two new Trematode Parasites from the Indian

Cobra. v(Mexbatie. 22 oe ctlcas ct aden ree ces fen Scceenneree

Statistical Note on the Worm Parasites collected from

the Animals dying in the Zoological Gardens, from

December, 1910 tll A oral POT Ae. ee eee ee ees eee

O’Donocuur, Cuartes H., B.Sc., F.Z.8., Assistant to the

Jodrell Professor of Zoology, University College,

London.

The Circulatory System of the Common Grass-Snake

(Tropidonotus natrix). (Pls. LXX-LXXII. and Text-

AIS, BOO |) 2 sericea ce Me oc ule Sets CaS AREA ARRAS Ae ee

Page

807

806

908

612

Xill

Percrvau, A. Buayney, F.Z.S.

Exhibition of photographs and lantern-slides of Game

Animals trom British Bast Africa <8. Biss... .c. ices cscacs.

Pixein, Miss Heten I. M., B.Sc., F.Z.S., Demonstrator

of Zoology and Reid Fellow, Bedford College,

University of London.

Polychzeta from the Pacific Coast of North America.—

Part I. SerpuLip#, with a Revised Table of Classification

of the Genus Spirorbis. (Pls. LX XXVII-LXXXIX.) .

Pocock, Reeinatp I., F.R.S., F.L.S., F.Z.8., Superintendent

of the Gardens and Curator of Mammals.

Exhibition of lantern-slide of two Polar Bear cubs

(Ursus maritimus) born in the Gardens ................0606+

On a rare Stag (Cervus wallichit) from Nepal recently

presented to the Zoological Society by His Majesty King

Georveene (lemt-tgs: GG (1)... csiie s+ cde teecevcesereossensne es

Exhibition of a skin and a living specimen of a fawn

variety of the Brown Rat (Hpimus norvegicus) ............

On Antler-Growth in the Cervide, with special reference

to Elaphurus and Odocoileus (Dorcelaphus). (Text-figs.

COLSTON Sy hee ee eles ORNs tna eres ie ye eet

Regan, C. Tate, M.A., F.Z.8.

Exhibition of lantern-slides of a Flatfish (Platophrys

podas), showing its power of changing its colour and

Tip ra STS 54: ch anpeeebceEQbeeeces Ob Ar pan) A eS ob pce ce eannoeeneeoe ras

Descriptions of new Fishes of the Family Loricariide

in the British Museum Collection. (Pls. LXXV.-

TONITE icant, coop Brac a en od o5 B55 SECAISSS aba tn coNEIRE IS ar

806

784

556

XIV

Serm-Smirn, Davip, F.Z.S., Curator of Birds.

Exhibition of lantern-slides of the display of the male

Peacock Pheasant (Polyplectron chinquis). Also of

photographs of the young Cariama cristata hatched and

rearegun the Gardens: (CPi. IX UX). oii. tee Pee

Exhibition of epidermal sheaths shed from the base of

the lower mandible of the King Penguin (Aptenodytes

JREIOLOHTOIION: Ge 00 Ore SEER RBRR ORO eraen Coon CAnGet Serisecadhocng e008

Exhibition of two living specimens of the Lory,

Caliiavilins solitarius, LVOML WU}iica...+.:oece sca scnatinecmtencsnile

Sexton, Mrs. E. W., Marine Biological Laboratory,

Plymouth.

Some Brackish-water Amphipoda from the mouths

of the Weser and the Elbe, and from the Baltic.

(Pls: XOX LXV) os. BS coats cae ete

Suann, Epwarp W., B.Sc., Demonstrator and Assistant

Lecturer in Zoology in the Victoria University of

Manchester.

Observations on some Aleyonaria from Singapore, with

a brief Discussion on the Classification of the Family

Nephthiyides., oss gs Xo SGI) er esos veins aie paaneia

Suuretpr, Roperr W., M.D., C.M.ZS.

Exhibitions of skins of, and remarks upon, two Virginia

Opossums (Didelphis virginiand) ......cscscecserenvererecnenes

Smira, Davip Seru-. See Sera-Smrra, D.

Srepprne, The Rev. Tuomas R. R., M.A., F.R.S., F.Z.8.

Notice of a Memoir on the Crustacea Isopoda collected

by the ‘ Porcupine’ Expedition in 1869-1870.........0s000

Page

671

806

656

505

556

Page

SreveNson-Hamitton, Major J., C.M.Z.8., Game Warden

of the Transvaal.

The Local Races of Burchell’s Zebra. (Text-figs. 102-

HOG ieee RSS scm iu ON a cts Meee nce Ms Niwa SUN ka Cok 757

Tuomas, Mrs. Rost Hare, F.Z.S.

Experimental Pheasant- breeding. (Pls. LXIV.-

POV) ones oo ae man atis Hats See eels e ca bce snge te alee 539

Exhibition of the Eggs of Phasianus formosanus,

P. versicolor, and their F.1 and F. 2 offspring. (Text-

SILT AG re ee Made eee Neder. nies Aenean soot stwen CL Aue coins 912

TuRNER, Rowianp H., F.Z8., F.ES.

Studies in the Fossorial Wasps of the Family Scoliide,

Subfamilies Elidine and Anthoboscine. (Pls. LX XXI.—

15 O09 Bg i oars bale eA oe I a aera eine es eee eet 696

Wa po, EK. G. B. Meapn-. See Meapr-WALpo, E. G. B.

Warp, Francis, M.D., F.Z.S.

Demonstration, illustrated by lantern-slides and the

cinematograph, of a method of observation of fishes, birds,

and: matmimals umder waber? <i. .2s....c..seeaces0 sna ceneteeces 672

Wiruers, THomas H., F.G.S.

Some early Fossil Cirripedes of the Genus Scalpellum.

Qhemt-Hes G4 dr Goi). tessa secede cy an Nepean diosa weteceldwnnon 528

NEW

XVI

GENERIC TERMS

PROPOSED IN THE PRESENT VOLUME (pp. 505-913).

Page

Alopecorhinus (Reptilia) ...... 864

Braunsomeria (Insecta)..,...... 697

Brevicella (Insecta) ...........- 549

Dasyuroteenia (Vermes) ......... 694

Dipteropeltis (Crustacea) ...... 762

Emydochampsa (Reptilia)...... 875

Emydops (Reptilia) ............ 871

860

Galeops (Reptilia)

poreceteecesece

eoeeercos

Hyracotenia (Vermes)

Ictidepsis (Reptilia)

eescce

Protoleospira (Vermes)

Scymnognathus (Reptilia)

Taurops (Reptilia)

woes escscessce

Urocystidium (Vermes)

Xenopharynx (Vermes)

PROCEEDINGS

OF THE

|" GENERAL MEETINGS FOR SCIENTIFIC BUSENESS

OF THE

| ZOOLOGICAL SOCIETY

ae OF LONDON.

1912.

Pe LEY,

CONTAINING Paces 505 20 756, witH 23 Pxares

AND 88 TEXT-FIGURES.

SEPTEMBER 1912,

PRINTED FOR THE SOCIETY,

SOLD AT. THEIR HOUSE IN REGENT’S PARK.

a ee. LONDON:

~ MUESSRSALONGMANS, GREEN, AND CO,,

nt 1) oS. PATURNOSTER ROW.

] =

LIST OF CONTENTS,

1912, Part III. (pp. 505-756).

- -

EXHIBITIONS AND NOTICES.

Page

The Secrerary. Report on the Additions to the Society’s eee during the months

of Mepruary-and March; LOU2 econ... suie ciecic'v «eiclne cleisneicie line Rocha SocnmoedS 555

Mr. R. I. Pococs, F.B.S., F.L.8., F.Z.8. Exhibition of lantern-slides of two Polar Bear

cubs (Ursus maritimus) born in the Gardens ...... siateis cis eer daere ain arate sities aise etaleti 556

Mr. C. Tarn Ruaay, M.A., F.Z.S. Exhibition of lantern-slides of a Flatfish (Platophrys

podas) showing its power of changing its colour and markings..........++..>+2++-. 9506

Dy. R. W. Suvuretpr, C.M.Z.8. Exhibition of skins of two Virginia Opossums Sage

BUT GUIADIG) soo. «a eictee 0 «ie esersherriafrietareiiselotees ave Toisesrareraj iwiseae atetta ste eeersceke oieaotoeepeRcreeene -- 606

Mr, D, Sers-Suitu, F.Z.8. Exhibition of lantern-slides of the display of the male Pea-

cock Pheasant (Polyplectron. chinguis). Also of photographs of the young Cariama

eristata hatched and reared in the Gardens. (Pl, LXIX.) ...........0..0ceeneee 557

The Sucrerary. Exhibition of a living specimen of a young female Dorsal Hyrax ae

(Dendrohyra« dorsalis) ......+.+. Neate ails an Bae Hate aioe awe Sas magne eee eesti!

The Secretary. Exhibition of photographs of an Elephant Kraal in Siam .......... 671

Mr. R. I. Pococn, F.R.S., F.LS., F.Z.S. Exhibition of a skin and a living specimen of

a fawn variety of the Brown Rat (Hpimys morvegicus) .. 2 ecescsecsccsececeunce 671

Mr. D. Setn-Suitu, F.Z.8. Exhibition of epidermal sheaths shed from the base of the

lower mandible of the King Penguin (Aptenodytes pennant?) .....e--seeeseeeeene 671

Dr. Francis Warp, F.Z.8. Demonstration, illustrated by lantern-slides and the cine-

matograph, of a method of observation of fishes, birds, and mammals under water. 67

PAPERS.

26, Observations on some Alcyonaria from Singapore, with a brief Discussion on the

Classification of the Family Nephthyide. By Epwarp W. Suann, B.Sc., Demonstrator

and Assistant Lecturer in Zoology in the Victoria University of Manchester, (Pls.

HX ToS NCET DD) ietaiee wie e's! e-ainicisc ais MMe ca calais Biec'e ott te ayn tetatereiere HAL orn. oft onecteun 505

27. Some early Fossil Cirripedes of the Genus Sealpellum. By Tuomas H. Wirmurs, F.G.S.

(Dext-frs; 64:85 OD, ), isrevs save sien sia'r v0 os eos ale 8 ou 0.0 0 ates calele eben \nilsieh iain eiela. Are DES.

Contents continued on page 3 of Wrapper,

P.Z.s97 1012. Flier

vex yy

Grout Intaglio et imp.

ALCYONARIA FROM SINGAPORE.

P.Z.S. 1912. Pl.. LXII.

bite ee

Grout Intaglio et imp.

ALCYONARIA FROM SINGAPORE.

P.Z.S. 1912. Pl. LXIII.

OSS |

$4,"

» bow’

v2 '@ee

Pe Miers

toe.

‘ee,

Grout Intaglio et imp.

E.W.S. del.

ALCYONARIA FROM SINGAPORE.

PROCEEDINGS

OF TIIE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF TILE

ZOOLOGICAL SOCIETY OF LONDON.

PAPERS.

26. Observations on some Aleyonaria from Singapore, with

a brief Discussion on the Classification of the Family

Nephthyide. By Epwarp W. Suann, B.Sc., Demon-

strator and Assistant Lecturer in Zoology in the

Victoria University of Manchester *.

[Received January 17, 1912 : Read March 19, 1912.]

(Plates LXI.-LXIII.+)

IypEx.

Systematic : Page

Sclerophytum pinnulatum, sp. 0. ......s00-02.222.-.- 5OT

Nephihiya Gedfordi, Sp. Wei.... 62:0 -02-<-.sessccscsee seen DIAL

ane TRAE ASU hls een esee chs ees | A

Wiig ntellennovustam sys Dee ae ee) eee eie nn Oey

Nephthya: Historical Summary aes Eis ek AA 510

Dendronephthya do. Saeee ee cs. oe OLS

Stereonephthya do. Se Geeac cseeee OLO

A collection of Aleyonaria, made by Mr. W. F. Lanchester

and the late Mr. F. P. Bedford during their residence in Singa-

pore, was sent to the Zoological Department at the Victoria

University of Manchester. “Professor S. J. Hickson kindly

* Communicated by Prof. S. J. Hickson, D.Sc., F.R.S., F.Z.S.

+ For explanation of the Plates see p. 527,

Proc. Zoou. Soc.—1912, No. XX XTV. 34

506 MR. E. W. SHANN ON

entrusted to me the task of identifying the specimens in the

collection. The collecting was carried on in shallow water around

the coast of Singapore; the precise localities will be cited under

the description of the several species. Several examples of

Nephthyide occur, and this necessitated facing the vexed ques-

tion of the classification of that family; I shall state below my

reasons for adopting the scheme of classification advanced by

Kiikenthal.

I wish to express my indebtedness to Prof. Hickson for his

constant advice as to my procedure and help in the pursuit of

literature bearing upon the subject. In the last-named branch

of the work I had frequent recourse to an exceedingly useful

catalogue of the Alcyonaria prepared by Dr. J. Stuart Thomson.

My thanks are also due to Mr. J. T. Wadsworth for preparing

the excellent photographs from which Plate LXII. figs. 7, 8, & 9

were made.

In all eleven species are described, of which four arenew. The

systematic positions of the several species are as follows :— —

Order ALCYONACEA Verrill (pro parte).

Family ALcyONTIIDA.

Sclerophytum pinnulatum, sp. n.

Family TELESTIDA.

Telesto rupicola F. Miiller.

Family Nerutuyips.

Nephthya bedfordi, sp. n.

Dendronephthya disciformis Kiikenth.

Stereonephthya lutea, sp. n.

Paraspongodes crassa Kiikenth.

Family SrpPHoNOGORGTIDA&.

Siphonogorgia variabilis Hickson.

Order GORGONACEA.

Suborder PSEUDAXONTA.,

Family Scterocoreps.

Suberogorgia suberosa Pallas.

Family Mexiropip&,

Melitodes albitincta Ridley.

Psilacabaria gracillima Ridley.

Wrightella robusta, sp. n.

ALCYONARIA FROM SINGAPORE. 507

Order ALCYONACHEHA.

Family ALCYONIIDA.

Genus SCLEROPHYTUM.

Sc. PINNULATUM, sp. n. (Pl. LXII. fig. 7; Pl. LXIII. fig. 10.)

A single complete colony was taken just below low-tide mark

from Blakang Mati.

In its form and mode of branching the colony closely resembles

Se. palmatum Pratt (1903). As in the last-named species, the

colony (fig. 7) is erect, branched, and shows a marked lateral

compression. The total height is 88 mm., of which the stalk

comprises 45 mm. The diameter of the stalk is 31x11 mm.,

and that of the capitulum is 60x 35mm. The stalk divides at its

distal end into two almost equal primary branches. From these

branches spring numerous secondary lobes which are arranged

in no definite order. The secondary lobes frequently show

lateral compression in their proximal regions, but their termi-

nations are usually bluntly conical, while a few are reniform.

The length of these lobes is 12-20 mm., and their diameter

6-9 mm.

The colour of this specimen in alcohol is slate-grey. The

consistency is tough; the stem is hard and brittle, but the

secondary lobes are soft and fleshy.

The autozoids are of a brown colour and are uniformly scattered

over the surface of the secondary lobes. The average distance

between two adjacent polyps is‘6 mm. Autozoids are also found

on the primary branches almost down to the level of the stalk,

but in this region the distance between them is very much greater,

viz. 3-4 mm. Many of the autozoids are more or less expanded,

but none of them are situated on raised rounded areas such as

occur in Se. palmatwm. The anthocodie are of medium size

when compared with those of the genus Sclerophytum as a whole ;

the diameter of an expanded crown is 1 mm., but the polyp-heads

appear only half that size when contracted. :

The tentacles (fig. 10) form the distinctive feature of this

species. They are ‘55 mm. in length, compressed, and expanded

at the distal end. There is a single row of free well-developed

pinnules on each side of a tentacle. The pinnules are -(06 mm.

in average length, but the larger ones attain a length of -10 mm.

. and a breadth of (035 mm, The larger pinnules are those which

are placed third, fourth, and fifth from the distal end; they

become shorter as they approach the base of the tentacle. There

are about twelve pinnules in each row.

The stomodeum is long and convoluted. The mesenteries

and mesenterial filaments are well marked. No sexual organs

were observed.

The siphonozoids, if present, are extremely degenerate. <A

very few minute diverticula of the superficial canals, which point

in the direction of the surface, but never open to the exterior,

34*

508 MR. E. W. SHANN ON

may represent rudimentary siphonozoids. The superficial canals

form a dense network just beneath the surface of the colony ;

their diameter is ‘(044-120 mm. ‘The vessels of the internal

system are clearly defined and circular in cross-section, and are

fairly numerous.

Zoochlorelle are numerous in the superficial canal-system, and

occur in less abundance in the interior. They are also found in

the endoderm of the polyps, and extend into the tentacles,

occupying even the lumen of the pinnules.

The spicules are quite characteristic of the genus Sclerophytum,

and indeed only differ from those of Sc. palmatwm in almost

insignificant detail. The little knotted clubs average ‘16 x °034 mm.,

and the spindles are sometimes only -058x'015 mm. These

minute spicules only occur immediately beneath the surface.

The spicules of the ccenenchyme are fairly numerous and are all

of the tuberculate warted type; they vary considerably, however,

in shape and size. The majority are spindles which narrow

rapidly towards their ends to rather acute points; their measure-

ments range from ‘4x‘ll mm. to 30x45 mm. Ivrregularly

branched forms are by no means uncommon.

The following considerations are advanced as an apology for

the creation of a new species of Sclerophytwm based on the

examination of a single specimen.

This colony had been assigned in a preliminary investigation

to the species Sc. palmatum Pratt (1903), and at first sight this

diagnosis appeared perfectly accurate. The external appearance,

and indeed the actual measurements, agree closely with the

description of thetype specimen. The characters and distribution

of the spicules, the well-marked mesenteries and mesenterial

filaments, the rudimentary condition of the siphonozoids, the

orientation of the canal-systems, and the distribution of the

zoochlorelle, all tend to enhance the resemblance between the

two species. It is only when the autozoids are examined that

the true specific difference is realized; those of Se. palmatwm are

distinctly larger than those of Se. pinnulatwm. The characters

of the tentacles which form the essential divergence of the two

species are tabulated below :—

Sc. palmatum Pratt. Sc. purnulatum, sp. n.

‘7 mm. in length. -55 mm. in length.

Almost of uniform length. Expanded at distal end.

Possess a double row of rudi- A single row of free well-deve-

mentary pinnules down loped pinnules down each

each side. side.

Sc. pinnulatum could be confounded with no other species of

Sclerophytum. At the same time, it is of interest to note that a

single row of free pinnules has been recorded on either side of

the tentacles of Se. viride by Thomson and Henderson (1906).

The possession of free pinnules, now recorded for two species of

ALCYONARIA FROM SINGAPORE. 509

Sclerophytum, tends to strengthen the relationship of the genera

Sclerophytum and Xenia which has been suggested by Pratt

(1903).

Order ALCYONACEA.

Family TELESTID2.

Genus TELEsro.

T. nuricoLaA F.. Miiller.

Carijoa rupicola F. Miiller, Arch. Naturg., Jg. 33, 1867, p. 33,

tab. 9. figs. 56 & 57.

Telesto (Carijoa) rupicola Wright & Studer, ‘ Challenger’

Reports, Zool. vol. xxxi. 1889, p. 262.

Telesto rupicola May, Jena Zeitschr. Naturw. vol. xxvi. 1900,

p. 98.

Telesto rupicola Hickson & Hiles, Willey’s Zoolog. Res. 1900,

p. 496, tab. 50. figs. 1 & 2.

Telesto rupicola Thomson & Henderson, Marine Fauna of

Zanzibar, 1906, p. 434.

Telesto rupicola H. Laackmann, Zoolog. Jahrb. Supp. 11,

Heth 1h 1908) pa si, Paty 2. fies. \; 2); Wat. 3. fig. 3.

Although the only species of Yelesto previously recorded from

Singapore is Z’. prolifera v. Koch, the numerous small colonies in

this collection appear to bear a closer resemblance to 7’. rwpicola,

and are therefore described under that name.

The largest of these colonies is of a grey hue. Its longest

axial polyp measures 85 mm. in length, it is 3 mm. in diameter

at the base and 1:25 mm. at the top. It bears six lateral polyps,

the longest of which measures 24 mm. Anthocodiz arise at

frequent, but pretty regular intervals, both from the axial and

lateral polyps; their average length is 3 mm., and breadth 1°5 mm.

The majority of the other specimens are pale yellow and of

smaller size. All these forms were obtained in shallow water

near Singapore. Some of the exact localities read as follows:

Pulo Brani, 6 fms.; Pulo Brani, 5-10 fms.; Blakang Mati, below

low tides; Tanjong Pagar, 10 fms.

Previously recorded from Rio de Janeiro (F. Miller); Brazilian

coast (Munich Museum); Blanche Bay, New Britain (Hickson

& Hiles); Zanzibar (Thomson & Henderson); Bahia (‘ Chal-

lenger'’).

The spicules show a very wide variety of forms. Not only is

this the case, but a different selection of spicules was found in

each of the five specimens examined.

One specimen contained spicules very much resembling those

of 7’. rvisei. Indeed, the two species are probably very closely

related, for Laackman (op. cit. pp. 72 & 82) is at some pains to

distinguish between them. It should be remembered, however,

that 7’. riisei has not yet been recorded from the Old World.

A very small specimen of a pale yellow colour was examined

510 MR. E. W. SHANN ON

and found to bear many points of resemblance with 7’. prolifera ;

the spicules, for instance, agree well with those figured by

Laackmann for that species (op. cit. p. 87), and the stem walls

ave very thin, At the same time, it was taken from the same

locality (Pulo Brani) as one of the larger specimens which wn-

doubtedly belongs to the species 7’, rwpicola. Moreover, another

specimen was intermediate in every particular between the

above exceptional form and the larger examples, which had been

assigned without difficulty to 7’. rupicola. As this small specimen

was evidently very young, one would not wish to attach too deep

a significance to the observations made or to draw any hasty

conclusions from them. At the same time, it is well within the

range of possibility that the accumulation of such knowledge

may lead eventually to a reduction in the number of species of

Telesto.

An Historical Summary of the Genera Vephthya, Dendronephthya,

and Stereonephthya, with reasons for retaining the definitions

of these genera of the family Nephthyide as enumerated

by Kikenthal.

Genus Neruruya Savigny.

1817. Nephthée Savigny, Descr. de Egypte, Hist. Nat. Suppl. i.

Atlas, Polypes, tab. 2. fig. 5.

1828. Vephthe« Audouin, Explication sommaire des Planches de

polypes de l’Egypte et de la Syrie, publi¢es par Jules-

César Savigny dans: Description de l’Kgypte, vol. xxii.

Paris.

1834. Vephthyo Ehrenberg, Die Corallenthiere des Rothen

Meeres, p. 284.

1846. Vephthya Dana, ‘ Zoophytes,’ Philadelphia, p, 610.

1857. Nephthya Milne-Edwards, Histoire Naturelle des Coral-

laires, vol. i. p. 127.

1877. Nephthya Klunzinger, Die Korallthiere des Rothen Meeres,

Th. 1, p. 33.

1887. Nephthya Studer, in Arch, Naturg., Jg. 53, vol. i. pp. 19,

20.

1887. Vephthy« (pars), Danielssen, in Norske Nordhavs Exp.

vol. v. p. 82.

1889. Nephthya and Spongodes (pars) Wright & Studer, ‘Chal-

lenger’ Reports, Zoology, vol. xxxi. p. XXv.

1895. Spongodes (pars) Holm, in Zool. Jahrb., Bd. 8, p. 24.

1895. Spongodes (pars) Kiikenthal, in Zool. Anz., Jg. 18, p. 428.

1896. Vephthya WKiikenthal, in Abh. Senckenb. Ges. Frankfurt,

vol. xxiii. pp. 89-91.

1899. Nephihya May, in Jena Zeitschr, Naturw. vol. xxxiil. p. 156.

1903. Nephthyw Kiikenthal, in Zool. Jahrb., Bd. 19, p. 141.

Tn 1817 Savigny described two genera of Nephthyide to which

he gave the names Ammothée and Nephthée.” Audouin (1828),

ALCYONARIA FROM SINGAPORE. 511,

to whom fell the task of describing Savigny’s plates, believed

that Savigny’s Tab. 1. fig. 8 represented Ammothée, and that

Tab. 2. figs. 5 & 6 represented Mephthée. he genus Nephthea,

as the author wrote it, was recognised by Ehrenberg (1834); but

at the same time he disputed the interpretation of Savigny’s

plates, maintaining that Audouin had given the name Vephthea

cordierit to the form, represented in Tab. 2. fig. 6, which Savigny

had intended to cail Anumothée. Ehrenberg’s view has been

accepted by all subsequent authors, and it is now generally agreed

that Tab. 2. fig. 5, correctly designated by Audouin Vephthea

chabrolit, represents the type of Savigny’s genus Vephthée, while

Tab, 2. fig. 6 represents his type of the genus Ammothée, namely

A, virescens. There can be no reasonable doubt with regard to

the authenticity of origin of the genus Vephthya, that is to say,

that the genus was based on the description of the species

N. chabrolii, which is figured in Savigny’s Tab. 2. fig. 5. In the

case of Ammothée, or Ammothea as the genus was known for

many years, the name was changed to Lithophytwm by Kiikenthal

(1903), since that author found that Savigny’s type species,

A. virescens, is identical with a form described forty-two years

previously by Forskil under the name Lithophyton arboreum ; in

deference to the International Rules of Zoological Nomenclature

the older name must be retained. Thus it is of little moment

whether or not Ehrenberg was justified in disputing Audouin’s

interpretation of Tab. 2. fig. 6.

_ Copies of Savigny’s plates are extremely scarce, so that it is

not always possible for the research worker to examine the

original figures; many have probably been compelled to content

themselves with descriptions by other authors. With this

difficulty in view, Professor Bourne very kindly had photographs

taken for Professor Hickson from Savigny’s plates, Tab. 2.

figs. | & 6, in the Radeliffe Library at Oxford. Prof. Hickson

has given me permission to publish these figures in this paper,

so that they may be readily accessible to all workers on the

Nephthyide. They are reproduced in Pl. LXI. figs. 1-5 and

Pl. LXITI. fig. 6.

Ehrenberg (1834) distinguished Vephthya from Ammothea by

the prominence of the polyp-spicules in the former genus, for he

says of Nephthya:—“ polypis in verrucas inermes retractilibus.”

We see, then, that Khrenberg recognised the distinction between

the genera Nephthya and Lithophytwm (Anmothea) which obtains

at the present day, namely the presence and absence of armed

polyps (polyps with “Stiitzbiindel ”) in these genera respectively.

Khrenberg’s definition of Vephthya was recognised by Dana (1846),

Milne-Edwards (1857), Klunzinger (1877), Studer (1887), and

Danielssen (1887). The numerous new species described during

this period were distinguished by their authors, on the one hand

from Ammothea by the presence of armed verruce, and on

the other from Spongodes Less. by the comparatively slight

development of the spicules which formed the armament of

512 MR. E. W. SHANN ON

the verruce. Wright and Studer (1889) described a specimen

under the name Spongodes nephthyceformis, concerning which

they observe :—‘“‘ The entire habit of the colony recalls much

more that of Nephthya than that of Spongodes, and this im-

pression is strengthened by the slight development of the spicules

surmounting the little heads, whence the colony does not appear

so prickly as other species.... The species must be referred to

the genus Spongodes, because the polyps are placed sideways

within a bundle of spicules, although these only project slightly.”

That is to say, Wright and Studer recognised on the anthocodiz

of Sp. nephthyeformis the presence of what is now known as a

“ Stiitzbiindel,” and here we have the starting-point of the

difficulty of discriminating between the genera Spongodes and

Nephthya. Holm (1895) faced the problem of reconstructing

the genus Spongodes in the light of the knowledge which had

accumulated since Lesson first described the genus in 1834. —

This author pointed out that Spongodes nephthyeformis W.& St.

is identical with Nephthya chabrolii Audouin, and added that

NV. chabrolii differs from Spongodes in many characters, such as

the branching of the colony and the arrangement of the polyps:

on these characters one can establish two genera, but it is

necessary then to add to the genus Wephthya many species

hitherto included in the genus Spongodes, including the type

Sp. celosia. Though Holm shrank from submitting a well-known

type like Spongodes celosia to such treatment, he proceeded

fearlessly to include all the species of Mephthya, including the

type WV. chabrolii, within the genus Spongodes; he retained

Nephthya, however, as a subgeneric title. Here was a step in

the direction of elucidation; Wephthya and Spongodes;as hitherto

defined, were shown to be synonymous; but Holm’s solution of

the problem threw too great a burden on Spongodes. Kiikenthal

(1895), writing during the year in which Holm’s paper was

published, accepted the genus Spongodes in its new distended

form; but the term Spongodes had become so obviously cumber-

some that this author (Kiikenthal), in a later paper *, reinstated

Nephthya with full generic honours. In this paper Kukenthal gave

a summary of the family Nephthyide, and divided the various

genera into two groups as they possessed or lacked a “ Stiitz-

bindel”; he summed up his remarks as follows :—“ Innerhalb

der Familie der Nephthyiden ist als wichtigstes Merkmal zu

betrachten, ob die Polypenképfchen terminal auf ihrem unteren

Teile, dem Stiele, sitzen oder seitlich davon. Letzterer Fall tritt

stets dann ein, wenn sich auf einer Seite, der oberen, ein Bindel

Spicula besonders stark entwickelt : das Stiitzbiindel.” The

genera possessing a “‘ Stiitzbiindel” were distinguished from one

another by the disposition of the polyps on the colony, as Holm

(1895) had already suggested; the name WVephthya was applied

to forms resembling the original type JV. chabrolii Audouin, in

which the polyps are collected on branchlets, the latter being

arranged in catkins or lappets, and the name Spongodes was

* Abh. Senckenb. Ges. Frankfurt, vol. xxiii. p. 88 (1896).

ALCYONARIA FROM SINGAPORE. 513

retained for forms in which the polyps are disposed sporadically

or in bundles. The net result of this reformation was the

inclusion in the genus Vephthya of all forms hitherto included in

the “ Spicate” group of the genus Spongodes.

In adopting this means of classification it became necessary

to include Sp. celosia, Lesson’s type of the genus Spongodes, in

the emended genus Vephthya; nevertheless, the name Spongodes

was retained by Kikenthal for his other emended genus, since

the latter included a large number of forms which during many

years had been described under Spongodes. The definition of the

genus Wephthya given in the paper under consideration runs

thus :—‘‘ Nephthyiden mit ‘ Stiitzbiindel.’ Die Kolonie ist

buschig veriistelt, die meist Kurzen und nur vereinzelt sterilen

Stammteile sind durch einen abgeflachten, oft membranosen

Basalteil verbunden. Die Polypen stehen in grosser Zahl und

zeimlich gleichmissig verteilt auf den Steinzweigen, die dadurch

die Form von ahrenformigen Lappen oder ‘ Katzchen’ erhalten.

Hervorragende Spicula der Polypenkopfchen fehlen.” May (1899)

accepted the foregoing definition of the genus Vephihya, aud in

his * Revision of the Nephthyide’ (1903) Kiikenthal has not had

occasion to modify it.

A “ Stiitzbiindel.”

The crucial point in Kiikenthal’s classification of the Neph-

thyide lies in the definition of the term “ Stiitzbtindel.” Much

of the: opposition to the above classification has arisen through

the different interpretations which various authors have attached

to the term. Kiikenthal, to whom we owe the word, uses it in

an extremely comprehensive sense, the range of which can be

circumscribed, however, by the following limits :—

A “Stttzbundel” is an aggregation of spicules disposed along

the abaxial aspect of an anthocodia and lying approximately

parallel to its axis. The spicules are usually spindle-shaped ;

they are not infrequently larger than those from any other

portion of a given specimen, and one or two of them commonly,

but not invariably, reach from the apex of the polyp-stalk into

the substance of the colony. A few of the spicules in a

characteristic example protrude beyond the polyp-head, but

such a condition is not essential.

A definite “Stiitzbiindel” may not be recognisable in every

polyp of a given colony; but if such is present it will appear

most obvious in the younger polyps near the distal ends of the

branches. A specimen in which a “ Stiitzbiindel” is demonstrable,

whether or not in every polyp, must be classified as possessing a

“ Stutzbiindel.”

In the genus Lithophytum, which closely resembles Vephthya

in external appearance, the anthocodie being massed on small

terminal lobes or lappets, there is no ‘“Stiitzbiindel.” The

anthocodiz contain very few spicules, some of which are loosely

arranged en chevron along the abaxial surface. Both the small

size and oblique position of the spicules so arranged prevent their

being described as forming a “ Stiitzbiindel.”

514 MR. E. W. SHANN ON

NEPHTHYA BEDFORDI, sp. n. (PI. LXII. fig. 8; Pl. LXITI.

figs. 11, 12.)

Two specimens which, while conforming with the characters

described above for the genus Vephthya, fail to agree in detail

with any of the large number of forms hitherto described, have

necessitated the creation of a new species of this genus.

The colony (fig. 8) is bilaterally compressed, the growth is

bushy, and the major diameter of the capitulum is approximately

equal to the total height, including the stem. The consistency

is tough and leathery. The stem, which shows signs of bilateral

compression, is short, and gives rise at its distal extremity to a

variable number of main branches. These main branches are

again divided into unequal secondary branches. From both

main and secondary branches spring the short terminal lobes.

The latter are conical in shape, but rounded; on them the

anthocodie are tolerably evenly distributed. As the polyps are

situated very close together, and the terminal lobes are ex-

ceedingly numerous, the whole capitulum appears to be covered

with anthocodie. The polyp-heads when at rest make an acute

angle with their stalks. The latter scarcely protrude from the

colony, with the result that the polyp-heads are very closely

apposed to the surface from which they arise.

Colour in alcohol cinder-grey, polyps brown.

Locality : below low-tide mark, Blakang Mati.

Detailed measurements :—

Region measured. Specimen I. Specimen II.

otal hete lit yor scceaeeen er 60 mm. 42 mm.

eieht of Stem cee ecoccs dc 17 mm. 14 mm.

Diameter of stem ............ 18 x 10 mm. 12 x 10 mm.

Height of capitulum......... 43 mm. 28 mm.

Breadth of capitulum ...... 52 x 20 mm. 45 x 20 mm.

In both specimens the length of the terminal lobes is about

4 mm., but the variation in this respect is between 3 and

6mm. The diameter of the terminal lobes is from 3 to 4mm.

The edges of adjacent polyps are seldom more than "175 mm.

apart. The polyp-heads have an average length of 1:1 mm. and

diameter of *6 mm.

The spicules show a wide range in size and shape. The

smallest forms are found in the tentacles, where they are

arranged in no very definite order, but for the most part lie at

an acute angle with the axis of the tentacle. The tentacle-

spicules are minute spiny spindles, some of which are straight,

others ecrescentic. The polyp-spicules (fig. 11, /) are very brittle,

longitudinally striated spindles. They are straight. or curved,

and usually smooth, but sometimes bear a few minute spines.

It is difficult in most cases to discern an arrangement of the

polyp-spicules en chevron, but some of the polyps show such a

condition more clearly than others.

ALCYONARIA FROM SINGAPORE. 515

The “Stiitzbiindel,” though undoubtedly present, is very ill-

defined. From four to six spindles can usually be observed

along the abaxial surface of each anthocodia, but in no instance

do these protrude beyond the polyp-head. It is exceedingly

difficult to dissect out a single polyp with its supporting spicules

intact; some preparations were obtained, however, after soaking

portions of the branches for twelve hours or more in oil of

cloves. A more satisfactory method of observing the “ Stiitz-

biindel” is obtained by placing one of the terminal lobes,

previously cleared in oil of cloves, under a low-power binocular

microscope. By this means a stereoscopic view of the polyp and

its supporting spicules is obtained, such as is represented in

fig. 12. The cortex of the capitulum contains numerous hori-

zontally disposed spindles, some of which are among the largest

spicules found in this species; they resemble in type those

described from the polyps. In the outer wall of the stem are

found small spicules with broad rays (fig. 11, a) which are usually

numerous, but vary considerably in size and concentration.

These forms are covered with warts -015—036 mm. long, and

interlock with one another, thus accounting for the tough

consistency of the stem. Among the forms described above

there are also in the outer wall of the stem some larger spindles

with remarkable spines (fig. 11, 6); the latter, which have an

average height of ‘(044 mm. and basal breadth of -02 mm., are

often larger and sharper on one side of the spindle than on the

opposite side, and, since the spines all lie approximately in one

plane, give the spicule a comb-like appearance. The spicules of

the canal-walls (fig. 11, c, d) are not very plentiful, but are

distinguished by their stout appearance. They may be described

as very thick, longitudinally-striated spindles, somewhat flattened

and bearing low rounded warts. Their ends are either rounded

or bluntly pointed, the body of the spicule is straight or slightly

curved. Among these regular forms there are found a few

branched spicules. The latter resemble the regular forms in

structure, but are either triradiate or show short irregular

processes springing from the central region of a typical spindle.

The following are characteristic measurements of the spicules,

length by breadth, in millimetres :—

a) RE Oly pi (Outenh ai. s.caraiectasowraen ge eek ee 3 X °025

(aye Folia: (irien)) “5. ezccsse cy. cabot eke es coving IEC OD “112 X *023

‘07 X 02

Key StniZbiMmdel 5. ae. atest eee neers cose ee eT CO 6 X 05

“4 X °06

G@)) Contex (of capitalmy 0...) seeesels chee 1:0 X 075 ‘9 X 08

(e) Cortex of stem, radiate forms .................. 115 X ‘181 "058 X *054

Thickness of branches *03--015

Do. spindles (including spines) ... °85 & 125 "22 X °125

G7)) Canal-walls; spindles) :./s.J2.(u.b.. 27.2 eS Ug Heb 2 a) Seer

4 X °08

Do. triradiate forms ‘41 X °4, thickness of ray *1

(measured by taking two ter-

minal points asa base line and

the third point as the extremity). -22 x ‘18 thickness of ray ‘05

516 ; MR. E. W. SHANN ON

On applying the key to the species of Wephthya given by

Kikenthal (1903, p. 145) to the case of WV. bedfordi, it is seen

that this species falls within the group characterised by forms

in which the terminal lobes are conical but rounded. The

inner polyp-spicules are smaller than the outer; moreover,

they have the form of smooth rods. These characters taken

together indicate that WV. bedfordi resembles closely WV. pacifica

and WV. albida; and, since the spicules do not form a ring around

the base of the polyp, it may be inferred that the new species is

most nearly allied to WV. albida Holm, so far as the key to the

species can be relied upon. Excepting in the character of the

inner polyp-spicules, however, WV. bedfordi appears to be more

nearly related to WV. elongata Kiikenthal than to J. albida.

This supposition is supported by the fact that NV. elongata is

reported from Ternate, while WV. albida isa Red Sea form. The

points of difference and of resemblance between JV. bedfordi and its

two nearest allies may be seen at a glance in the appended table :—

Character, LV. elongata. NV. bedfordi. NV. albida.

aes Long ... 5 6-3 9

Terminal lobes Thick 3 4-3 4”

Polyp-bends thick | 98 “8 “

| Angle between eee p and

hl eee 45° Acute. Right.

ABEL ES ( Outer ... 27xX— 3X 025 *3X°03

Polyp-spicules § UInner ...) -13X— 112-023 “AX:015

(Spiny spindles.) | (Smooth rods.) | (Smooth rods.)

“Stiitzbiindel” ............] Projects slightly. | Does not project. | Projects slightly.

Spicules of “Sttitzbiindel ”

Long ...... 15 6 12

niches ase 0/5) "12

Spicules of Upper Cortex

TOME: scree 8 1 14.

Thick ... hi 075 "12

Spicules of Lower Cortex

Long ...... 8 "35-22 85

Thick ... (Compact.) 125 22

(Comb-like and

radiate forms.)

Spicules of Canal-walls:

Long ...... 12 11-4 1

Thick... (Scattered.) 2-08 2

(Spindle and tri-

radiate forms.) |

Colour (in alcohol) —...... Greyish yellow. Cinder-grey. | Greyish white.

JUCOEEUDUT fats eaeae eden on eceabao eRe Ternate, 5 fms. Singapore. Ted Sea.

All measurements are in millimetres.

Lest the use of an artificial key to the identification of species

should have led me to overlook a member of the genus not

included in the group to which the Singapore specimen apparently

ALCYONARIA FROM SINGAPORE. 517

belongs, but at the same time exhibiting a close resemblance to it

in structural detail, I have read descriptions of all the species of

Nephthya which might possibly have proved to be identical with

WV. bedfordi. In no instance is there a greater resemblance

between JV. bedfordi and another species of Vephthya than exists

between any two established species of that genus. Indeed, the

exceedingly feeble development of the ‘“Stiitzbiindel” and the

presence in the stem-cortex of exceptionally small, though

curiously comb-shaped, spicules are characters which serve clearly

to distinguish WV. bedfordi from any species previously recorded.

Among some unpublished notes by Miss Coward, which

Professor Hickson has kindly placed at my disposal, the following

paragraph occurs:

“In his work on the family Nephthyide (1903) Kiikenthal

names a specimen Wephthya chabrolii. In doing this he refers

to Hickson and Hiles’ (1900) description of WV. chabrolit. These

writers, however, state that the spicules of their specimen are

just as described by Klunzinger (1877)—that is to say, they

do not form a ‘ Stiitzbtindel.’ In the description of his specimen

Kiikenthal says the spicules only rarely project beyond the

polyp-heads—and yet his diagnosis is the presence of a

‘Stiitzbiindel.’”

This observation came to hand after the account of the new

species WV. bedfordi had been written and the ‘ Stiitzbiindel”

described. As stated above, the ‘ Stiitzbiindel” was very poorly

developed. I then thought it advisable to examine JV. chabrolii

to satisfy myself as to the nature of the “ Stiitzbiindel” in that

species. ‘The specimen at my disposal was the identical one

described by Hickson and Hiles (1900), in Willey’s collection.

Preparations, which have been cleared in oil of cloves and

examined under the binocular microscope, reveal the presence of

distinct bundles of spindles supporting the polyps. One or two

of these spicules not infrequently project beyond the polyp-

heads. One must admit that WV. chabrolit is characterised by

the presence of a small but clearly defined “ Stiitzbiindel.” It

seemed possible that the “ Stiitzbiindel” was so degenerate in

NV. bedford that the polyps might be described as being without

this characteristic. Were this the case, the specimen in question

would come under Kiikenthal’s definition of the genus Litho-

phytwm Forsk. I accordingly made and examined preparations

of Lithophytum arboreum ; but in this case there was no trace of

a bundle of spicules on the abaxial surface of the anthocodize

in the least degree comparable with the condition which I have

described in WV. bedford.

The genus Wephthya, as at the present time accepted, contains

a large series of species showing every gradation in the develop-

ment of a “Stiitzbiindel.” At one end of the series is found

such a form as JV. celosia Lesson, in which one or two of the

“ Stiitzbtindel ” spicules reach 2 mm. beyond the polyp-heads ;

in the middle WV. chabroli, in which they only project slightly ;

and at the other end WV. bedfordi, in which they do not project.

518 MR. E. W. SHANN ON

Genus DenpronerutTaya Kikenthal.

1791-97. Alcyoniwm Esper, Pflanzenthiere, pp. 49, 50, tab. 16.

1834. Nec Spongodes Lesson, Illustrations de Zoologie, vol. ii.

; part 2, p. 89.

1834. Nephthya (pars) Ehrenberg.

1846. Spoggodia (pars) Dana, p. 625.

1857. Spoggodes (pars) Milne-Edwards, p. 127.

1862. Spoggodes (pars), Morchellana Gray, in Proc. Zool. Soc.

London, p. 27.

1877. Spongodes (pars) Klunzinger, pp. 34, 35.

1889. Spongodes (pars) Wright & Studer.

1895. Spongodes (pars) Holm, p. 16.

1896. Spongodes (pars) Kiikenthal, p. 97.

1899. Spongodes (pars) May.

1905. Dendronephthya Kikenthal, Zool. Jahrb. xxi. p. 526.

The genus Spongodes was founded by Lesson (1834) on the

type Sp. celosia, and for many years workers in the field of the

Nephthyide found no difficulty im discriminating between

Spongodes Lesson and Wephthya Savigny. I have endeavoured

to show in my section on the genus Wephihya that as the number

of species of the above-mentioned genera increased it became

more and more difficult to draw a hard-and-fast line between

them; further, that this fact became manifest when, in 1889,

Wright and Studer described a specimen as Spongodes nephthyw-

formis, which was subsequently relegated to the genus Vephthya,

and, indeed, shown to be identical with J. chabrolit, the type of

that genus. A new distinctive feature was required: such a

feature was discovered by Holm and applied in a practical form

by Kiikenthal. The feature in question is the arrangement of

the polyps upon the stem, and its application has already been

mentioned above. Kiikenthal’s (1896) definition of the emended

genus Spongodes is as follows :—‘* Polypenstock baumartig veras-

telt, unterer Stammteil nackt. Die Polypen sind in Biindeln

vereinigt oder stehen vereinzelt.” Under the genus Spongodes

thus defined, Kiikenthal included as subgenera Spongodia and

Spongodes ; he further subdivided the latter into three groups,

namely, Glomerate, Umbellatz, and Divaricate. May (1899)

and others adopted this classification ; and so the matter stood

until the year 1905, when Kiikenthal published the second

part of his ‘‘ Revision of the Nephthyide.” In this paper was

introduced the division of the time-honoured genus Spongodes

Less. into the two new genera, Dendronephthya and Stereonephthya,

which has provoked such a storm of criticism. It must be.

remembered, however, that the group Spongodes ‘‘ Spicatee,” and

with it the original type Sp. celosia, had already been relegated

to the genus Vephthya; so that the genus Spongodes, as then

accepted, no longer retained its ancient prestige ; moreover, since

the genera Nephthya and Spongodes were shown to be synony-

mous, Kiikenthal was justified by the International Rules of

Zoological Nomenclature (Art. 28) in retaining the older name—

ALCYONARIA FROM SINGAPORE. 519

Nephthya—to designate the merged genera. The genus Dendro-

nephthya is almost synonymous with Spongodes (Glomerate,

Umbellate, and Divaricatze), as defined by Ktikenthal in 1896,

and Stereonephihya has a like relation with his Spongodes

(Spongodia).

There is no doubt that here, again, Kiikenthal was acting in

accordance with the International Rules of Nomenclature. Had

these Rules been extant at the time Kiikenthal published his

classification of the Nephthyide in 1896, the name Spongodes

would doubtless have been discarded by him then, for in the

second section of Art. 30 it is written: “In no case, however,

can the name of the original genus be transferred to a group

containing none of the species originally included in the genus.”

Thus Spongodes, as used by Kiikenthal in 1896, would not be

considered as a valid generic name by those who accept the

International Rules of Zoological Nomenclature, which appeared

in 1905. One cannot-but feel the loss of a name which for

nearly a century has been familiar to students of the Alcyonaria,

and regret the inconvenience caused in Museums and Zoological

Laboratories throughout the world by its suppression. Nomen-

clature is a matter of convenience and cannot be regulated

absolutely by arbitrary laws. Generally speaking, of two

synonyms, that which is most familiar to the majority is to be

preferred ; a more practical reason should be required than a

mere regulation before supplanting an old familiar generic name

by a new one. If we are required to obey literally the Inter-

national Rules of Zoological Nomenclature, such everyday names

as Astacus, Holothuria, Actinia, Madrepora, and many others

will be forfeited, as has been Spongodes. While regretting that

Kiukenthal did not exercise his authority in retaining the name

Spongodes, in deference to his extensive knowledge of the Neph-

thyidz and his able reorganisation of that family, I have adopted

the term Dendronephthya for a genus of Nephthyide defined as

follows by Kiikenthal (1905) :—

““Nephthyiden von baumformig verzweigtem Aufbau, deren

Polypen stets in Biindeln vereinigt sind, Polypen mit ‘Stiitz-

biindeln.’”

DENDRONEPHTHYA DIsciFoRMis Kiikenthal 1905. (Pl. LXIII.

fig. 13.)

Three specimens in the collection agree so nearly in all

essential features with the description given by Kiikenthal

(1905) for D. disciformis that there is little doubt that they

may be assigned to that species. Kiikenthal’s specimen came

from the China Sea.

The largest of the three colonies is 8 em. high, 7 em. broad, and

3°5 em. thick. It was taken off Pulo Brani in 5 fms. of water.

Two other specimens were taken in 7 fms. of water between

Pulo Hautu and Blakang Mati. Their measurements, height by

breadth by thickness, are 4x 4x 2°5 em., and 3°5x2°5x1°5 em.

respectively.

520 MR. E. W. SHANN ON

It may be noted that the two last-mentioned specimens are

much smaller than any recorded by Kikenthal. The colour

of the upper branches and “ Stiitzbindel” is pink, whereas the

typical colour for this region is deep red. Moreover, no radial

spicules were observed in the lower portion of the stem.

The exact systematic position of a fourth specimen is rather

doubtful; but, until further evidence is forthcoming, it has

been deemed advisable to describe the specimen under the specific

title D. disciformis.

This specimen shows externally the typical ‘ disciformis”

features. It is 9°5 cm. in height and 8:5 x.3:0 cm. in breadth,

thus exceeding somewhat in dimensions the largest specimen

described above. The characters and distribution of the spicules

are identical with those of typical examples, only excepting those

from the lower portion of the stem. The spicules found in the

last-mentioned area are spiny clubs (fig. 13), among which are

scattered a few of the typical radiate forms. The clubs vary in

length from ‘70-27 mm., but the majority are ‘45 mm.; the

thickness is almost the same even in forms of different length,

the average width of the handle of the club is ‘05 mm., that

of the head -12 mm., excluding the spines, which sometimes

reach -044 mm. in height. The colour of the branches and

“ Stiitzbiindel ” is a dull yellowish brown, and there are no red

spicules such as are found in these areas in all the other recorded

specimens. Kiikenthal (1905) described a colony whose branches

were pale brown and whose polyps were white, but even in this

specimen red spicules were observed.

Doubtless one might be led to base a new variety of Dendro-

nephthya on the characters of the colony described above, a

colony possessing a very characteristic form of spicule which has

not been observed, either from the same region or from any

other region, in any previously recorded specimen of D. diset-

formis. Such a course might have been adopted had not this

aberrant form been taken at the same time and from the same

place as the two smaller of the three colonies described above,

namely, between Pulo Hautu and Blakang Mati, 8/xii/98, in

7 fms. It seems probable, then, that the species D. disciformis

is liable to considerable variation in the colour of the terminal

branches and “ Stiitzbiindel,” and in the character of the spicules

which are found in the lower region of the stem. The latter

phenomenon is very remarkable, and, it is hoped, may act asa

check to the frequency with which new species are created to

separate two or more specimens of Alcyonaria which only differ

from one another in the shape of their spicules.

Genus STEREONEPHTHYA Kiikenthal.

1869. Spoggodia (pars) Gray, in Ann. & Mag. Nat. Hist. (4)

vol. 111. p. 128.

1877. Spoggodia (pars) Klunzinger.

1889. “ Divaricatee ” (Spongodes) Wright & Studer.

ALCYONARIA FROM SINGAPORE. 521

1895. Spongodia (pars) Holm, p. 25.

1896. Spongodia (pars) Kiikenthal.

1904. Spongedia (pars) Holm, Weiteres iiber Nephthya und

Spongodes.

1905. Stereonephthya Kiikenthal, p. 694.

“ Spongodes”-like forms in which the polyps were scattered

irregularly over the stem were recognised at an early date, and

were classified sometimes as a distinct genus, at others as a sub-

genus of Spongodes, under the name Spoggodia or Spongodia.

Wright and Studer (1889) recognised this group, but included

its members in the genus Spongodes under the designation

“ Divaricate.” Later writers, such as Holm (1895) and Kiiken-

thal (1896), agreed that Spongodes and Spongodia were synony-

mous. Hence Spongodes dragged Spongodia with it in its

downfall in 1905. For similar reasons to those stated in the

section on Dendronephthya I have adopted the new genus Stereo-

nephthya which Kiikenthal (1905) defined thus: ‘Sehr starre

Nephthyiden, deren Polypen weder in Liippchen noch in Biindeln

angeordnet sind, sondern einzeln oder in kleinen Gruppen direkt

vom Stamm wie den nicht oder wenig veraweigten Hauptiisten

entspringen. Polypen mit ‘ Stiitzbiindeln.’”

STEREONEPHTHYA LUTEA, sp. n. (PI. LXIII, fig, 14.)

The specimen, on the observation of which this new species is

based, agrees in every respect with the diagnosis given by

Kiikenthal (1905) for the genus Stereonephthya, At the same

time it possesses characters which serve clearly to distinguish it

from other species hitherto described.

The colony is upright and branched. Numerous tapering

branches leave the insignificant stalk either singly or in groups,

and spread in all directions, but with a distinct tendency to

assume an upright position. When they arise in groups, the

members forming a group are frequently united for a considerable

distance by a common cortex. The primaries sometimes, but not

invariably, bear a few stationary branches, which also show a

tendency to grow in a vertical direction. From both the primary

and secondary branches short lateral lobes are given off at

irregular intervals.

Measurements :—

Eleiohitial colowy iit .ath Thao PEG Be Rude gon 7

Brest buitat, Waser ii). Jens. tele, SANs aee ney Dyk i

Maximum breadth of capitulum ..............0......... 5

hensth’ ofprimary branchess yi. 55.. Mids dos tte wn ve 3°5-5

; 1

Breadth of primary branches | sit xGheetellagl eco ss) 0:

Length by breadth of secondary branches ............ 15x

Length by breadth of lateral lobes .................0... , 05x 0°

Proc. Zoou. Soc.—1912, No. XXXV. 3D

ey MR. E, W. SHANN ON

The polyps arise sporadically from all parts of the colony.

They are closely aggregated on the lateral lobes, and are most

scarce on the basal portion of the primary lobes. The anthocodize

are supported by a “Stiitzbiindel ” (fig. 14), three horny spindles

of which project -2--4 mm. beyond the polyp-head. The polyp-

heads make an acute angle with their stalks ; the stalk is 1:2 mm.

high, the head is 1 mm. in length and -7—85 mm. in maximum

breadth. The polyps contain thorny spindles of various sizes

and a few clubs. These are sometimes placed horizontally, but

are more usually oblique, and in the distal area are arranged

en chevron. These spindles measure 45-035, -27x:04, °2x

025 mm. Among the above and extending into the tentacles

are numerous little spicules which are nearly rectangular in

outline; they measure 110-036, :095 «:015, 050 x-015 mm,

The “ Stiitzbiindel” is composed of a number of thorny spindles

of a bright yellow colour (they resemble at first sight crystals of

sulphur). These spicules are ten in number round the base of |

the polyp-stalk, but there are only three at the distal end, The

four spicules which underlie the point of origin of the polyp-

head are semi-lunar in shape, the concave sides facing the

polyp-head, The “ Stiitzbiindel” spicules reach 1-8 mm. in length

and ‘12 mm.in breadth. The cortex of the colony derives its

characteristic colour from the enormous number of slim yellow

spicules which it harbours. There are also thorny spindles ; they

lie parallel to and very near the surface, and vary very much in

size and shape. Some are twice, others only half, the size of the

“‘Stiitabiindel” spicules. Few are straight, most are semi-lunar

or S-shaped, and very exceptionally a triradiate form occurs.

The ends of the spindles are usually bluntly pointed, but

occasionally forms were observed with their ends frayed out into

a number of points. The largest measure 3°0 x-28, 2-7 x <30,

2-1 x °15, the smallest °67 x ‘07, °67 x :04, 55 x 05 mm., but every

conceivable intermediate stage can be found, A remarkable

feature of this species is the entire absence of spicules from the

canal-walls.

The colour of the colony in alcohol 3 is pale yellow, the polyps

are cream-coloured.

The specimen was taken in Imbiab Bay, where it was left

exposed at the lowest tides.

Two other specimens taken in the littoral region at Teluk Ayer

have also been assigned to this species. At first sight they do

not resemble closely the type specimen, for they are shorter,

broader in proportion, and more bushy in appearance than the

colony described above. One of these colonies is 4°3 cm. in

height, its base is ‘8 cm., and the maximum diameter of the

capitulum is 3:1 cm,; the measurements of the other, taken in

the above order, are 3x‘7 cm.: 455 cm. Theapparent difference

in the branching from that of the type is due to the fact that

the secondary branches are no larger than the lateral lobes, and

that both these kinds of offshoot are relatively numerous. The

polyps are arranged much more densely than in the type,

ALCYONARIA FROM SINGAPORE. 523

especially at the tips of the branches. The colour of these

specimens, moreover, is of a duller yellow. These slight differences

appear quite insignificant when one considers that these two

specimens resemble the type of Stereonephthya lutea in the

following important characters :—

The size and shape of the anthocodie. The size, distribution,

and remarkable sulphur-yellow colour of the spicules; the dis-

tribution of spicules agrees even to the disposition of those

forming the “ Stiitzbiindel” ; the large internal canals, with thin

walls devoid of spicules.

Genus ParasponcopEs Kiikenthal.

P. crassa Kiikenthal, 1896.

This species was first described by Kiikenthal in 1896, and

further reference to it igs made by the same author in his

‘Revision der Aleyonarien’ (1907), where it is described under

‘“ Species incerti generis.” Two small colonies of a pale brown

colour are placed without hesitation in this species.

One colony was taken in 5-9 fms, of water from the New

Harbour. It measures 20 mm. in hejght; the diameter of the

capitulum is 13 mm., thag of the stem 7°5 mm.

The other colony was taken in 5 fms. of water, the precise

locality is not recorded. Jt measures 20 mm, in height; the

diameter of the capitulum is 11 mm., and the stem is broken.

The only important character in which these specimens differ

from the type is the size of the gpicules in the stem-cortex.

Measurements, length by breadth in millimetres, for three of the

largest spicules from this region are 1:2 x18, 1:0 «+15, :95 « °22.

Kiikenthal found spicules measuring as much as 1:8 x °24 mm. in

the stem-cortex. The spicules from the Singapore specimens

show, however, the typical warts (702 mm, high).

Previously recorded from ernate, at a depth of 30 fms,

‘

Family SIPHONOGORGIID4.

Genus SIPHONOGORGIA.

S. VARIABILIS Hickson (olim Chironephthya varjabilis Hickson). _

Two beautiful little specimens taken in ]0 fms, of water south

of Blakang Mati are referred to this very variable species, Each

possesses a very slender stem, from the apex of which two main

branches are given off, so that the appearance of the colgny is

roughly Y-shaped. These primary branches give rise tg secondary

branches, which in one or two cases are again divided. The

polyps occur most frequently on the tips of the branches, but a

few are scattered on all portions of the colony except the stem.

Measurements :—

Specimen A. Specimen B.

mm, mm,

ilersinpot eben? ii) yeas ce 18 11

Diameter of stem... 22.6.0. 48 she. 6 3

Length of primary branches ...... 23 and 23 24 and 18

3o*

5o4 MR. E. W. SHANN ON

The spicules are of the usual shape and of medium size, and

assume the colour of the part of the colony in which they are

embedded. The large warty spindles of the cenenchym are

tinted with mauve and do not exceed 3:0 x°3 mm. In the polyp-

area the spindles are more frequent than the clubs ; all the polyp-

spicules are more or less bent, they are either bright red or bright

yellow in colour, and the larger ones measure *6x°‘05 mm.,

‘47x °065 mm. (the red spicules mainly compose the ‘ crown,”

the yellow ones the “ points ”),

The colour of the colony in alcohol is a delicate pinkish mauve.

The polyps are deep red, and the tentacles bright yellow. It is

difficult to conceive how the colouring could have been more

vivid or the blended tints more pleasing even before these little

colonies were removed from their natural habitat.

For a list of the colour-variations to which this species is liable

to run, see Hickson’s ‘“‘ Aleyonaria of the Maldives,” Part I.

p. 488 (under Chironephthya variabilis), and Thomson and

Simpson (1909, p. 125).

Previously recorded from Mahlos Atoll, S. Nilandu, Persian

Gulf, Andamans, and the Arakan Coast.

Order GORGONACEA.

Suborder PSHUDA XONTA.

Family Sc LEROGORGIID&.

Genus SUBEROGORGIA.

S. SUBEROSA Pallas.

Gorgonia suberosa Pallas, Elench. Zooph. p. 191.

Suberogorgia suberosa Gray, Proc. Zool. Soc. 1857, p. 159.

Sclerogorgia suberosa Kolliker, Icon. Histiol. p. 142, pl. xix.

fig. 13 (2).

Sclerogorgia suberosa Studer, Monatsber. K. Akad. Wiss.

Berlin, 1878, p. 666.

Suberogorgia subercsa Ridley, Journal Linn. Soc. Zoology,

vol, xxi. p. 243.

Suberogorgia suberosa Wright & Studer, ‘Challenger’ Re-

ports, vol. xxxi. p. 166,

Suberogorgia suberosa Brundin, ‘ Aleyonarien aus dem Samm-

lung der Zool. Mus. in Upsala,” Svenska Vet.-Akad. Hand.

Xxll. pt. iv.

There is in the collection a bottle which contains a number of

fragments, apparently the component parts of a single colony of

S. suberosa. The basal portion is present and has a diameter

of 6 mm.; the diameter of the terminal branches is 2°5 mm.

The specimen was taken in about 14 fms. of water off Pasir

Panjang,

ALCYONARIA FROM SINGAPORE. 525

Previously recorded from the coasts of Western Africa and the

West Indies (Pallas and Esper); the Mermaid Straits, Dampier

Archipelago, and N.W. Coast of Australia (Studer); Port

Denison, Queensland, and Torres Straits (‘ Alert’ Coll.); Mauritius

(Coll. Brit. Mus.); Admiralty Islands (‘Challenger ’ Coll.); and

Sumatra (Brundin).

Family MELITopDID«,

Genus Metiropgs.

M. Avpirincta Ridley.

This species is represented by numerous fragments, but none

of these represents an entire colony. The specimens were taken

in 15 fms. of water from Blakang Mati.

Previously recorded from Port Molle, Queensland.

Genus PsILACABARIA.

P. GRAciuiima Ridley.

Psilacabaria gracillima, n. gen. et sp., Ridley, Rep. Zool. Coll.

H.M.S. ‘ Alert,’ Aleyonaria, p. 363.

After a lapse of nearly three decades this delicate little species

is to be recorded once more. It is represented in the collection

by four portions of colonies, which show the mode of branching

and length of the internodes, and by sundry fragments. None

of the colonies are complete, but the largest intact portion

measures 45 mm. in height. The internodes vary in length, the

majority, however, fall within the range given by Ridley, viz.

12-16 mm. The branches are much more delicate than those of

the type, measuring only 1-2 mm. in diameter, as compared with

3-7 mm. In the mode of branching, size of polyps, and details

of spicule-distribution the Singapore specimens resemble the

type very closely. The spicules show the typical shapes; but

they are in each instance somewhat smaller than their prototypes.

The colour of the ccenosare and polyps is fawn, that of the axis

white.

Locality. Salat Sinki, in 4-5 fms.

Previously recorded from Port Molle, Queensland, 12-20 fms. ;

Port Darwin, 8-12 fms.; E. Australia, 42 fms. (Ridley).

Genus WRIGHTELLA.

W. Ropusta, sp.n. (Pl. LXII. fig. 9; Pl. LXIII. fig. 15.)

This species has been formed to include a single well-developed

colony, which agrees more closely with Wrightella Gray (1870)

than with any of the other genera of the family Melitodide.

The form of the colony differs, in its stouter dimensions and in

its erect position, from the members of the four existing species

of Wrightella ; but it was thought expedient to associate it with

526 MR. E. W. SHANN ON

these, rather than to create a new genus from the observation of

a single specimen.

Wright and Studer (1889, p. xxxvi) give the following de-

finition of the genus Wrightella :—“‘ The branches and twigs are

compressed ; the projecting polyp calyces occur especially on the

sides. In the cortex there are foliaceous clubs. There are no

nutritive canals in the axis.” The specimen fulfils all these

conditions.

The single main stem arises abruptly from a strong reticulate

base (fig. 9). The base, which is broken at the edges, measures

15 mm. in diameter, and the gaps in its meshes average 1 mm.

in diameter. The colony is 120 mm. in height and 45 mm. in

breadth (some of the lateral branches are broken, so it is probable

that the true breadth exceeded 45 mm.). The branching of the

colony is dichotomous, and takes place at the swollen nodes. The

nodes are less prominent in the distal branches. All the branching

takes place in one plane; the terminal twigs are markedly flattened

in the plane of branching. Anastomosis of the upper branches

takes place at infrequent intervals. The nodes near the base are

globular, having a diameter of 4 mm.; the internodes in this

region are circular in section, their diameter is 2°5 mm., which is

slightly exceeded by their length. The internodes beyond the

lowest three become more elongated and show an average length

of 10 mm. The terminal twigs are only 1 mm. wide. The

varruce, which measure *75 x°75 mm., are not densely crowded,

and show a tendency to arrange themselves on the Jateral aspects

even of many of the lower branches, but more especially in the

terminal twigs. Both cortex and verrucz are yellow in colour.

The axis is white, and is not traversed by nutrient canals.

The precise locality is not. recorded, but, like the other speci-

mens in this collection, it was taken in shallow water near

Singapore.

The spicules attain all manntr of shapes (fig. 15); they

are quite colourless: The foliaceous clubs characteristic of the

genus Wrightella are present in large humbers, and show the

following range in measurement : length by breadth -27 x -12 mm.,

°15 x -06 mm. ; handlés of clubs ‘02 mm.if diameter. Numerous

spindles occur, some are foliaceous and measure *27 X-07 mm.,

-20 x ‘08 mm., 12 x-06 mm.; others are spiny, the spines fre-

quently bemg confined to the central region, and measure

22x05 mm:, ‘17-03 mm:, -12x‘03 mm. A few stellate

forms are found which have a diameter of ‘10-15 mm. Minute

scales abound; they measure °05 ‘025, °036x:030 mm.,

028 X 014 min,

Literature:

1870, Gay, J. E.—Cat. Lithophytes Brit. Mus. p. 31.

1900. Hickson, 8. J:, atid Hires, Isa L.—‘The Stolonifera and

Aleyonacea collected by Dr. Willey in New Britain,

ete.” Willey’s Zool. Results, pt: iv; pp: 493-508.

1877.

1903.

1905.

1907.

1903.

1906.

1909.

1889.

ive} [e'3) “Io

ALCYONARIA FROM SINGAPORE, BAT

Kuiunzincrer, C. B.—Die Korallthiere des Rothen Meeres.

Part I. p. 33, tab. 2. fig. 5:

Koxentaat, W.—“ Versuch einer Revision der Aleyo-

narien.” IT, Die Familie der Nephthyiden: 1 Theil.

Zool. Jahrb. Syst. xix.

Do. do. 2. Theil, .Op, cit. xxd.

Do. do. 3. Theil. Op. cit. xxiv.

Pratt, Eprra M.—‘The Aleyonaria of the Maldives.

Part IIL. Sarcophytuwm, Lobophytum, Sclerophytwm, and

Alcyoniwm.” Fauna and Geog. of the Maldive and

Laccadive Archipelagoes. (J. Stanley Gardiner), Vol. ii.

Part I.

Tuomson, J. ARTHUR, and HEenpERson, W. D,—“ Aleyo-

naria of Zanzibar and British East Africa.” Proc,

Zool. Soc. pp. 393-443,

Tuomson, J. ARTHUR, and Simpson, J. J.—“ An Account of

the Alcyonarians collected by the Royal Indian Marine

Survey Ship ‘Investigator’ in the Indian Ocean.”

Part I.

Wricut, EK. P., and Sruper, Tu.— Report on the

Scientific Results of the Voyage of H.M.S. ‘Challenger.’ ”

Vol. xxxi. Aleyonaria. ;

EXPLANATION OF THE PLATES.

Puate LXI.

- Ammothea virescens Sav. (= Lithophytum arboreum Forsk.), entire colon ys

Savigny, Tab. 2. tig. 1.

. Nephthya chabrolii Audouin, terminal lobe magnified; Savigny, Tab. 2.

TS Ban

. WN. chabrolit, single polyp magnified, lateral aspect; Savigny, Tab. 2.

fig. 5. 3.

. Same, abaxial axis; Savigny, Tab. 2. fig. 5. 4.

. Same, more highly magnitied ; Savigny, Tab. 2. fig. 5. 5.

Priate LXII.

. NV. chabroltt, entire colony ; Savigny, Tab. 2. fig. 5. 1.

- Sclerophytum pinnulatum, sp. n., entire colony; from a plictograph,

slightly reduced.

. Nephthya bedfordi, sp. n., entire colony; from a photograph, natural

S1Ze.

. Wrightella robusta, sp. n., entire colony; from a photograph, X 2.

Pirate LXIII.

. Sclerophytum pinnulatum, sp. n., tentacle bearing pinnules, X 245.

. Nephthya bedfordi, sp. n., spicules, < 72.

. WV. bedfordi, terminal lobe bearing polyps, x 25.

- Dendronephthya disciformis Kiikth. (aberrant specimen), spicules from

the stem-cortex, X 72.

. Stereonephthya lutea, sp. n:, polyp and “ Sttitzbundel,” x 72.

. Wrightella robusta, sp. n., stem-spicules, x 72.

* Figures marked thus are reproduced from photographs procured by Professor

Bourne,

and kindly given by him to mé for the purpose.

528 MR. T. H. WITHERS ON

27. Some early Fossil Cirripedes of the Genus Scalpellum.

By Tuomas H. Wrrumrrs, F.G.S.*

[Received January 31, 1912: Read March 19, 1912.]

(Text-figs. 64 & 65.)

INDEX.

Page

Structure of carina of Sealpellwm ..........c. cee 528

Evolution of early fossil forms of Senna sent 530

Distribution (Geological) of Cretaceous Scalpellum 537

Scalpellum arcuatum Darwin ............000cceeeeeeeees 532

S. trilineatum Darwin .......... paetiseapes(pgOO

Scalpellum spp.—discussion of aimitios, Serenade Gizis)

Among the Cirripede remains from the Albian (Gault) of Folke-

stone in the collection of the British Museum (Natural History),

three detached valves were noticed, which, for certain reasons to

be explained later (see p. 532), appear to be the original valves

upon which Darwin founded the species Scalpellum arcuatum t.

A portion of one of the valves, a carina t, happened to be broken

away from its matrix, showing that the intraparietes, said by

Darwin to be absent in this species, were really present. These

parts, however, are developed in such a way that they can be

seen only by cleaning the valve free from matrix. The finding

of the intraparietes in Darwin’s type of S. arcwatwm led to the

examination of further examples of the carina, and the fact was

established that intraparietes are developed in all carinz of this

species.

S. trilineatum Darwin (1851, p. 38, pl. i. fig. 5), from the

Cenomanian (Grey Chalk) of Dover, was said by Darwin to come

nearest to S. arcuatwm, owing to the absence of intraparietes.

The carina, which is the holotype of S. trilineatwm, is in the

British Museum (Natural History), registered 38461. On freeing

this specimen from the gum and matrix which obscured the inner

portion of the valve, it was at once apparent that not only were

the intraparietes present, but that they were developed almost

exactly as in S. arcuatwm.,

Other Lower Cretaceous species of Scalpellwm in which the

intraparietes of the carina are said to be absent are S. simplex

Darwin (1851, p. 39, pl. i. fig. 9) and S. accumulatum Withers §,

both of which come from the Aptian (Lower Greensand). The

unique carina of 8. simplex cannot at present be traced, but since

the parietes do not reach to the basal margin of the valve (see

text-fig. 64, 7) as in those carinee which have intraparietes, it

does not appear probable that intraparietes could be present. In

* Communicated by Dr. W. T. Caiman, F.Z.S.

+ C. R. Darwin, 1851, Pal. Soc. Monogr. Foss. Lepadide, p. 40, pl. i. fig. 7.

+ For the namies of the various valves see text-fig. 65.

§ T. H. Withers, 1910, Geol. Mag. dec. v. vol. vii. p: 152; figs. 1-4.

FOSSIL CIRRIPEDES, 529

S. arcwatum (text-fig. 64, 3) and 8. trilineatum (text-fig. 64, 4)

the parietes reach to the basal margin, and in my opinion the

intraparietes will be found to be developed in S. accumulatum

(text-fig. 64, 2) as in those species, since S. accumulatum is

similar in its general external form. The only carina known

Text-fig. 64.

Early types of the Carina of Scalpellum.

la-5a, side view; 3b, 4b, 5 6, inner view ; 1c-5 ¢c, transverse section near apex.

t., tectuin ; p., parietes ; z.p., intraparietes.

of S. accwmulatwm is embedded in such a hard matrix that it is

impossible to expose its inner surface without danger to the speci-

men, so that the matter must here remain until further specimens

are forthcoming.

So far, then, we have proved the existence of intraparietes in

530 MR. T. H. WITHERS ON

the carine of the two species S. arcuatum and S. trilineatwm, and

pointed out the possibility of their presence in S. accwmulatum.

The intraparietes in these species, instead of forming a thin wall

on each side of the carina as in the form represented by text-

fig. 64, 5, are bent inwards almost at right angles, and the upper

regions of their inner margins meet to a greater or less extent

(see text-fig. 64, 36, 46); the upper part of the valve is solid, and

must have projected freely to the extent indicated by the meeting

of the intraparietes*. The peculiar development of the intra-

pavietes in the carine of these species is therefore of importance

as showing a development of the carina distinct from that in

which the intraparietes form a thin wall on each side of the

carina.

Three types of carina, all having an apical umbo, were there-

fore already developed amongst these early forms of Sealpellwm,

and the geologically oldest of these more closely resemble the

carina of Pollicipes, from which Scalpellwm is considered to be

derived. These are (1) represented by S. simplex (text-fig. 64, Z)

from the Aptian (Lower Greensand), which has no intraparietes,

the tectum being flatly-arched transversely, the parietes bent

almost at right angles to the tectum and not extending to the

basal margin ; this type is distinguished from the Pollicipes type

of carina only in the parietes being separated from the tectum

by a distinct angle; (2) represented by S. accwmulatum (text-

fig. 64, 2) (Aptian, Lower Greensand), S. arcuatwm (text-

fig. 64, 3) (Albian, Gault), and S. trilineatwm (text-fig. 64, +)

(Cenomanian, Grey Chalk), which have intraparietes, these parts

being bent inwards almost at right angles and joining, the upper

part of the valve being solid and projecting freely to a consider-

able extent ; in the solidity and free projection of the upper part

of the valve this type is allied to Pollicipes ; (3) represented by

S. hastatuwm (text-fig. 64, 6) and other species from the Ceno-

manian which have intraparietes also, but are characterized by

these parts forming a thin wall on each side of the carina, the

apex of which projects freely, slightly, or not at all; this latter

type is more typical of the genus Scalpellwm, and, owing to the

upward growth of the intraparietes in some forms, subsequently

gave rise to the species with an angularly bent carina having the

umbo in a subcentral position, a type which is not known below the

Upper Senonian. The only species with an angularly bent carina

known from the English Chalk is S. darwinianwm Bosquet T, but

this has an early specialized form of carina in which the upward

* A somewhat similar development of the intraparietes can be seen in the carinie

of S. maximum var. cylindraceum Darwin (1851, p. 33, pl. ii. fig. 2) from the Upper

Senonian (Belemnitella mucronata-zone) of Norwich, Norfolk, and S. solidulwm

Steenstrup (1839, Kroyer, Naturhist. Tidsskrift, Bd. ii. p. 412, pl. v. figs. 14, 14* ;

Darwin, 1851, Pal. Soc. Monogr. Foss. Lepadide, p. 42, pl. i. fig. 8) from the Upper

Senonian of Kjuge, Scania.

+ J. Bosquet, 1854, Les Crust. Foss. du Terrain Crétacé du Duché de Limbourg,

p. 46, pl. iv. figs. 6-12; T. H. Withers, Jan. 1911, Geol. Mag. dec. v. vol. vill.

p. 28, figs. 3-4.

FOSSIL CIRRIPEDES. 531

extension of the valve is due, not merely to the upward growth of

the intraparietes, but to the almost equal upward and downward

growth of the valve from the umbo, the whole external surface of

the valve being ornamented. It is, therefore, probable that

species with an angularly bent carina, due to the upward growth

of the intraparietes, which is apparently the more primitive type,

existed even earlier than in the Upper Senonian *.

Extreme interest attaches to the remarkably complete example

of S. arcuatwm from the Albian (Gault) of Folkestone, Kent, here

described and figured (p. 534, text-fig. 65, 6). To find so many

valves of the capitulum in position is a remarkable circumstance,

very few specimens of Scalpellum having been discovered with

several valves associated. Those known up to the present come

from the Upper Senonian, and, with the exception of a single

incomplete capitulum of S. maximum, belong to the species S. fos-

sula. This specimen, therefore, represents the only fossil species

of Scalpellum obtained from below the Upper Senonian with any

considerable number of valves in position. Previously described

species of Scalpellum from rocks below the Upper Senonian have

all been founded on detached valves, and in most cases on single

valves. Neither the upper latus, rostral latus, or rostrum, are

included in these descriptions, but are represented in the present

specimen. A further example of S. arcuwatwm (described, p. 533)

has furnished some scales of the peduncle, which so far are not

known in any species found below the Upper Senonian. SS. arcu-

atum is the only representative of Scalpellum known from the

Albian (Gault) of England.

Consequent on the discovery of the intraparietes in the carine

of S. arcuatum and S. trilineatwm, and since the carina is the

typical valve of the genus, it is here proposed to give fresh

diagnoses of those species and to describe the new material of

S. arcuatum. From the two examples of S. arcuatum it is

possible to construct a restoration of the capitulum, to which only

two valves are diagrammatically added (see text-fig. 65, 7, p. 534).

ScALPELLUM TRILINEATUM Darwin,

1851. Scalpellum trilineatum C. R. Darwin, Pal, Soc. Monogr.

Foss. Lepadide, p. 38, pl. i. fig. 5.

1854. Scalpellum trilineatum C. R. Darwin, Ray Soc. Monogr.

Sub-class Cirripedia, Balanide, Synopsis et Index

Systematicus, p, 633.

1854, Scalpellum trilineatum J. Morris, Cat. British Fossils,

2nd ed., p. 97.

1877. Scalpellum trilineatum H. Woodward, Brit. Mus. Cat.

Brit. Foss. Crustacea, p, 143.

* Since the above was written, two exceedingly small, angulatly bent carine

(length respectively 1°7 mm. and 1°8 mm.) with the umbo subcentral, due ‘to the

upward growth of the intraparietes, have been obtained by Mr. F. Mockler from the

Cenomanian (Chalk Marl) near Carabridge. The specimens are incomplete and

poorly characterized, but of much importance as showing that species with an

augularly bent carina existed even so far back in time as the Cenomanian.

532 MR. T. H. WITHERS ON

This species was founded on a carina and tergum from the

Cenomanian (Grey Chalk) of Dover. The tergum cannot now be

found, but the carina, which Darwin considered to be the typical

valve of the genus and can therefore be regarded as the holotype,

is in the British Museum (Natural History) registered 38461.

Diagnosis—Carina with three prominent, rounded, longi-

tudinal ridges on its tectum—one central, and one on each side

separating the tectum from the parietes; intraparietes bent

inwards almost at right angles, the inner margins meeting a

short distance below the apex, upper part of valve solid and

projecting freely.

Description of Carina.—Carina narrow, widening very gradu-

ally from the apex, considerably bowed inwards, basal margin

obtusely angular. Tectum flatly-arched transversely, with a

central, prominent, rounded ridge extending from the apex to

the basal margin, and bounded on each side by a slightly coarser

but flatter ridge on the angle separating the tectum from the

parietes. Parietes narrow, less than half the width of the tectum,

bent almost at right angles to the tectum, slightly concave.

Intraparietes very narrow, bent inwards almost at right angles,

the inner margins meeting about one-sixth the length of the

valve from the apex; the upper part of the valve is solid, and

must have projected freely to the extent indicated by the meeting

of the intraparietes. Lines of growth plainly marked.

SCALPELLUM ARCUATUM Darwin. (Text-fig. 65.)

1851. Scalpellum arcuatum C. R. Darwin, Pal. Soc. Monogr.

Foss. Lepadide, p. 40, pl. i. fig. 7.

1854. Scalpellum arcuatum C. R. Darwin, Ray Soc. Monogr.

Sub-class Cirripedia, Balanide, Synopsis et Index

Systematicus, p. 633.

1854. Scalpellum arcuatum J. Morris, Cat. British Fossils, 2nd ed.

poo.

1865. Scalpellum arcuatum J. W. Salter & H. Woodward

Cat. & Chart Foss. Crustacea, p. 27, pl. i. fig. 14.

1877. Scalpellum arcuatum H. Woodward, Brit. Mus. Cat. Brit.

Foss. Crustacea, p. 142.

The species S. arcuatwm was founded on three detached valves,

namely, carina, scutum, and tergum, from the Gault of Folke-

stone, Kent. These valves were considered by Darwin to belong

to the same species, and the material now to be described proves

such to be the case. Darwin further stated that these valves

were in the Bowerbank Collection. This collection was acquired

in 1865 by the British Museum (Natural History), and among

the specimens are three valves, a carina (I. 13796), a scutum

(I. 13797), and tergum (I. 13798), mounted together on Bower-

bank’s original tablet, and labelled in Darwin’s handwriting

“ §. arcuatum.”’ The carina has been much broken, presumably

since Darwin described it, but the scutum and tergum are in good

FOSSIL CIRRIPEDES. 533

condition. These specimens are also exactly half the size of

Darwin’s figures, which are enlarged to two diameters, and in

these circumstances there seems little doubt that they are the

original valves figured by Darwin. The carina, since Darwin

considered it to be the typical valve of Scalpellwm, is consequently

regarded as the holotype.

Two specimens in the British Museum (Natural History) add

considerably to our knowledge of this species.

One, registered (I. 13577), text-fig. 65,6, first appeared to

consist of a carina with fragments of the scuta and terga. Careful

clearing away of the matrix and the abundance of gum which

covered the specimen showed, however, that several of the valves

of the capitulum were preserved. These consist of the carina, the

paired scuta, the paired terga, an upper latus, a rostral latus, and

the rostrum. The left side of the capitulum is uppermost, and the

carina is so exposed that the intraparietes can readily be seen.

Portions of the inner surfaces of the right scutum and tergum are

also exposed owing to the fact that the upper valves are somewhat

displaced and broken.

The second specimen (I. 13580) has not so many valves pre-

served and those present are somewhat broken. The right side is

uppermost, and the plates shown are the carina, the pair of

terga, the upper portion of the left scutum showing its inner

surface, and the left upper latus. This last valve showed only its

inner surface, but on removal from the matrix it was found to

be the left upper latus. This specimen is of interest since it

furnished eighteen scales of the peduncle. A supposed shell-

fragment was removed from between the two terga, and on being

cleaned was found to be a peduncle scale. On removing and

washing the remaining matrix from the same position, the

number of scales was increased to eighteen. There is little

doubt that these scales belong to S. arewatwm, since it is the only

species of Scalpellum known to occur in the Gault. Moreover,

they are ridged like the valves of the capitulum of that species,

and the fact that they were found in such close association is

prima facie evidence that they belonged to the same individual, and

were washed into the position in which they were found. They are

somewhat similar in shape to the scales of the peduncle of the

Upper Senonian species S. maximum and S. fossula as figured by

Th. Marsson *, but are easily distinguishable from them by their

longitudinal ridging.

Diagnosis.—Capitulum composed probably of fourteen valves f,

which are ornamented with numerous fine ridges radiating

from their apices; umbo of all valves apical. Carina with

tectum flatly arched transversely, parietes rectangularly inflected,

* Th. Marsson, 1880, “ Die Cirripedien und Ostracoden der weissen Schreibkreide

der Insel Riigen,” Mitth. naturwiss. Vereine Neu-Vorpommern und Riigen, xii,

pl. i. figs. 26, ¢, d, 3d, e.

+ It is possible that this species had a subcarina, in which case the number of

valves would be fifteen. A higher number of valves is not likely.

534

MR. T. H. WITHERS ON

Text-fig. 65.

ies

CER,

ga. bis gh

Fig. 6. Scalpellum apcuatum Darwin, Nearly complete capitulum showing the

Fig. 6 a.

Fig. 65.

Fig. 6c.

Fig. 6 d.

Fig. 6 e.

left side uppermost, with the valves somewhat displaced and broken. X 2

diam, Albian, Gault: Folkestone, Kent. Brit. Mus, (Nat. Hist.), I, 13577.

c., carina, showing the inflected intraparietes; ¢.1, outer portion cf left

tergum ; ¢.2, inner view of right tergum near apex, showing the ridges

evidently connected with the firm attachment of the cortum; s,1, outer

portion of left scutum ; s.2, inner view of right scutum showing the pit

for the addyetor scutorum; J., upper latus; 7./,, rostral latus with ypper

portion broken off; 7., rostrym.

Id. Outer view of incomplete rostral latus,

Id. Inner view of same. ‘

Id. Outer view of rostrum. * 4 diam,

Id. Hypothetical transverse section of same.

Id, Side view of same.

Fig. 7. Restored capitulum of Sealpellum ayreuatum Darwin. ‘This figure is based

on the nearly complete capitulum represented by fig. 6, with the addition

of a carinal latus and an infra-median latus. These two yalves have not yet

been discovered in the Gault. XX 2 diam, Albian, Gault; Folkestone,

Kent. ./., carinal latus ; 7./., infra-median latus,

Fig, 8, Scalpellum arcuatum Darwin. ‘Two scales of the peduncle, a, outer yiew,

b, inner view; c, outer view; d, inner view. X 8 diam, Albian, Gault;

Folkestone, Kent, Brit, Mus. (Nat. Hist,), I, 138980,

FOSSIL CIRRIPEDES. 535

intraparietes bent inwards and meeting for about one-fourth the

length of the valve from the apex, upper part of valve solid and

freely projecting. Scutum with tergo-lateral angle almost in line

with the middle of the valve, Tergum subrhomboidal, a delicate

furrow extending from the apex to the basal angle. Upper latus

subtriangular, apex acutely angular, slightly bowed towards scuta,

basal margin rounded. Rostral latus acutely angular transversely,

about 23 times as long as wide. Rostrum subtriangular, with a

strong median keel extending from the apex, widening towards

the convex basal margin.

Description of valves.—Carina narrow, widening gradually from

the apex, considerably bowed inwards, basal margin obtusely

V-shaped. Tectum flatly-arched transversely, obscurely carjnate,

and ornamented with numerous fine longitudinal ridges, Parietes

narrow, less than half the width of tectum, not longitudinally

ridged, bent almost at right angles to the tectum, slightly concave,

Intraparietes very narrow, bent inwards almost at right angles,

the inner margins meeting about one-fourth the length of the

valve from the apex, above which the valve is solid and must

have projected freely,

Scutum moderately convex, divided unequally by a prominent;

ridge running from the apex to the basi-lateral angle, the basi-

lateral angle being slightly produced, Apex acuminate. Basal

margin sinuous, about two-thirds the length of the valve;

eccludent margin slightly convex and nearly parallel to the lateral

margin; tergal and lateral margins of almost equal length and

forming an angle of about 145°, either margin being about half

the length of the valve. Surface of valve ornamented with fine,

closely-set, longitudinal ridges; a narrow slip along the tergal

margin is somewhat bent downwards and is devoid of ridges,

Tergum sub-rhomboidal in general outline, slightly convex,

with a delicate furrow extending from the apex to the basal

angle. Apex and basal angle acuminate, more so than is indicated

in Darwin’s original figure; scutal angle somewhat protuberant,

Carinal margin convex ; scuta] margin sinuous, longer than the

oceludent margin, which is nearly straight. Surface of valve

ornamented with numerous fine longitudinal ridges. Inner

surface, in the region of the apex, is marked on its edges with

oblique lines of growth, these indicating the extent to which the

valve projected freely. A little below the apex, nearer to the

occludent margin, are three or four small ridges ending abruptly

about one-fourth the length of the valve from the apex. These

ridges were evidently connected with the firm attachment of the

corium, or membrane which lined the inner surfaces of the valves,

and are homologous with the series of tubercles on the inner

surfaces of the terga of S. darwinianum Bosquet.

Upper latus subtriangular, slightly curved towards the scuta,

almost flat transversely, convex longitudinally ; umbo slightly

projecting, with a thick ledge formed beneath it, which thins out

towards the lateral angles; tergal margin slightly convex ; scutal

536 _ MR. T. H. WITHERS ON

margin slightly concave and about the same length as the tergal

margin, the two margins if represented by lines from the apex to

the lateral angles would enclose an angle of about 55°; basal

margin rounded, and indistinctly marked off into three lines ;

a portion of the valve on either side, parallel with the scutal and

tergal margins, is somewhat raised, and the lines of growth on

these parts are upturned sharply towards the umbo. Surface of

valve between the raised portions ornamented with several longi-

tudinal ridges.

Rostral latus.—This valve is imperfect, the upper portion being

broken off. Valve very narrow, acutely angular transversely,

about 22 times as long as wide, widening gradually from the

inner acute extremity to the rostral margin, which is abruptly

truncate; umbo apical; the inner extremity is marked by a

strong rounded keel; outer (rostral) half of valve ornamented

with fine longitudinal ridges, two fine ridges close together almost

dividing the basal margin into two equal portions. At the point

where these two ridges reach the basal margin the valve is, some-

what convex, no doubt indicating the extent to which the valve

was bounded by the inframedian latus.

Rostrum sub-triangular, strongly convex transversely, bowed

inwards ; lateral margins bounded by strong ridges; basal margin

convex ; a strong median rounded keel extends from the apex,

widens considerably towards the basal margin, and is bounded

on either side by indistinct longitudinal ridges,

Peduncle scales varying in shape from semilunar to trapezoidal,

the basal margin of the former being straight, while that of the

latter is somewhat concave; immediately below the apex the

trapezoidal scales are slightly constricted, the truncated top

appearing to overhang ; scales thickest at one-third from the base,

below which, on the inner surface, they are somewhat excavated,

this indicating the extent to which the scales were covered by the

corium, the upper two-thirds no doubt overlapping the contiguous

scales. Outer surface ornamented similarly to the valves of the

capitulum with a number of longitudinal ridges, These ridges

number about seven on the larger scales, and four or five on the

smaller scales.

Measurements.—Owing to the fact that the valves in these two

specimens of S. arcuatwm are broken, it is impossible to give

their accurate measurements. Since, however, it is desirable

that we should have some idea of the relative sizes of the valves

in an individual, approximate measurements are given where the

correct measurements cannot be obtained,

Specimen I, 13577,

Carina. Length circa 21 mm.

Scutum (right valve). Length (from apex to rostral angle)

13°5 mm.; breadth 7-4 mm.

Tergum. Length circa 17 mm.; breadth 8*°2 mm.

FOSSIL CIRRIPEDES. 537

Upper latus. Length 6-8 mm.; breadth cirea 7°5 mm.

Rostral latus. Length 6°38 mm.; breadth 2°6 mm.

Rostrum. Length 3:4 mm.; breadth 2:2 mm.

Specimen J. 13580.

Carina. Length circa 20 mm,

Tergum. Length circa 17 mm.; breadth 7-5 mm.

Upper latus. Length 5-9 mm.; breadth circa 4:2 mm.

Scales of peduncle. Length from 0°5 mm. to 1-2 mm.; breadth

1°3 mm. to 2 mm,

Remarks, and comparison with other Species—The restoration

of S. arcuatwm (text-fig. 65, 7) is based on the nearly complete

capitulum figured on the same page, and, to complete the capitu-

lum, a carinal latus and an inframedian latus have been added.

The carinal latus figured in the restoration was found amongst a

number of detached plates of S. arewatum from the Cambridge

Greensand, and is longitudinally ridged as in the valves of

S. arcuatum. It possibly belongs to the same species. In any

case the only carinal latera known to the writer from the Lower

Cretaceous rocks are of the type figured, although they evidently

belong to several different species. Judging from the hiatus

between the carinal latus and rostral latus, an inframedian

latus was undoubtedly present, and was probably very like the

homologous valve in S. fossula from the Upper Senonian.

The specimen of S. arcuatwm here figured (text-fig. 65, 6) is,

up to the present, the oldest fossil Scalpellum from which one

can gain any idea of the appearance of the complete capitulum.

It comes from the Albian (Gault) of Folkestone, Kent, and the

only undoubted valves of Scalpellum older than this occur in the

Aptian (Lower Greensand) *. These Lower Greensand forms

comprise only three species, two of which—sS. simplea Darwin

and S. accumulatum Withers—are respectively represented by

a single carina; the third, S. comptwm Withers, is represented

‘by two detached terga.

Our knowledge of these early forms of Scalpellwm is therefore

not very extensive, and the fact that they are represented by

such a small number of valves, and those only of carine and

terga, serves to emphasize the importance of this fine example of

S. arcuatum (text-fie. 65, 6).

A comparison of the carina and tergum of S. arcuatum with the

corresponding valves of S. sclidwhim Steenstrup, as figured by

Darwin (1851, p. 42, pl. i. fig. 8), shows how closely these two

species resemble each other. They are evidently related. The

* This statement is made with full knowledge that certain valves from the

Paleozoic and Jurassic rocks have been referred by various authors to the genus

Scalpellum. There is, however, not sufficient evidence to justify the reference of

these valves to the genus Scalpellum. Notwithstanding this, it is possible that

some of the Mesozoic Cirripedes referred to Pollicipes may eventually be shown

to be ancestral forms of Scalpellum, but this cannot be done until more is known of

the various valves comprising the capitulum.

Proc. Zoot. Soc.—1912, No. XXXVI, 36

DB8 ON FOSSIL CIRRIPEDES.

carine are easily distinguished, for whilst in S. arewatum the

tectum is flatly-arched transversely, and the intraparietes are

bent inwards almost at right angles, the tectum in S. solidulum

is strongly convex, and the intraparietes join to form a prominent

crest. Darwin considered the scutum of S. solidulum to be like

that of S. arcuatum, with the exception of the longitudinal ridges

being proportionally broader and further apart, closely resembling

those in the carina of S. solidulwm.

Unfortunately the only known complete capitulum from the |

Cretaceous rocks with which that of S. arcwatwm can be compared

is that of the Upper Senonian species S. fossula. In the relative

positions of the valves both species are alike, but in the structure

of the carina and scutum there are important differences. The

intraparietes of the carina of S. arcwatwm are sharply bent

inwards, the upper part of the valve is solid and must have

projected freely to some considerable extent. The carina of

S. fossula, on the contrary, projected freely but little, and the

intraparietes form a thin wall on each side of the carina. In

S. arcuatum the tergo-lateral angle of the scutum is situated

much further from the apex than in S. fosswla, and in this

respect is further removed from the more advanced scuta which

have the tergo-lateral angle almost in line with the apex, above

which the valve is added to, the umbo consequently being sub-

central. The valves of S. arcwatum are longitudinally ridged,

while those of 8. fossula ave smooth.

Affinities of the Species mentioned,

8. arcuatum is no doubt an ancestral form of a group of almost

exclusively deep-sea species, which Dr. P. P. C. Hoek* has

separated as a sub-genus under the name Arcoscalpellwm.

S. trilineatum, S. accumulatum, S. comptum, S. maximum var.

cylindraceum, and S. solidwlwm, which appear to be related to

S. arcuatum, possibly belong to the same group, but we know too

little of these species to say much about them, SS. simplex

probably does not belong here.

The species embraced by the sub-genus Arcoscalpellum have

no sub-carina, and it is impossible at present to say whether

S. arcuatum had a sub-carina or not. The Senonian species

§. fossula also comes nearest to the subgenus Arcoscalpellum, but

this species is said by Ed. Hébert? to have a sub-carina, and

in view of this it is possible that the Albian S. arcwatwm also

may have had one. Assuming this to be the case, we have two

forms agreeing in all essential characters with the forms of

Arcoscalpellum, except that they (possibly) have a sub-carina.

Moreover, the two species differ in the form of the carina, S. arcua-

twm having the intraparietes bent inwards at right angles and

* P, P. C. Hoek, Oct. 1907, Siboga-Expeditie, Cirripedia Pedunculata, p. 59.

+ Ed. Hébert, 1855, Mém. Soc. Géol. France, 2° sér, vol. v. p. 356, pl. xxviii. fig. 1

(S. gallicum = 8. fossula).

IAS diver, Jel | Wy CIV)

Wij “tee

West,Newman ad nat. chr.

FEATHERS FROM PHEASANTS, d&@.

"6 82 ‘SINVSVHHd WOU SHUSHLV aA

‘ayo Jeu pe UPWMEeh ‘soy

ISAM Wal ei ieils 1S cal

Basie ihSilra Tell Ju OVAL,

VE 8 9

West,Newman adnat chr

FRATHERS FROM PHEASANTS, 2.

PZ Ss 1o4o- Ph sayin

West,Newman ad nat. chr.

Seed By A Sl ee REN MV Re ON SINS A NEN se eee

ON EXPERIMENTAL PHEASANT-BREEDING, 539

joining, the upper part of the valve being solid and projecting

freely, while the carina of S. fossula has the intraparietes

forming a thin wall on each side of the carina, which is a more

advanced type. There is not as yet sufficient evidence to prove

whether S. fossula was derived from S. arcuatuwm or not. It is

patent, however, that from the foregoing species subsequently

arose the forms grouped as Arcoscalpellum and characterized by a

reduction in the number of plates of the capitulum, by a sup-

pression of the rostrum, and a tendency towards the reduction

of the pair of infra-median latera.

For help in connexion with this paper I wish to express my

indebtedness to Dr. F. A. Bather, Dr. W. T. Calman, Mr. C. P.

Chatwin, and Dr. P. P. C. Hoek.

Key to Species mentioned.

A. Carina without intraparietes, parietes not

reaching to the basal margin and bent almost

at right angles to the tectum ...................

B. Carina with intraparietes bent inwards and

joining, the upper part of the valve solid and

projecting freely.

1. Carina with basal margin rounded.............., S. accumulatum Withers.

2. Carina with tectum flatly-arched transversely,

and marked with numerous fine longi-

fradinmalleny lores! oh Pen emeeiee ae an

3. Carina with tectum flatly-arched transversely,

and marked with three prominent longi-

tudinal ridges, one central, and one on

either side separating the tectum from the

[DEN OTCUGES ce aetemaneente cnarcc an nee Dene meee at reena S. trilineatum Darwin.

4, Carina with tectum strongly convex trans-

versely and marked with several longitudinal

TPIUGSSEIS) isa Ips ae Act ee a a

5. Carina with tectum strongly convex trans-

versely with smooth surface..................... S. maximum var. cylindra-

C. Carina with intraparietes forming a thin wall [eeuwm Darwin.

on each side of the valve. é

1. Carina with tectum and parietes smooth,

dorsal surface and inner margin much

S. simplea: Darwin.

S. arcuatum Darwin.

S. solidulum Steenstrup sp.

arenated( tpl a en oS hastatum Darwin

2. Carina with tectum bordered on each side by

a large, protuberant, flat-topped ridge ... .. S. fossula Darwin.

28. Experimental Pheasant-breeding,

By Rose Haig THomas, F.Z.S.

[Received December 4, 1911: Read February 6, 1912. |

(Plates LXIV.—-LXVII.*)

The experiment with which I deal in this paper was under-

taken to test the truth of the result of one previously made

(P. Z.8. 1909, pp. 884-885), in which it was shown that a male

Pheasant had transmitted to his female offspring of the second

* For explanation of the Plates see p. 545.

30”

540 MRS. R. HAIG THOMAS ON

generation the female plumage of his species, under the following

scheme of mating :—

Silver 2 X Swinhoe ¢

F.1 2 X Swinhoe 2

is ——— =

(1) F.2 2 Sw. (1) ¢ Sw. (8) & Sw. hyb.

It is to be observed that the Swinhoe male transmitted the

pure Swinhoe plumage only to one of his male offspring. As

such an isolated instance, though interesting, had small value

without the support of other evidence, the following experiment

was arranged. JI chose species which have mature plumage in

five months from hatching, so as to get quicker at the results,

instead of, as in the case of ees, waiting eighteen months

for the adult plumage of F. 2. Phasianus formosus, one of the

species selected, is found only on the Island of Formosa ; somewhat

resembling P. torguatus, it differs from it in one special feature ;

in the cock, the collar, instead of being an even white ring

forming a complete circle round the neck, is formed by a deep

Vandyke on each side with a narrow connecting-line round the

back of the neck but not connected round the throat, an incomplete

circle. The other species, P. versicolor, the Japanese Pheasant, is

too well known to require description ; it has no white collar.

Below is appended a table of the differences between the males

and females of the two species which might be expected to prove

Mendelian pairs.

P. formosus 3. P. versicolor ¢.

Moult: Rapid, early. Moult: Slow, late.

Leg: Lee:

Colour: Pale bluish grey. Colour: Dark, reddish grey.

Dimension: Thick. Dimension: Thin.

Bill: Large. Bill: Small.

Crest: Pale. Crest : Dark.

Breast : Rich brown. Breast : Metallic green.

Tail: Long. Tail: Short.

All rectrices banded. Some laterals not banded.

Neck: White collar. Neck : White collar absent.

Bird; Large. Bird: Small.

P. formosus 2. P. versicolor °.

Leg: Leg :

Colour: Pale bluish grey. Colour: Reddish grey.

Dimension: Thick. Dimension: Thin.

Bill: Laree. Bill: Small.

Colour: Greyish green, paler at Colour: Darker greyish green, same

gape. all over.

Crest; Pale ground, black oval | Crest: Dark ground, — irregular

spots. : black patches.

Breast : Pale buff, no pattern. Breast : Dark buff, thickly patterned.

Tail; Banded pattern on pale | Tail: Banded pattern on dark

ground, ground.

Bird: Large. Bird: Small.

Reg: Large. Egg : Small.

Calin temperament: quiet, tame. Excitable temperament: wild, untame-

able.

EXPERIMENTAL PHEASANT-BREEDING. HAL

Scheme of mating in second experiment :—

P. formosus 9 X% P. versicolor 3

oS Ee Ee

(3) F.1 ¢ F129 “Y” F.12“Z” x P. versicolor 3

(5) F.2 ¢ Ve. (2) Ve. hyb.

The second experiment confirmed the results obtained in the

first.

Four skins of the F, 2 2’s are exhibited (the fifth female has

been kept to breed from), together with skins of pure Versicolor

and pure Formosan females, also the skin of the F.1 9 “Z” parent

of F. 2. If these are all turned breast uppermost, it is seen at a

glance that Versicolor pattern and coloration are present to a

certain extent in F.1 9 “ Z,” mother of the F. 2 generation, and

that the F. 2 females appear to be identical with Versicolor in size,

pattern, and coloration, except for small differences which might

be found existing between individuals of any species. Besides

plumage, these five hens had the habit and temperament of the

Versicolor, the leg-colour and eye-skin also.

The records of this experiment constantly refer to the quick

sharp movements, the wild scared appearance of the F, 2 females,

so characteristic of the habits and ways of.the untameable

Versicolor. If the pen was entered for any purpose, even when

exercising the greatest care, there was always a chance the birds

might break their necks in their terrified flights and violent

dashes against the wire netting, and in this manner one or two

were scalped to the bone.

A list of the Versicolor characters found in the F. 1 females is

interesting, for there are certain characters which can only be

classified in the living bird: Leg-colour, eye-skin, bill, moult, habit,

temperament, voice.

F.1 29 “Z” (P. formosus 2 x P. versicolor ¢).

Parent of Eye-skin. ft

F. 2 (Fo. x Crest. |

Ve. x Ve.). Neck. bus

Flank. ss

Breast. -

Interscapulars. 4 ie

Scapulars. SS

Secondaries. SS

Back.

Size of egg. 4

EN ing OR BL lech Formosan.

Bill. ~

Primaries. ] ie

Dimension. =

Leg.

549 MRS. R. HAIG THOMAS ON

F190 «y

Parent of HKye-skin.

inter se K. 2. Crest.

Neck.

Breast.

Flanks.

Interscapulars. e

Scapulars. |

Secondaries. |

ere

Versicolor.

Back.

Size of eg

1 By ak tm tah bee teit Formosan.

Bill Gs ~

Primaries. =

c = LS

Dimension. | SS

Tail.

In both F. 1 females the size of egg was transmitted by the

Versicolor male.

1 Oe Crest. ‘

Neck.

Flank. |

Breast.

Legs. | .

Bill,sizeand colour. | $

Interscapulars. =

Scapulars. Sea

Secondaries. 2

Primaries. 2

Tail. az

Back. |

Dimension.

Habit of moult. |

Temperament. 9)

The fifth female, reserved to breed from, has a pale grey stripe

down the back of the shank of one leg.

The presence of a mosaic of pale grey and dark grey seen in

the legs of F. 1 in this experiment led to the inference that the

two parents had severally pale grey legs and dark grey legs,

which an immediate examination of the Formosan and Versicolor

species confirmed. A curious independent double segregation of

allelomorphs was observed in the autumn of 19]1 in the crests of

the F. 1 males. The centre feathers were dark—colour Versicolor ;

fully developed, rapid early moult—habit Formosan : the feathers

on the outer edge of the crest were pale—colour Formosan ;

EXPERIMENTAL PHEASANT-BREEDING, 543

undeveloped, many still in the quill—habit Versicolor: two

Mendelian pairs coupled in each parent yet repulsing one

another in the same area in F. 1. An examination made at the

same time of the Formosan male showed the crest fully developed,

and an inspection of the Versicolor male showed the crest

undeveloped, mostly stil] in the quill.

F. 1 generation is to me always the most interesting in these

artificial Pheasant crosses. For Mendelian segregation already

shows, and it 1s sometimes possible to select the strain of parentage

desired to reappear more strongly in the F. 2 generation, Also

F. 1 occasionally produces remarkable mosaics of sex—a sort of

sex-hybridism accompanied by sterility and extraordinary develop-

ments of plumage in the female, phenomena I will not touch upon

now. ‘To illustrate more clearly the points I wish to bring to your

notice, here is a selection of secondaries (3rds from the last primary)

extracted from the wings of the parent species and of all the

birds connected with the experiment (P]. LXIV.): the Formosan

male secondaries have a peculiar Vandyke pattern like a feather

laid on a feather, and the female secondaries of this species are

banded, both are extremely light in colour. The Versicolor male

secondaries have a mottled grey oblique banding on a very dark

grey ground; the female secondaries of this species are rich

brown, with wide bands of darker brown ; also placed in the same

frame are the secondaries of F.1 and F.2 males and females.

After examining the parents’ secondaries, we perceive that the

Versicolor male has transmitted to his female offspring of the

F. 1 generation the female secondaries of his species, and that

conversely the Formosan female has transmitted to her male

offspring of the F. 1 generation a pattern resembling the male

secondaries of her species on one vane, though not on the other

vane of the feather. In the F. 2 generation (‘Fo x Ve. x Ve.”

Pl. LXV.) the influence of the Versicolor male on his female

offspring continues in pattern and size and more or less in colour,

for in F. 2 females the secondaries seem to be Versicolor, with

slight differences of colour not unlikely to be found between

individuals of the same species; whilst the F. 2 male secondaries,

though most resembling Versicolor, are somewhat hybrid in size

and pattern, and still show slight traces of Formosan influence.

A selection of interscapulars extracted from the parents and

from the two generations of the Formosan Versicolor cross is

shown ; the females are in one frame, the males in another, and in

these the same phenomena appear. Those of the F. 1 females

seem to be Versicolor in pattern and colour, as also do the five

F. 2 females, with certain modifications that might be readily

found amongst individuals of a pure race (Pl. LXYI.). The

frame containing the male interscapulars shows amongst the F. 1

males a Formosan and also a hybrid pattern, whilst the F. 2 males

are also hybrid (Pl. LX VII).

The phenomenon of pattern-transference has occurred in all

my Pheasant crosses ; sometimes from the male to the female, or

544 MRS. R. HAIG THOMAS ON

conversely from the female to the male, or a pattern may be

transferred from one area in the parent species to another area

in the F. 2 offspring. I have noticed that these pattern-tians-

ferences are inclined to remain fixed and constant.

Colour-transference also takes place, and sometimes where it has

occurred seemed to inhibit the appearance of pattern.

One instance was noticed where the breasts of the males of the

two races crossed, 7h. amhersti and Th. picta, differing widely in

colour, both colours were found on the breast of F. 2 Th. obscura,

the red of picta overlying the metallic green of amhersti.

These appear to be the results of the second experiment :—

(1) The male parent transmitted to his F. 1 female offspring

much of the female plumage of his species and the

dimension of the egg.

(2) The female parent transmitted to her F. 1 male offspring

much of the male plumage of her species.

(3) In the F. 2 generation, the offspring of F. 1 female x

Versicolor male, the Versicolor male seems to have trans-

mitted every character—bill, leg-colour, plumage, habit, and

temperament—of the female of his species to his F. 2

female offspring, whilst he has not transmitted every

character of the male of his species to his F. 2 male

offspring; repeating exactly the results of the original

experiment with genneus.

Are we, then, to suppose that some of the gametes of this

Versicolor male contained all the factors representing the tem-

perament and habit, the colour, pattern, and dimension of plumage,

leg, and bill, and the bulk of the female of his species and

even the factor for size of egg, with the one exception of the

factor for the sex to which these belonged ?

To the practical experimentalist, to the non-mathematical

simple observer, the hypothesis is difficult to conceive.

These phenomena seem to be of the nature of a sex-limitation

opposed to expectation.

Thave read with much interest Mr. Doncaster’s account of game-

togenesis in the Gall-fly, also his researches on sex-limitations

published in ‘Genetics,’ and am interested to know how he

would consider the above facts in relation to his theory of sex:

Male gametes ¢ O female gametes ¢ 2 with selective fertilisation

between the male gamete O (a non-determinant of sex) and the

female gamete ¢.

I hope the material collected in these two experiments may be

thought of sutticient importance for the higher students of

Genetics to give it some attention, when probably the apparently

complicated problem will receive a simple explanation.

EXPERIMENTAL PHEASANT-BRIEEDING. 545

EXPLANATION OF THE PLATES.

Pratt LXIV.

6 & §% Secondaries (8rd from primary) of the two Parent species

and of F. 1 (Fo.x Ve.).

1. P. versicolor 2. Horizontally set, broad bands, bifurcated ends; ground-

colour brown,

2. P. versicolor §. Mottled light bands V-shaped, apex towards rachis ; ground-

colour dark grey.

3. P. formosus 9. Obliquely set narrow bands, pointed ends; ground-colour

nearly white.

4. P. formosus 8. Vandyke pattern, light, mottled; ground-colour light grey.

5. F.19 “Z” (P. form.2 XP. vers.$). Colour, banding, length of feather,

(Parent of F. 2, Fo.x Ve.x Ve.). versicolor; a similar: mottling to that

here seen is found on the 3rd Second-

aries of a pure versicolor 9 unrelated

now living in my pheasantry.

6. F.1g“C” (2. form. 9 XP. vers. 8). Lett vane Vandyke, Formosan ¢ pattern.

Right vane hybrid banding.

Colour hybrid.

Prats LXY.

6 & & Secondaries (8rd from primary) of F. 2 (Fo. Ve. X Ve.).

1. F.2 g¢ “G” (F.1¥Fo.XVe. 9 XP. vers. $). Pattern of banding nearest to

versicolor 6.

Colour hybrid.

2. 3. 4. 5. 6. F.2 9 9(F.1 Fo.xXVe. 9 XP. vers. ).

eo Deane tree | eed [avec HY 220 fhe eb 2?0 |

Although slight differences exist between these five secondaries, yet the length

and colour of them, the horizontal setting of the bands, their breadth and bifurcated

eds, are all pure female versicolor characters.

PratEe LXVI.

2 posterior interscapulars of the two Parent species, of F. 1 and F. 2.

1. P. versicolor 2.

2. P. formosus 2 (parent of F. 1).

3. F.12“Y” (parent of inter se F.2). Pattern transference. Pattern found on

several anterior interscapulars of P. ver-

sicolor 2.

4. ¥1 2 “Z” (parent of F. 2, Fo.X Ve. x Ve.).

5. F.2 9 “EK” (moult completed). 9 versicolor posterior interscapular pattern.

6. F.2 9 “EF” (moult completed). Pattern transference. The pattern seen on

7. F.2 2 “H” (moult completed). these two feathers is found on some anterior

interscapulars of 2 versicolor.

8. F.2 9 “C” (aged 4 months, moult incomplete). Pattern found amongst

posterior interscapulars on 2 versicolor.

9. F.2 9“B” (moult completed). Pattern found amongst anterior and posterior

interscapulars on 2 versicolor.

Note, July 2nd, 1912.—A number of posterior and anterior interscapulars extracted

from a living female versicolor in my pheasantry showed all the patterns figured

on Plate LX VI.

Prate LXVII.

6 posterior interscapulars of the two Parent species, of F. 1 and F. 2.

ei ; P. versicolor g. ; a } P. formosus 3.

46 SIR GEORGE H. KENRIGK ON

5. F.1 g “A” (Fo.XVe.). Similar patterns found on posterior interscapulars of

6 versicolor.

6. KF. 1 ¢ “B” (Fo.XVe.). Coarser lines but same pattern as found in some

posterior interscapulars on ¢ versicolor.

7. F.1 $6 “C” (Fo.XVe.). Similar pattern found amongst posterior inter-

scapulars on g versicolor.

8. F.2 ¢ “A” (Fo.x Ve.X Ve.).

9. F.2 g “G” (Fo.XVe.X Ve.). Pattern found amongst interscapulars on

6 versicolor.

Note, 2nd July, 1912.—A number of posterior and anterior interscapulars were

extracted from a male versicolor now living in my pheasantry ; amongst these were

found all the various patterns pictured on Plate LXVII. There is a difference

between central and lateral interscapulars, the centrals have the pattern both sides,

the lett laterals have the pattern on the lett side, the right laterals have the pattern

on the right side.

27. A List of Moths of the Family Pyralide collected by

Felix B. Pratt and Charles B. Pratt in Duteh New

Guinea in 1909-10 ; with Descriptions of new Species.

By Sir Georce H. Kewrics, F.Z.8.

[Received January 30, 1912: Read March 19, 1912.]

(Plate LX VIIL*)

INDEX.

Systematic :— Page

Macalia arruensis, Sp> MWe 0.0... ..cecesee ses sseccs eee DAT

ME SOU UQUISHSp SNe, Ae 5: Sa eee ces cde ea

DE CCOSUBES OAS 05 WG Acs avodeGabe.anoscemdeneebuban ndanasoce ) OL

SUUBPOCUE) FUN AOUIS, So Warde coacee ddcnsntes dae enn cebasoncs eae 548

Perisyntrocha suffwsd, Spo. ...... cece eee eee 548

Ambia novagwinensis, SP. Wei... ci. ecc cence ccc cce eee ee 549

Margarosticha plumbealis, sp. n. ............-000..-.. 54D

Brevicella emarginata, gen. et sp.n. ................. 549

NSceliodes grisealisy Sp. Ms ince -ekesaer een) eee OO)

Entephria miosward, Sp. W. ......-....ceteceeee sees DDO

JOR GRO EGI SS S105 In Regarenonnondnnsndesdseduedondesc sebeccccs” SU)

LEGGY AC UTMOSH |S) Mel lentes eke cere eee ee OSI

SMG Ri NCI S05 Ie. geenosssadeAcoepagbRacKinsosc-2c.54ne Dah

Glyphodes magnificalis, Sp. MW. ...... 02.00 eee eevee. 52

GAG TACINICLAy SP. Wenn ese eee ee eee OO

G. pseudocesalis, SP. Wy... .2.2-..22ecstsscreeseee-se--1. | ODA

Omiphisa wariegata, 8p: We ie. -...c.ccese see see ones 553

Noord a Cnfaensis, Sp. ie. ke ees eee nee ee ODS

Pilocrocis angulifera, sp. n. cies Ogto sesh coe 553

Pyrousta flammealis, SPW. 6... ee.eeeeteee esses. 8O4

Variation :—

CHT NOOUES TO OUPARE: ceo cab obo boise. ondounseséeyaebasn aeceen 5a4

Calamachrous albipunctalis ............ccc cesses BSA

After some considerable experience in British New Guinea,

the two sons of Mr. A. E. Pratt made several attempts to explore

various portions of the Dutch territory, and although disappointed

in some directions they made a very successful ascent of the Arfak

Mountains, formerly visited by d’Albertis, and spent a considerable

* Yor explanation of the Plate see p. 555.

ght del.et lith.

orace Kni

Lal

MOTHS FROM DUTCH NEW GUINEA. AT

time in the vicinity of two lakes, which proved to be excellent

collecting-grounds for the lepidopterist.

Although for the most part the insects were the same as those

collected in British New Guinea in the south, a fair number of

different species occurred, a list of which I give her ewith, together

with descriptions of new species.

The island of Mioswar, on the north-east coast, was also visited ;

but the nights were cmfayone ole for collecting and few moths

were tolken although it proved te be an excellent locality for

Lycenide.

The specimens were nearly all taken at light, as in the case of

those enumerated in my former list.

Sub-fam. HPreASGHIINA,

MAGCALLA SYRICHTUSALIS Walk.

MACALLA ARRUENSIS, sp.n. (PI. LX VIII. fig. 3.)

Head, legs, antenne, and palpi fawn-colour; abdomen the

same, orn darker segments at the tuft; tarsi jane ringed with

paler.

Fore wing fawn-colour, with a dark brown basal patch out-

wardly oblique followed by a whitish irregular spot on the costa

and reaching as far as the bottom of the cell. Following this on

the costa is a darker mark developing into a faint central shade.

Beyond this is a curved postmedial line ending in a white mark

on the costa, and at the apex there is a dark patch.

Hind wing pale ochreous ; fringes pale.

Exp. 52 mm.

From Arfak Mts., 4000 ft.; but there is an undescribed spe-

cimen from Arru in the British Museum.

MACALLA OLIVALIS, sp.n. (Pl. LX VIII. fig. 1.)

Head, legs, palpi, and thorax olive and white ; antenne brown ;

abdomen pale grey.

Fore wing olive-green with white markings ; three equidistant

white spots” on the costa, an antemedian interrupted white band,

a median interrupted white line, and a postmedian distinct white

line preceded by an irregular white blotch which does not reach

the costa or the inner margin; a subterminal white line; fringe

orange and white; underside of fore wing with a number of large

silvery scales in the cell and at the base of the wing.

Hind wing grey, semi-hyaline ; fringes paler.

Exp. 29 mm.

Arfak Mts., Dutch New Guinea.

MACcALLA MIoswARI, sp.n. (PI. LX VIII. fig. 5.)

Head, thorax, antenne, and palpi reddish ochreous ; legs darker,

ringed with ochreous.

Fore wing pale ochreous with chestnut patches, one at the base

548 SIR GEORGE H. KENRICK ON

terminating in a dark antemedian line, one at the apex, and another

at the angle. Between these two is an extension of the median

pale ochreous area extending to the hind margin; an angulated

dark postmedian line ; a small dark central spot and a dark mark

on the inner margin between the lines.

Hind wing grey. Fringes of both wings bright chestnut.

Exp. 27 mm.

Island of Mioswar, Dutch New Guinea.

STERICTA RUREALIS, sp.n. (Pl. LX VIII. fig. 2.)

Head, antenne, and palpi pale ochreous; patagia chestnut ;

abdomen pale at the base, but with chestnut tuft; legs dark,

paler at the joints.

Fore wing ochreous, with a yellower shade along the costa and

the raised scales red. A basal patch reddish chestnut, bounded

obliquely and extending along the inner margin nearly halfway.

An apical chestnut patch and a smaller patch at the angle.

Hind wing pale ochreous, with a fine marginal line darker.

Fringes ochreous, but darker at the apex.

Exp. 44 mm.

Artak Mts., 4000 ft.

Sub-fam. PyraLin”®.

CURENA EXTERNALIS Walk.

Sub-fam. Hyprocampin&.

AULACODES BRUNNEALIS Hmpsn.

AULACODES METALOXALIS Hmpsn.

NyMPHULA CHRYSEIS Meyr.

NYMPHULA POLYSTICTALIS Hmpsn.

CLUPEOSOMA POLUSALIS Walk.

PILETOCERA GGIMISALIS Walk,

PILETOCERA DIPLATYLA Walk.

PILETOCERA TORSICUTALIS Hmpsn.

NEOGENESIS FLAVOPLAGIALIS Hmpsn.

PERISYNTROCHA SUFFUSA, sp. n. (PI. LXVITI. fig. 4.)

Head, antenne, palpi, and legs pale ochreous ; thorax darker,

with paler patagia; abdomen grey, with pale band at base and

paler rings at the segments.

Fore wing hyaline, pale golden; a darker margin to the costa.

A patch of dark scales at end of cell, developing into a triangular

fuscous mark reaching to the inner margin; the apical area of

the wing is occupied by a fuscous patch, with the exception of a

pale patch extending from the costa to the central nervure.

MOTHS FROM DUTCH NEW GUINEA. 549

Hind wing pale with fuscous margin, broad at apex but dis-

appearing at the angle ; fringes pale.

Exp. 24 mm.

Arfak Mts., 4000 ft.

AMBIA NOVAGUINENSIS, Sp. n. (Pl. LX VIII. fig. 6.)

Head, legs, antenne, palpi, and thorax brown.

Fore wing whitish, with numerous brown markings ; two ante-

median lines containing a white spot near the costa : a dark spot

at end of cell, with a white triangular centre; an oblique line

running from near apex to middle of inner margin, followed by

a darker space containing five triangular white spots with dark

outer margins.

Hind wing white, with an irregular dark median line and a

dark subterminal line.

Exp. 18 mm,

This insect has veins 4 and 5 stalked in both wings.

Warmasin Lake, Arfak Mountains, 6000 ft.

MARGAROSTICHA PLUMBEALIS, sp. n. (Pl. LX VIII. fig. 9.)

Head, legs, antenne, and thorax pale chestnut ; patagia white ;

abdomen bright chestnut, with black tuft.

Fore wing bright chestnut, with silvery-white blotches: the

first is square and extends from the median nervure to the hind

margin, the second is triangular and near the apex; before the

apex itself is a narrow white streak.

Hind wing chestnut, with a broad white transverse band;

beyond this is a lead-coloured round patch reaching the outer

margin : it is in reality black, with a number of scattered white

scales; on the outer edge of this are three black dots, each having

a centre of chestnut, and between them silvery-white dots.

Underside silvery, with the markings of the upper wings

showing through ; fringes darker,

Exp. 22 mm.

Arfak Mts., 4000 ft.

New genus under sub-fam. Hydrocampine :—

BREVICELLA (type emarginata, ° ).

Palpi porrect, twice the length of head, not scaled : antenne

sunple; legs not hairy; tibie with usual spines. Hind wings

with long hairs on parts of the costa.

Venation—Fore wing: 2 and 3 stalked; 4, 5, 6,7 from cell;

9 and 10 stalked, with 8-11 present. Hind wing: 2 and 3

stalked slightly ; 4, 5, 6 from cell; 7 stalked with 8,

BREVICELLA EMARGINATA, sp.n. (Pl. LX VIII. fig. 8.)

Body and wings dark brown ; a postmedian curved line bordered

within by seven triangular whitish dashes; at the upper end of

550 SIR GEORGE H. KENRICK ON

this line is an oblique intensely black dash reaching nearly to the

costa.

Hind wing with an antemedian whitish spot and a postmedian

darker band. There is a narrow darker marginal line to both

wings, and the fringes, which are dark, are strongly emarginate.

Exp. 30 mm.

Momi-Arfak, 4000 f§.

ScELIODES GRISHALIS, sp.n. (Pl. LX VIII. fig. 7.)

Head, antenne, legs, and palpi ochreous ; thorax and abdomen

pale brown ; tuft pale.

Fore wing ochreous, powdered with black scales and with three

white angulated streaks, the first much angulated before the middle,

the second beginning in the middle of the costa and narrowing to

a point on the lower edge of cell, the third also from the costa

extending as an angulated fine line to the middle of the wing; a

subterminal ill-defined whitish band, in which is a dark sub-

terminal line followed by a dark cloud.

Hind wing whitish, with two dark dots on the disc, a dark

subterminal line followed by a dark cloud ; fringes brown.

Exp. 30 mm.

Arfak Mts., 4000 ft.

Sub-fam. Pyraustin«.

XANTHOMELENA SCHEMATIAS Meyr.

ENTEPHRIA GLAUCIAS Meyr.

ENTEPHRIA MIosWARI, sp. n. (PI. LXVIII. fig. 10.)

Head, antenne, legs, and palpi silvery white. Thorax and

abdomen white, the latter with black markings on three or four

of the last segments. Fore wing white, a small soot-black spot

near the base of the cell and a larger round one at the end.

Before the apex is a costal black spot, from which springs a pale

transverse ochreous line reaching the inner margin. In some

specimens there isa dark spot at the apex, and in all the hind

margin is dark.

Hind wing white, with a small dark spot near the base and a

thin median line; hind margin and fringes darker,

Exp. 24 mm.

Island of Mioswar, Dutch New Guinea.

ENTEPHRIA GRISEALIS, sp.n. (Pl. UX VIII. fig. 12.)

Head, palpi, antenne, and legs silvery white; thorax and

abdomen white, the last three segments darker above.

Fore wing white shaded with grey; two transverse ochreous

antemedian lines, the first somewhat obscure; a median line from

inner margin to vein 2, where it disappears in a broad shade; a

transverse distinct line from costa to vein 2 beyond the cell,

MOTHS FROM DUTCH NEW GUINEA. 551

Hind wing white; a transverse dark line from angle to costa

anil another from costa beyond cell to vein 2.

The margins of both wings are bordered by a dark, thin, double

line enclosing a paler portion. Fringes white.

Exp. 32 mm.

Arfak Mountains, Dutch New Guinea.

REHIMENA CIssopHoRA Turner.

AINCKENIA LOPHOCERALIS Hmpsn.

AGROTERA PyRosticraA Hmpsn.

PAGYDA BOTYDALIS Snell.

PacypDA Fumosa, sp.n. (Pl. LX VIII. fig. 13.)

Head, legs, antenne, and palpi straw-colour ; thorax and abdo-

men rather more orange,

Fore wing straw-colour, a short, orange, transverse line near

base; an antemedian and a postmedian line, the middle line

curved, dark orange; an oblique curved band of pale orange from

apex to hind margin and a paler marginal band.

Hind wing suffused with a smoky patch in which there are

three whitish transverse lines and a dark fine marginal line;

fringes paler.

Exp. 22 mm.

Arfak Mts., 4000 feet.

CAPRINIA DIAPHANALIS Walk.

PHRYGANODES EREBUSALIS Hmpsn.

PHRYGANODES ANALIS Hmpsn.

PHRYGANODES UsITALIs Guen.

PHRYGANODES TETRAPLAGALIS Hmpsn,

PHRYGANODES CENTRABALIS Hmpsn,

NACOLEIA PERDENTALIS Hmpsn.

PRORODES MIMICA Swinh.

SYLEPTA PoLYDonTA Hmpsn.

SYLEPTA LEUCoDoNTA Hmpsn.

SYLEPTA DINAWA, sp.n. (PI, LX VIII. fig. 11.)

Head, thorax, legs, antenne, and palpi dark brown ; abdomen

dark brown, with a conspicuous pale ochreous tuft and a pair of

black tufts of hair at base. Fore wing pale ochreous clouded with

dark grey ; an irregular dark antemedian line and a vague post-

median line with a branch towards the angle; the apex is broadly

552 SIR GEORGE H. KENRICK ON

mareined with dark; below the costa at end of cell is a lunated

mark and there is anather smaller one nearer the base.

Hind wing pale semihyaline, with traces of dark ante- and post-

median lines, the hind margin with a broad dark shade; fringes

pale.

Exp. 42 mm.

Dinawa, British New Guinea.

GLYPHODES POLYZONALIS Hmpsn.

GLYPHODES MAGNIFICALIS, sp. n. (Pl. LX VIII. fig. 17.)

Head, palpi, and antenne black; legs and underside golden.

Thorax black, with collar and patagia ochreous ; abdomen black

above.

Fore wing pale ochreous ; an oblique black band beginning as a

fine line on the base of the costia reaches a point two-thirds along

the inner margin, from this there stretches a nearly vertical

black band with curved outer side to the costa. There is a sub-

terminal cloud of brown intersected by two fine metallic purple

lines, on the inner side of this are two triangular black marks

united in the middle. All the black marks are shot with metallic

blue.

Hind wing pale ochreous, with an interrupted dark postmedian

line. On the middle of the edge of the wing are three white-

centred black dots on a faint bright line.

Exp. 40 mm.

Arfak Mts., 4000 feet.

The only specimen is a female.

GLYPHODES NIGRICINCTA, sp. n. (PI. LX VIII. fig. 16.)

Head, palpi, and antenne dark brown; legs white, the joints

dark brown on the outer side; front of thorax black ; patagia

white ; abdomen grey, with a black band at base.

Fore wing white, a dark grey spot in the middle of the cell and

a second at the end; the outer half of the wing very dark grey.

Hind wing white, with grey lunule and dark grey apical patch.

Fringes smoky, fading into white towards the angle of the hind

wing.

Exp. 26 mm.

Arfak Mts., 4000 feet.

GLYPHODES PSEUDOCHSALIS, sp. n. (Pl. LXVIII. fig. 18.)

Differs from cesalis Wlk. in the absence of hyaline patches, in

having the fringes unspotted whereas in cesalis the dark spots

give the wings a distinctly emarginate appearance, and in the

markings of the fore wing: these consist of a basal line indicated

by dots, antemedian, median, and postmedian lines, each of which

is double and the inner lines of the median and postmedian are

connected. In ewsalis these double lines are wider apart, more

irregular, and filled in with darker instead of with the ground-

colour as in pseudocesalis. Beyond the postmedian line in cesalis

MOTHS FROM DUTCH NEW GUINEA. 553

is a dark patch with four pale lunules, which are absent in pseudo-

ceesalis,

Exp. 30 mm.

Arfak Mts., Island of Mioswar, Mt. Goliath, and British New

Guinea.

It is possible that there is another allied species, but it may be

simply a larger form of cwsalis.

Py@osPrLa tyres Cram.

CrocrpoLoMIA BINOTALIS Zell,

SAMEODES POLYTHLIPTALIS Hmpsn.

OMPHISA REPETITALIS Moore.

OMPHISA VARIEGATA, sp.n. (PI. LXVITI. fig. 19.)

Head, legs, antennex, and palpi chestnut-brown ; thorax chest-

nut-brown, the patagia pale; abdomen darker, spotted with paler

on back.

Fore wing pale ochreous with brown markings.

Fore wing: basal patch brown, an antemedian and a postmedian

line enclosing a darker band which does not reach the costa, but

which surrounds a conspicuous subtriangular pale spot near the

end of the cell; beyond this is an angulated line followed by a

darker shade; costa with six lunules,

Hind wing pale, with an irregular sharply-defined median band

and a double outer dark line.

Exp. 32 mm.

Island of Mioswar, Dutch New Guinea.

Noorpa NigRripuNcTALIs Hmpsn.

NoorDA ARFAKENSIS, sp. n. (Pl. LX VIII. fig. 14.)

Antenne and palpi brown; legs white, with brown tips to the

joints; head ochreous; thorax white ; abdomen whitish, the last

segment grey, anal tuft white.

Fore wing creamy white, the costa reddish ; a dark brown dot

at base, another at the beginning of the cell, and a third at the

end; a dark brown annulated subterminal line; the remainder

of the wing to the outer margin of a chestnut colour mottled

with darker.

Hind wing white. Fringes pale brown, shading into white at

the angle of the hind wing.

Exp. 30 mm.

Arfak Mts., 4000 feet.

DAUSARA AMETHYSTINA Butler.

Pinocrocrs (a subdivision of Crocidophora) ANGULIFERA, sp. n,

CELE TTT fae V5. yee

Head, legs, antennz, and palpi straw-colour; thorax and abdo-

men pale golden, the last few segments becoming fuscous.

Proc. Zoou. Soc.—1912, No. XX XVIL. 37

554 SIR GEORGE H. KENRICK ON

Fore wing pale golden, a very short basal dark line, an oblique

antemedian line, a sharply angulated median Jine, a regular

curved postmedian line (all these are dark fuscous and the last is

lost in a fuscous patch which extends from the middle of the hind

margin halfway up the hind margin). i

Hind wing pale at base, with the whole of the outer portion

dark fuscous, the boundary between these is marked by a very

irregular angulated line. Fringes rather paler.

Exp. 28 mm.

Arfak Mts., 4000 feet.

APHYTOCEROS GROSALIS Meyr.

CuRICTA OPPOSITALIS Walk.

PYRAUSTA OCCULTILINEA Walk.

PYRAUSTA ALENTIALIS Snell.

PyRAUSTA DEDUCTALIS Walk.

PYRAUSTA FLAMMEALIS, sp.n. (Pl. LX VIII. fig. 20.)

Male with a tuft of hair at base of fore wing below and a strong

fold of costa extending to end of cell; female with a smaller tuft

of hair; both sexes have the base of the antennz surmounted by

a tuft of hair, but this is larger in the male. Head, legs, and an-

tenn pinkish brown, also palpi; abdomen ochreous above, darker

below; thorax and patagia flame-colour.

Fore wing flame-colour, with three transverse fine dark lines: the

first ante median and oblique, the second median and angulated in

the middle, the third angulated and not reaching below the third

nervure; all the margins of the wing are darker.

Hind wing uniformly straw-colour with fringes slightly darker ;

fringes of fore wing dark. In the female the flame-colour is

replaced by Indian yellow.

Exp. 30 mm.

Arfak Mts., 6000 feet.

In addition to these species I have figured two varieties of

species already described :—

Pl. LX VIII. fig. 21 represents a form of Glyphodes pfeiffere in

which the marginal border to both wings is much deeper than any

specimen I have received from New Guinea.

Pl. LXVIII. fig. 22 represents a form of Calamachrous albi-

punctalis in which the deep maroon-purple of the wings of this

species is replaced by bright chestnut.

Both of these forms are from the island of Mioswar.

Although in some cases a good series of the species was obtained,

in others only single specimens or two examples were taken, so

that the differences in sex could not be ascertained.

From this circumstance I am inclined to think that many other

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 555

species would turn up if the localities could be worked throughout

the year, and I believe any collector willing to face the difficulties

of a camp life would be amply rewarded by the results.

I desire to tender my sincere thanks to Sir George Hampson

for advice and assistance in regard to species of this large group

and for permission to study the excellent collection at South

Kensington.

EXPLANATION OF PLATE LXVIII.

Fig. 1. Macalla olivalis. Fig. 13. Pagyda fumosa.

2. Stericta rurealis. 14, Noorda arfakensis.

3. Macalla arruensis. 15. Pilocrocis angulifera.

4, Perisyntrocha suffusa. 16. Glyphodes nigricincta.

5. Macalla mioswari. 17. Glyphodes magnificalis.

6. Ambia novaguinensis. 18. Glyphodes pseudocesalis.

7. Sceliodes grisealis. 19. Omphisa variegata.

8. Brevicella emarginata. 20. Pyrausta flammedlis.

9. Margarosticha plumbealis. 21. Glyphodes pfeiffere (var.).

10. Entephria mioswari. 22. Calamachrous albipunctalis

ll. Sylepta dinawa. (var.).

12. EHntephria grisealis.

EXHIBITIONS AND NOTICES.

April 2, 1912.

Dr. A. Smiru Woopwarp, F.R.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 months of

February and March, 1912.

FEBRUARY.

The registered additions to the Society’s Menagerie during the

month of February were 128 in number. Of these, 50 were

acquired by presentation, 29 by purchase, 13 were received on

deposit, 19 in exchange, and 17 were born in the Gardens.

The number of departures during the same period, by deaths

and removals, was 236.

Amongst the additions special attention may be directed to :—

1 Snow-Leopard (Felis uncia) and 1 Musk-Deer (Moschus

moschiferus), from Nepal, presented by H.M. Tar Kine on

February 29th.

2 Snow-Leopards (Helis wncia), from Kashmir, presented by

Capt. G. Douglas Oliver, F.Z.8., on February 26th.

2 Jaguars (Felts onca), born in the Menagerie on February 20th.

2 Striped Hyenas (Hyena hyena), from N. Nigeria, pre-

sented by H. P. Lobb, Esq., on February 4th.

1 Spurrell’s Dormouse (Graphiurus spurrelli), en Duiliena,

556 DR. R. W. SHUFELDT ON VIRGINIA OPOSSUMS.

Gold Coast, new to the Collection, presented by Dr. H. G.

F. Spurrell, F.Z.S., on February 12th.

1 White-capped Redstart (Chimarrhornis leucocephalus), from

the Himalayas, presented by Alfred Ezra, Esq., F.Z.S., on

February 29th.

Marca.

The registered additions to the Society’s Menagerie during

the month of March were 110 in number. Of these 45 were

acquired by presentation, 32 by purchase, 21 were received on

deposit, 8 in exchange, and 4 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 192.

Amongst the additions special attention may be directed to :—

1 Wallich’s Deer (Cervus wallichii) $, new to the Collection,

1 Burrhel (Ovis burrhel) 3, 1 Tahr (Hemitragus jemlaicus) 9,

and 6 Tibetan Mastiffs (Canis familiaris), from Nepal, presented

by H.M. the King on March 13th and 25th.

1 Alpaca (Lama pacos) 3, from Peru, received in exchange on

March 5th.

1 Racket-tailed Drongo (Dissemurus paradiseus), from India,

presented by Alfred Ezra, Esq., F.Z.S., on March 14th.

1 African Tantalus (Pseudotantalus ibis), from Southern

Nigeria, presented by Sir Walter Egerton, K.C.M.G., F.Z.8., on

March 15th.

1 Ashy-headed Goose (Chloéphaga poliocephala) bred in

Holland, received in exchange on March 22nd.

3 Prairie-hens (Tympanuchus americanus), from North America,

received in exchange on March 21st.

Mr. R. I. Pocock, F.BR.S., F.LS., F.Z.S., Superintendent of

the Gardens, exhibited a lantern-slide of two Polar Bear cubs

(Ursus maritimus) born in the Gardens in November 1911, and

made some remarks upon the causes of the difficulty experienced

in Zoological Gardens in rearing the offspring of this species.

Mr. C. Tare Reaan, M.A., F.Z.8., exhibited some lantern-

slides, prepared from photographs taken by Dr. F. B. Sumner, of

a Mediterranean Flatfish (Platophrys podas) on sand, gravel, and

various artificial backgrounds, showing its power of changing its

colour and markings to resemble the ground on which it lies.

Dr. R. W. SHuretpt, C.M.Z.S., sent for exhibition the skins

of two young Virginia Opossums (Didelphis virginiana). These

specimens were each about ten weeks old and belonged to the

same litter.

The bones of the entire skeleton at this age were quite

pins

Prise

Gers

ier

S <i

2 2 & iSi2 el ina.

Photos by D. Seth-Smith. Bale & Danielsson Lt¢ Imp.

YOUING (CAIRIVAINUA CIRIUSIVATIA,

MR. D, SETH-SMITH ON A YOUNG CARIAMA. 557

elementary in character, especially the terminal vertebre of the

tail, the bones of either carpus and those of the tarsi, and the

epiphyses of the long bones, etc. The marsupial bones were well

formed in both sexes, and upon either side were nearly as long as

the corresponding ilium, and about one-fourth the size in bulk.

According to Flower, the number of vertebre in the spine of

the Virginia Opossum was 7 cervical, 13 thoracic, 6 lumbar, and

26 caudal. This was probably correct for the adult animal of

this species, while in subadults, of an age here considered, the last

three caudals were not developed, and the three or four anterior

to them were in the most rudimentary condition possible.

In both these specimens the dentition in use was as follows :—

a3, cp pm + m 2 = 42,

the third of the cheek-teeth being the molar-like predecessor of

the one premolar which changes in Marsupials. Two further

molars would have come up in later life, making the adult

formula :—

- 5 1 3 4

tp Cp pms, m4= 90.

The premolars were triangular, sharp-pointed, and flattened from

side to side; the molars had numerous sharp cusps and the canines

were large and curved.

Mr. D. Seru-Suire, F.Z.S., Curator of Birds, exhibited, by

means of lantern-slides, photographs of the male Peacock Pheasant

(Polyplectron chinquis) displaying to the female.

The typical display, as depicted in the photographs, resembled

very closely that of the Argus Pheasant, the bird facing the

female while he lowered the breast to the ground and expanded

the wings and tail like a shield, the head being held sideways

against one wing.

Mr. Seth-Smith also exhibited photographs (Pl. LXIX. *) of

the young Cariama cristata hatched and reared in the Gardens in

1911, and remarked that although young of this species had been

hatched in the Menagerie on previous occasions, he believed that

this was the first occasion on which the young had been reared

to maturity.

The nest was formed of sticks and small twigs on a platform of.

branches that had been specially erected at about eight feet from

the ground in the Eastern Aviary. Of the two eggs laid, one

was accidentally broken by the birds, but the other hatched on

June 6th, incubation, performed chiefly by the female, having

occupied twenty-nine or thirty days. The young bird was

covered with down of a pale brown colour mottled with darker

brown on the back, that on the head being extraordinarily long

and hairlike. It was fed by the parents with small pieces of meat,

* For explanation of the Plate see p. 558.

558 MR. R. I. POCOCK ON A

mice, hard-boiled egg, and cockroaches. It left the nest at a month

old, and at five weeks old may be described as follows :—

Upper parts pale brown, darker and with a rufous shade on the

mantle, the rump and upper tail-coverts being dark brown. The

crown of the head and neck barred with dark brown; the wings

also covered with fine wavy lines of dark brown on a paler ground.

The abdomen and under tail-coverts whitish.

The bill very dark brown, the legs and feet dark grey, and the

iris very pale silvery grey.

EXPLANATION OF PLATE LXIX.

Fig. 1. Young Cariama cristata 24 days old, in nest.

Fig. 2. ae BS a 30 days old, with parent.

Figs. 3 & 4.,, Ko 58 2 months old.

PAPERS.

30. On a rare Stag (Cervus walliehii) from Nepal recently

presented to the Zoological Society by His Majesty

King George. By R. I. Pocock, F.R.S., F.LS.,

F.Z.8., Superintendent of the Gardens.

[Received March 19, 1912: Read April 2, 1912.]

(Text-figures 66-71.)

INDEX.

Systematic : Page

Cervus wallichii Cuv., Western Tibet, text-figs.

GGEG9) pacbacepelt 28s Baek 4. co castahe Seep eoDO

» hanglu Wagn., syn. cashmeriensis Adams,

cashmeerianus Falc., Kashmir, text-fig.70 566

» affinis Hodgs., Choombi Valley, Lhasa, ete. 567

» macneilli Lydd., Szechuen, text-fig. 71 ... 570

» kansuensis, nov., Kansu .........-.....0000000) 502

Characters of these species tabulated .................. 574

Cervus albirostris, Tibet ............c0cccccceeeeeeneee eee 574

From the scientific standpoint the most interesting animal

in the collection from Nepal presented by H.M. the King to the

Zoological Society was a large Stag belonging to a species

(Cervus wallichii) which has never been previously exported

alive from India, and, so far as available records show, has

never been shot or preserved by any English sportsman, and is

therefore unrepresented in any of the large museums of the

world. The arrival of this specimen, moreover, has made it

possible to classify a species which for nearly a century has

been a puzzle to all systematic workers on the Deer.

RARE STAG FROM NEPAL. 559

Cervus wallichit was described by G. Cuvier (Rech. Oss. Foss.

ed. 3, iv. p. 504, 1825*; ed. 4, vi. pp. 88-89, 1835) from a

native sketch, sent to him by Duvaucel, of an animal, one of a

pair, according to Blyth, living in the menagerie at Barrackpore

and said to have come from Muktinath, north of Dwalagiri in

Nepal. This sketch was reproduced by F. Cuvier & Geoffroy

St. Hilaire (Hist. Nat. Mamm. iv. no. 356, 1823), and modifications

of it were reproduced subsequently by Jardinet and Hamilton

Smith §.

Text-fig. 66.

Cervus wallichii.

Photograph of Cuvier’s figure of the type-specimen.

The main characters of the animal are made clear by the

descriptions and figures of the French authors. It was described

as dark grey-brown or yellowish grey-brown with pale legs, a

* T have not seen this edition.—R. I. P.

+ In support of this Blanford (Mamm. Brit. India, p. 538), copying Blyth, cites

Hardwicke (Tr. Linn. Soc., Zool. xiv. p. 581, 1823). Hardwicke, however, does not

there state that the stag from Muktinath he identified as ‘ C. pygargqus, Pall.’ was

the type from which the sketch was taken. The coincidence of the dates, however

leaves little doubt on this point.

{ Naturalistst’s Library, iii. pl. 10, p. 161, 18365.

§ Griffith’s Animal Kingdom, iv. p. 103, pl., 1827.

560 MR. R. I. POCOCK ON A

very short white tail and a large white disk upon the croup*.

The figure shows no less clearly that the caudal disk in size and

extension over the croup, reaching nearly to its summit, resembled

that of a Wapiti but was white instead of yellowish buff in

colour. The antlers were short, only a little exceeding the head

in length, and carried a brow and a bez tine, the beam being

simply forked and the trez tine absent. The brow and bez tines

were close together at the base and diverged at about an angle of

45°, the brow tine projecting straight forwards and the bez in-

clining obliquely upwards. The beam, which receded at an obtuse

angle of about 135° from the bez, appears from the figure to

have formed a fairly regular arch, there being no indication that

either of the terminal tines was turned inwards.

For reasons given below I am: quite of Hamilton Smith’s

opinion (Griffith’s An. King. iv. p. 103, 1827) that these antlers

show the stag to have been an old animal at the time the

illustration was made.

A pair of the antlers of this species was figured and described

by Blyth (J. A. S. Bengal, x. fig. 7 of pl. facing p. 750, 1841)

and subsequently stated by this author in the same journal

(vol. xxx. p. 188, 1861) to have belonged to the animal of which

a sketch was sent to Cuvier. They were judged by Blyth

to have been produced by a stag in its third year. It will be

noticed, however, that neither antler of the pair has a trez tine ;

and since in the Red Deer this tine normally, at all events,

appears in the antlers of the third year, one would expect it to

be present in a specimen of C. wallichit of that age. It will

also be seen that in the left antler figured by Blyth, the beam is

straight and terminates in a single spike, while in the right

antler the beam is forked and much more curved. The want

of symmetry between the two in these and other particulars,

coupled with the absence of the tines mentioned, forcibly

suggests to me degeneration with age. It is, at all events, quite

clear that these antlers were not those carried by the stag at the

time the figure sent to Cuvier by Duvaucel was drawn. They

were probably a later growth and still more decadent. Even the

right antler was less curved than were those of the type, as figured ;

and although the basal juxtaposition and mutual divergence of

the brow and bez tines were much the same in the two cases, the

angle of their divergence from the beam was in both instances

much smaller. But, although these differences and resemblances

between the two sets of antlers are interesting, I do not think

they afford much clue to the aflinity of C. wallichii, since these

antlers were, in my opinion, obviously degenerate.

The specimen the Society has just received is a comparatively

large Stag, though not approaching a Wapiti in size. It stands

about 4 ft. 3 ins. (just under 13 hands) at the shoulders. It

has a long face, a small sleepy looking eye, and its longish ears

* The italics are mine.—R. I. P

RARE STAG FROM NEPAL. 561

differ from those of all living Deer that I have seen in having

the upper edge markedly sinuous (emarginate) towards the tip.

The hairs on the neck and throat are longer than elsewhere

and constitute a small mane. The tail is quite short, the legs

strong, the hoofs rather broad and the metatarsal tuft a little

below the hock. The colour (in March) is remarkably pale, being

a tolerably uniform yellowish or sandy-brown all over, except on

Text-fig. 67.

Cervus wallichii.

The King’s specimen shortly after its arrival at the Gardens in March.

the forehead where the matted hair is browner, on the cheeks

and the backs of the ears which are greyer, and on the muzzle

and chin which are pale fawn with an indistinct darker patch at

the corner of the mouth. The backs of the thighs are white,

the pale portion of the rump being only as wide as in the

Hangul or Kashmir Stag, but from the level of the ischia the

white bends sharply forwards and extends nearly as far as

562 MR. R. I. POCOCK ON A

the summit of the croup, forming a very large pure white disk

without trace of median dividing dark line. Upon the croup

this disk is not emphasized by a bordering of darker hairs; but

below the level of the tail, the white on the back of the thighs

is set off by a margin of brown. This disk, so far as size is

concerned, differs from that of the Wapiti in being narrower

below, its width on each side on a level with the tip of the tail

only equalling the width of that organ, whereas in the Wapiti

the disk at this point is equal to twice the width of the tail.

It is also noticeable that the hairs adjoining the croup-disk are

quite white at the base, so that if they were elevated or deprived

of the yellowish-brown pigment of their distal portions, a far

larger white rump patch would be displayed than is normally the

ease. None of the hairs show signs of speckling apically or

elsewhere, and their concealed portion is greyish brown on the

body, though darker on the neck. Both on the body and over

the croup a very large number of them are curled forwards

apically, like those on the croup of the type specimen of C.

wallichit as described by Cuvier, but a mounted specimen of the

Kashmir Stag in the British Museum shows, to a lesser degree,

this same peculiarity, which is no doubt a sign of old hair.

The antlers, which were shed early in March on the voyage

from India, are short but massive. The right one is abnormal,

the left probably normal but, possibly, reduced in size from age

degeneracy. Both are brown with pale tips to the tines. In the

left antler the burr is short, the brow tine rising about an inch

above it. This tine is about 84 inches long, projects nearly

horizontally forwards and is bent slightly down just before the

tip. The bez tine is straight, about half an inch shorter than the

brow and diverges from it at an angle of about 45°, their basal

separation being about 1 inch. Just above the root of the bez

on the inner side there is a short snag. Beyond the bez the

beam curves gently outwards and upwards for about 74 inches to

the origin of the trez, which is about 7 inches long and lightly

curved, and lies in approximately the same plane as the bez and

brow tines. Above the trez the beam ascends, showing upon the

anterior aspect a slight inward inclination, but from the external

profile view it exhibits a decided inclination backwards with a

light upward curve, its axis lying almost at an angle of 45° to

that of the trez, and forming a very obtuse angle with the lower

part of the beam, there being a marked concavity behind the

root of the trez. It terminates above in two tines, one short and

erect and continuing the line of the beam, the other twice as

long and inclined upwards and inwards.

The antler is thus five-pointed. It resembles the antlers of

the typical specimen in the basal juxtaposition and marked

divergence of the brow and bez tines, but, apart from the

presence of the trez tine, differs from them in the more upward

curvature of the basal part of the beam, which is thus inclined at

sharper angles to the axis of the brow and bez tines. In this

RARE STAG FROM NEPAL. 563

latter particular, however, the antlers figured by Blyth are inter-

mediate between those of the example figured by Cuvier and the

one just described.

Text-fig. 68.

Cervus wallichii.

A. Anterior aspect of the shed antlers.

B. External aspect of left antler.

In the right antler the brow and bez tines resemble those of

the left antler; but above the bez the beam bifurcates into

564 MR. R. I. POCOCK ON A

anterior and posterior branches ; the former, occupying almost the

position of the trez in the normal antler, is stout, 15 inches long,

evenly curved upwards and outwards and two-tined, one tine

being short and forwardly directed, the other, which is terminal,

being long and lightly curved upwards, backwards and inwards.

This branch may be described as an attempt at a reduplication

of a normal beam. The posterior branch is thinner than the

anterior and ascends with an even curvature upwards, outwards

and then inwards, to terminate in two tines, an external which

is merely a short bud, and an internal which follows the line

of the beam without showing any marked inward inclination,

although it clearly corresponds to the inwardly inclined terminal

tine of the left antler.

Judging from their small size and asymmetry, these antlers

may be decadent; but, if so, the left antler has suffered less

than the right from that process and less than those of the

type specimen of C. wallichii described above. Still I do not

think it safe to infer that this left antler resembles that of a stag

of this species in full vigour. The possibility, however, must be

borne in mind.

Of the exact geographical area inhabited by C. wallichii we

have at present no trustworthy information. The type of the

species in the Barrackpore menagerie was said by Hardwicke

(Tr. Linn. Soc. xiv. p. 581, 1823) to have been brought from

Muktinath, away to the north of Dwalagiri in Central Nepal.

The example now in the Gardens was presumably in captivity in

Nepal, since it was presented to King George by the Maharajah

of that country (see note below).

The only other Stag, known to me, which seems to belong

to this species is the one which Mr. Lydekker identified from a

photograph as C. affiiis (P. Z.S. 1909, p. 599, fig. 182). This,

too, was a captive animal and was alleged to have come from

Sikhim. I judge from the size of the rump patch that this stag

belongs to C. wallichit. Myr. Lydekker described the colour

as “very like that of the Hangul with a large white rump-

patch.” This seems to be correct so far as the size of the

rump-patch is concerned; but if the colour of the body was

inferred from the photograph, the inference is untrustworthy

since the camera is notoriously deceptive in accurately indicating

the differences between dark and light tones * (text-fig. 69).

* | Note added June 1st, 1912.|—While this paper was in the printers’ hands

Mr. Lydekker wrote to the ‘ Field’ (May 11th, 1912), on behalf of Col. J. Manners-

Smith, to say that the Stag represented by the photograph above discussed is the

same individual as the one presented to the King by the Maharajah of Nepal. The

history of this animal was recorded by Col. Manners-Smith in the ‘ Field, July 31st,

1909, p. 289. It came from the upper reaches of the Sanpo Valley, close to Lake

Mansarowar, where it was captured as a fawn, and was in its second year in 1909.

It is therefore in its fifth year at the preseut time, and is not, as I supposed, an old

animal. That the colour was the same three years ago as now may be inferred from

Col. Manners-Smith’s reference to it as “very light.” As regards the distribution

of the species, reports stated that the deer was plentiful in various places in Western .

Tibet, near the source of the Brahmaputra River, the hills north of Mount Kailas,

RARE STAG FROM NEPAL. 565

Since then the only known specimens belonging, or presumably

belonging, to C. wallichit weve menagerie animals, no confidence

can be placed in their alleged localities.

Text-fig. 69.

Cervus wallichii.

The photograph, forwarded to Mr, Lydekker, of the specimen shown in text-fig. 67

(p. 561).

For a variety of reasons, into which it is needless to enter,

C. wallichit has hitherto found no abiding place in the chronicles

of the Cervide. Some authors have added its name to the

synonymy of the Kashmir Stag Cervus hanglu (= cashmeriensis),

others have supposed the Stag to be identical with Cervus affinis ;

others have left it as indeterminable. ‘To settle, as far as may

be, its status and affinities it is necessary to examine in some

detail the characters of the two above mentioned species with

which it has been confounded, and of certain other Stags

belonging to the same group of the genus Cervus.

and other less well-known localities being mentioned; the available evidence indi-

cating, in Col. Manners-Smith’s opinion, that the species does not cross the southern

watershed of the Brahmaputra and is not found in Nepalese territory. He suggested,

therefore, that the original specimen of C. wallichii had been brought to Muktinath

from somewhere in Western Tibet round about Lake Mansarowar. It may be

added that Col. Manners-Smith, apparently unaware of the discrepancies between

the descriptions of C. walliehii and OC. affinis, believed these two forms to be

identical.

566 MR. R. I. POCOCK ON A

Cervus hanglu*, better but wrongly known as cashmeerianus, is

a not uncommon Stag. Moderately good figures of it have been

published by Dr. P. Ll. Sclater (Tr. Zool. Soe. vii. pl. 30, 1870),

and by Mr. Lydekker (Deer of all Lands, pl. iv.), while there are

several photographs in existence of specimens that have been

exhibited in the Gardens or kept at Woburn. One recently

Text-fig. 70.

Cervus hanglu.

Specimen, living in the Gardens in March, to show the resemblance of the

caudal disk to that of C. affinis, as described by Hodgson.

published in the ‘ Field, Dec. 30, 1911, shows very clearly the

shape and extent of the caudal disk characteristic of the species.

The white on the inner side of the back of the thighs extends a

short distance upwards above the root of the short tail, but the

* Waener in Schreber’s Saug. iv. p. 352 (note), 1843. Various modifications of

the word cashmeerianus are also in use; but of these cashmeriensis appears to be the

oldest, since it was first introduced by Leith Adams in connection with a description

of this species (P. Z.S. 1858, p. 529).

RARE STAG FROM NEPAL. 567

yellowish or dirty-white right and left moieties of this area do

not meet in front of the base of that organ but are separated by

a median brown band, continuous with the colour of the back ;

‘this band may or may not run along the middle of the tail

itself. Thus the caudal disk is small, smaller in fact than in

most Deer of the Elaphine group. Nevertheless it is very

conspicuous by reason of the blackish setting formed by the

adjacent hairs of the hind quarters. So far as my experience of

two specimens goes, this caudal disk does not vary appreciably

with the season, but remains fairly constant in extent, shape and

distinctness. Nor does it alter with age, judging from the fact

that in a very old animal with worn teeth, which was recently

shot in the Gardens for senile decay, the disk was exactly like

that of a younger animal in perfect condition now on exhibition.

No doubt, however, it exhibits a certain amount of individual

variation. It will thus be seen that C. hanglu may be at once

distinguished from C. wallichii by its totally dissimilar caudal

disk *.

It is also darker in colour, being very decidedly brown but

fading to a rather paler hue on the flanks, which are lighter than

the legs, the latter being as brown as the back both externally

and internally. The coat, moreover, is markedly speckled owing

to the presence of a subapical pale annulus on the individual

hairs.

Another very noticeable distinction between all the specimens

of O. hanglu and the one example of C. wallichii that 1 have seen

ig in the colour of the hairs about the mouth. The chin and

lower lip of C. hanglu are white and the upper lip up to the

nostrils is cream or dirty-white, so that the muzzle is rather

sharply contrasted in colour with the browner hue of the rest of

the face. Light rings round the eyes, too, are very marked.

They are not so marked in C. wallichw and, as has been stated,

the muzzle and chin are decidedly fawn-coloured and darker than

the rest of the face at least in the winter coat.

Cervus affinis Hodgson was based upon the skull and antlers

of a Stag wrongly alleged to have come from the Sal forests in

Nepal (J. A. Sg Bengal, x. pt. 2, pp. 721-724, pl., 1841). The

brow tine was long, projected forwards over the face and was

straight or had a sharply upturned tip. The bez was subequal

and subparallel to it and also had an upturned tip, the distance

between their points of origin being 23 inches. Above the bez

the beam reclined backwards and outwards for a short distance,

then bent sharply upwards below the origin of the trez and ended

in a pair of subequal terminal tines. This sharp upward bend of

* Blyth, who believed the Kashmir Stag and C. wallichit to be specifically the

same, explained away this difference by assuming that the caudal disk in the figure

of the type of C. wallichit was exaggerated (J. A. S. Bengal, xxx. p. 188, 1861).

568 MR. R. I. POCOCK ON A

the beam, obvious both in profile and front view, is absent in the

known antlers of C. wallichii.

In the J. A. S. Bengal, xix. p. 466, pl., 1851, Hodgson figured

and described another pair of antlers sent to him by Dr. Campbell

from Ding-cham, north of Sikhim. These he referred to

C. affinis; but they only serve to illustrate the variability to

which the antlers are liable.

Subsequently (J. A. S. Bengal, xx. pp. 388-394, pl. vii., 1851)

Hodgson redescribed the species ‘from abundant supplies of the

spoils [also sent by Dr. Campbell] ..... exhibiting both sexes

in various states of maturity ..... the skulls and leg-bones

being attached to the majority of the specimens.” He thus had

at his disposal what he described as “ unusually copious and

adequate material.” This is an important point to remember.

The description of the antlers given in this paper agrees

tolerably closely with that which he published in 1841. Certain

individual variations are pointed out, and it is stated that the

basal interval between the brow and bez tines varies from two to

over four inches, two being the usual distance.

After remarking that the specimens were in winter coat,

having been killed in February, Hodgson described the colour as

“earthy brown, more or less lutescent, the head and neck being

concolorous with the back; but the flanks are conspicuously

paled, and the belly as conspicuously darkened... .. the neck,

though paler below than above, is not very noticeably so. But

the paling of the flanks is as decidedly so as the nigrescence of

the belly; the white and black forming a conspicuous contrast

on these parts..... The limbs are paler than the back, darker

than the flanks and they have an earthy-brown list down their

external and anterior aspect.”

Finally he said, “The tail is very short, and the caudal dise

remarkably small but conspicuous from strong contrast of colours.”

In another place he also spoke of “the small caudal disc,” adding

“ The little tail is white, like tts disc, a dark mesial line dividing

the latter along the culmenal (sic) line of the tail.” *

This description of the small caudal disk with its median

dividing dark line extending on to the tail applies tolerably

accurately to the caudal disk of C. hanglw; and if Hodgson’s

published figure, bad though it be, of C. afinis be compared with

those of the former species published by Mr. Lydekker and

Dr. P. L. Sclater, it will be seen that the principal differences

between the two species lie in the form of the antlers and the

rather larger caudal disk with narrower median line in C. affinis.

Taking Hodgson’s description as a whole, however, it amply

justifies and explains Blanford’s statement (Mamm. Brit. India,

p- 537) that the coloration of the two species is the same, and

equally discredits Mr. Lydekker’s assertion that “as regards

* The italics in this paragraph are mine,—R. I, P,

RARE STAG FROM NEPAL. 569

coloration [of afinis] accurate information is wanting” (Game

Animals of India, p. 217, 1907).

The precise nature of the relationship, however, between the

two species is by no means so easily and certainly settled as the

literature seems to indicate. ,

In the British Museum there are three skins and a mounted

head referred to C. affinis. The head and one of the skins came

from Hodgson, the former being labelled “ North of Bhotan,” the

latter ‘“‘ North India” and bearing the date 1857. It is not cer-

tain, so far as I know, that these specimens are part of the material

that author described in 1851. Very possibly they are. If so, the

head is very much faded—which is likely enough considering that

it is exhibited in the gallery—because it is now a tolerably uniform

sandy fawn and not “earthy brown.” It is noticeable, however,

that the lips and chin are fawn-coloured as in the example of

C. wallichii in the Gardens, and not white as in C. hanglu. The

skin, on the contrary, which has not been exhibited, agrees fairly

well with Hodgson’s description, being for the most part dirty

brown and much darker both on the body and legs than is our

specimen of C. wallichii. The hair, moreover, is very coarse and is

of winter growth. The tail is cut away; but the white disk is not

nearly so distinctly divided mesially as Hodgson’s description would

lead us to believe, nor as is the case in C’. hanglu. Remnants of

a brown dividing line are, however, traceable. Again, although

the disk is smaller than in C. wallichii and is set off by a margin

of darker hairs, the bases of these adjacent dark hairs are white,

and the disk could be made to approach that of C. wallichii in

size if the brown tips of these hairs were removed or deprived of

pigment. But be it remembered, the bases of the hairs adjoining

the croup-disk in our example of C. wallichii are also white, so

that the disk of the latter is actually, with the hair undisturbed,

as large as, or larger than, the disk in Hodgson’s affinis with the

hairs interfered with in the way supposed. In both specimens

the hairs are short and crisp.

The other two skins named C. afiinis in the British Museum

belonged to the late Dr. Blanford and are labelled “Sikkim,

L. Mandelli”*. The coat is in a better state of preservation

than in Hodgson’s specimen above described. The general

colour is greyish brown, the hairs being pale basally, brownish

distally with a distinct subapical pale annulus, and a darker tip,

imparting a marked speckled appearance to the coat. The rump-

patch is small and white, but as in Hodgson’s specimen, and in

Hodgson’s published figure of C. affinis, it spreads on to the

posterior part of the croup in front of the base of the tail. In

one of the specimens it is very clearly defined all round by a

bordering of brown hairs unspeckled and considerably darker than

the back, but with white bases. In the other the hairs bordering

* Dr. Blanford once informed me that Mandelli’s localities are untrustworthy.

Proc. Zoou. Soc.—1912, No, XX XVIII, 38

570 MR. R. I. POCOCK ON A

the patch are only a little darker than the back and are speckled

to the edge of the disk. As in Hodgson’s skin the tails have

been cut away, and the croup-disk has a median abbreviated

irregular dark line, much more clearly defined in one skin than in

the other. This disk is of about the same size as in Hodgson’s

specimen, although the hairs of the coat generally are longer and

less close and crisped. The legs are paler, being fawn-brown down

the front and on the fetlocks, and the lips and chin are also fawn,

and not cream and white as in C. hanglu.

Reverting once more to the mounted head in the British

Museum, it may be added that the left ear shows signs of the

emargination of its upper border so evident in the living example

of CO. wallichii. The right ear nevertheless has a straighter upper

rim. I suspect, however, that C. affinis has the ears shaped as

in C. wallichia.

Putting all these facts together, it appears to me that C. affinis

is nearly intermediate in coloration between C. wallichit and

O. hanglu, especially in the general tint of the body and limbs

and the size and division of the caudal disk, the most marked

character in which it resembles C. wallichii and differs from

C. hanglu being the fawn colour of the lips and chin.

There is evidence also that both C. wallichii and C. affinis are

larger than C. hanglu and have longer faces*; but judging by

the standard of specific and subspecifie differences usually adopted

in the Cervide, it appears to me to be doubtful whether more

than subspecific importance should be granted to the differences

above described between C. wallichii, C. affinis, and C, hanglu.

Tt must be remembered, however, that wallichii is the oldest

name of the three.

Exact partieulars of the range of C’. affinis are much wanted.

Only two districts are mentioned by Mr. Lydekker in the table

of horn-measurements in Rowland Ward’s ‘ Records of Big

Game,’ 1910 (p. 38), namely, the Tibetan Frontier and the

Choombi Valley; but the valleys of Bhotan near the Choombi

are added under the diagnosis of the species. The Stag has

also been recorded by Col. H. A. Iggulden from the Tsan-po

basin, near Lhasa (‘ Field, Oct. 1906, p. 736); but whether the

specimens were accurately determined or not, it is impossible,

without the evidence of skins, to say.

Another Stag belonging to this same group is the animal from

Szechuen described by Mr. Lydekker as C. cashmirianus macneilli

(P. Z.S8. May 11, 1909, p. 588, pl. Ixix.). The coat is finely

speckled all over owing to the apical annulation of the hairs, as

in Mandelli’s skins of C. affinis and as in fresh-coated examples of

C. hanglw; the prevailing colour, however, is strikingly grey,

especially on the sides and legs, but the back is rather darker,

* This difference is very noticeable between the examples of C. hanglu and

C. wallichii living side by side in the Gardens.

RARE STAG FROM NEPAL. 571

that is to say more fawn, than the sides, and the top of the head,

the face, and the nape of the neck are browner than the back.

There is no mane and the neck shows signs of coat change. The

legs are less distinctly speckled than the sides of the body and

are darker in front than behind. The belly is whitish, as in the

females of C. affinis described by Hodgson. Apart from the

prevailing greyness of the pelage, the most interesting features

about this Stag are the brownish fawn colour of the chin and lips

Text-fig. 71.

Cervus macneilli.

Photograph of the plate of the type-specimen (P. Z.S. 1909, pl. Ixix.).

and the coloration of the rump. The upper side of the tail is

black with a narrow edging of white. On each side of the tail

there is a narrow white area, which however barely surpasses

the root of that organ dorsally. This white area is bordered with

black, and there is a blackish-brown unspeckled croup-disk almost

as extensive as the white croup-disk of C’. wallichii.

From the distribution of this Stag, its affinities might be

inferred to be rather with C. affinis than with C. hanglu ; and in

38*

572 MR, R. I. POCOCK ON A

spite of the lesser extent of the white on the rump, a feature in

which it more resembles C. hanglu, this conclusion is borne out

by the coloration of the chin and lips and by the shape of the

ears, which are pointed and appear to have had a sinuous upper

edge.

a as Mr. Lydekker records, Capt. McNeill declared

the Stags of this kind that he saw in Szechuen to be nearly as

large as Wapiti. Even making allowance for exaggeration in

this estimate, the animals must have been of considerable size.

CO. hanglu, however, is not a large Stag, although strongly made

and ‘“ cobby” in build.

Unfortunately, there is no record of the date when the type

specimen of this Stag, a young hind standing, as mounted, forty

inches, was shot.

Another type of this group of Stags is represented by a skin and

skull kindly lent to me for identification by Mr. W. F. H. Rosen-

berg, F.Z.S. The animal, an adult hind, was shot by Dr. J. A.

C. Smith on Mareh 23rd, 1911, 30 miles S.E. of Tao-chou, Kansu,

in China, at an altitude of 11,000 feet.

The colour is a tolerably uniform earthy brown relieved by fine

close-set speckling due to a subapical pale annulus on each hair.

On the sides the main shaft of the hairs is greyer and less brown

than dorsally, and low down towards the belly the subapical

annulus is longer, so that the general tint is markedly paler.

The belly is white, but not the chest. On the neck the hairs are

longer with longer apical annulus, the shaft of the hairs being

browner along the nape than on the sides of the neck, so that

there is an ill-defined dorsal neck-stripe as in C. macneilli ;

the front of the neck (throat) is paler than the sides ; the legs are

fawn-brown down the front and sides, paler behind. The forehead

is brown, the face grey-brown and closely speckled ; the lips and

chin are fawn-brown, unspeckled and without white, and the

black patch below the corner of the mouth is well marked. There

is a blackish-brown unspeckled croup-disk, as in C. macneilli,

and the hairs of this disk become gradually more and more white

towards the root of the tail*. The white on the buttocks is of

the same extent approximately as in C. macneilli, but the tail

itself is much whiter than in that animal, since it merely has a

narrow median dark stripe as in most examples at all events of

C. hanglu. The ears are long and pointed, with apparently a

sinuous upper edge such as is seen in CO. wallichit.

The following measurements in the flesh, taken by Dr. Smith,

may be of future use, although at the present time they indicate

nothing, because there are no corresponding measurements of

other deer wherewith to compare them :—

Head and body 1745 mm. (a little under 6 feet ; tail 145 mm.

* This is probably also the case in C. macneitlli, but I was unable to touch the

mounted specimen of the animal in the British Museum,

RARE STAG FROM NEPAL. 573

(under 6 inches); foot (? hock to heel) 525 mm. (21 inches) ; ear

225 mm. (9 inches). ‘The skin is too shrunk to make an estimate

of the animal’s height of any value.

The skull and teeth show that the specimen was a full-grown,

but not old, hind. Its basal length from the notch between the

condyles to the tip of the premaxille is 13? inches (about 344 mm.);

and the length from the anterior edge of the orbit to the tip of

the premaxilla about 9 inches (= 225 mm.), and the width across

the orbits 67 inches (=156 mm.).

On comparing these dimensions with those recorded by Blanford

for C. hanglu and C. affinis, it would appear that this Kansu

Stag is somewhere about the size of the former. For example,

he quotes the length of the Hangul (probably of males) as from

7 to 74 feet and the length of a skull of this sex as just

over 15 inches. Probably the basal length of the latter would

be about 142 inches.

The smaller of two skulls of C. affinis had a basal length of 163

inches and a width across the orbits of about 74 inches. Thus,

allowing for the fact that the Kansu specimen is an adult female,

it may be inferred that the males are about as large as those of

C. hanglu and considerably smaller than those of C. affinis.

The description given above of the colour of this Stag agrees in

a general way tolerably closely with that of the examples of

C. affmis that Hodgson had in his hands, and it is important to

remember the close correspondence in date, Hodgson’s specimens

having been killed in February and the Kansu specimen in March ;

and since deer of the Elaphine group moult in April and May,

it is evident that the examples in question were still carrying

their winter coats. In their generally dark earthy brown color-

ation, both differ markedly from the living example of C. wallichi

in March.

_ The coloration of the Kansu specimen, however, is not identical

with that of C. affinis. I do not think it safe on the evidence of

one skin to trust much to the greater uniformity of the colour of

the body as shown by the absence of a distinct darkening of the

back and “lutescence” of the sides to which Hodgson refers in

C. affinis, but the large size of the dark croup-patch and the

smaller extent of white at the base of the tail are probably more

dependable. It is in both these particulars especially that the

Kansu stag resembles the Szechuen stag C. macneilli. Since,

however, it differs from the latter in general coloration and in the

greater amount of white in the tail; from C. affinis in having no

white above the root of the tail and a larger dark area on the

croup; from C. wallichit in having no white on the croup at all,

except such as is concealed by the overlying black ends to the hairs ;

and from C. hanglu in the dark colour of the chin and upper lip,

the Kansu stag seems to deserve a name; and I propose to call

it C. KANSUENSIS.

574 MR. R. I. POCOCK ON A

The following analytical key to the species above discussed

represents my views as to their probable affinities :—

a. Chin and lower lip white, muzzle pale fawn, markedly lighter

than the rest of the face; ears bluntly pointed and with

straight upper rim; no white on croup above root of tail... hanglu Wagn.

a’. Chin fawn or brown, muzzle brownish fawn with at most

a little white on the lip in front; ears long and pointed

with sinuous upper edge (? in affinis).

b. White on back of thighs not spreading on to croup

above root of tail; tail dark down centre; a large

blackish-brown croup-patch reaching nearly or quite to

the summit of the croup.

c. Prevailing colour grey, rather darker on the back and

still darker on the head; practically the whole of the

uppersidelon thertallvblacksry. se, ceccsncoven sce cccnceess es. macneilli Lydd.

ce’. Prevailing colour (March) brown; upper side of tail

with an irregular median dark stripe..................-..... kansuensis, nov.

b'. White on back of thighs spreading upwards above the root

of the tail and encroaching more or less on the croup;

dark patch on croup smaller or absent.

d. White area above the tail comparatively small with a

more or less distinct median dark longitudinal stripe;

prevailing colour (February) earthy brown, paler

laterally sie cH ESAS, radars ee ese

d’. White area above tail very large, reaching summit of

croup and undivided by dark line; prevailing colour

(March) pale fawn-browN............... 2.200. sseeeseeseeneee es wallichit Cuv.

affinis Hodgs.

In selecting the colour of the chin and muzzle as the character

for eliminating C. hanglu from the rest of the species, I am not

unmindful of the fact that Cuvier wrote of C. wallichi ‘comme

Vordinaire, le tour de l’ceil, celui de la bouche, sont plus pales, et

il y a du blanc sous la machoire et une tache noire sous langle

des lévres.” These words may suggest that the muzzle and

chin were coloured as in C. hanglw; but the coloured plate of

C. wallichii, from which the description was taken, does not bear

out this supposition nor does it quite justify Cuvier’s phraseology,

for the lips and chin are washed with the same yellowish tint as

that of the rest of the body.

If the caudal disk were taken as the primary basis for the

classification of the species, they would be grouped as follows :—

C. macneillit+ C. kansuensis; C. hanglu+C. affinis; C. wallichii.

I prefer on the whole to regard these forms as species rather than

subspecies because we have at present no proof that they inter-

grade, and the differences between them are perhaps greater than

those between the various races of Wapiti (Cervus canadensis)

and Red Deer (Cervus elaphus). As a group they resemble the

Wapiti and differ from the Red Deer in the shortness of their

tails.

The only other Stag which should perhaps come into the same

category, judging from the shortness of the tail, the size of the

caudal disk, and the shape of the ears, is the Tibetan species

commonly known as (C. albirostris which Blanford described as

C. thoroldi (P. Z. 8. 1898, p. 444). Judging from the mounted

specimen of this stag in the British Museum the coloured figures

of this species published by Blanford and by Mr. Lydekker

RARE STAG FROM NEPAL. 575

(‘Deer of all Lands,’ pl. v.) have the yellow caudal disk too

large and extending too high up the croup, and the ears, which

are really very long and pointed, with the upper edge sinuous,

represented as much too short and blunt. In the whiteness of

the chin and muzzle this species goes a stage beyond C. hanglu ;

but of course the stag differs from those discussedin this paper

by the reversal of the hairs along the spine between the croup

and withers and, so far as is known, by the absence of the bez

tine.

In the above given table no use has been made of the antlers

as distinguishing features. These vary so much with age and

from a variety of unknown causes that I am convinced too

much confidence is placed in them in most systematic treatises

on Deer*. At the same time it may be useful to bear in mind

that the recorded antlers of C. wallichii appear to differ from

average antlers of C. hanglu and C. affinis in the basal juxta-

position and marked distal divergence between the brow and bez

tines, and at least from those of C. affinis in the comparative

straightness of the beam which lacks the sharp upward curvature

of its distal half. It will be highly interesting to see if the

example of C. wallichit now living in the Gardens grows, under

the changed conditions of its existence, antlers resembling those

that it shed on its journey from India. This and the colour of

its summer coat I hope to have the chance of recording later on.

As regards the stature of the Deer described above, Blanford

gave the height of C. hanglu as ranging from 48 to 52 inches.

The example of this species now in the Gardens is barely 48 inches.

C. affinis, according to Hodgson, stands from 54 to 60 inches

and thus rivals a Wapiti in stature. A small Canadian stag

Wapiti now in the Gardens is about 56 inches, and an example

of C. xanthopygus 58 inches. But since Hodgson’s measurements

of C. affinis were taken from dried skins, his estimate must have

been largely guess-work and probably erred on the side of

exaggeration than otherwise. Blanford, or the authorities from

whom his information was derived, measured in all probability

freshly killed specimens of C. hanglu; and since dead animals

give higher stature-measurements than living ones, exaggeration

in connection with this species must also be allowed for.

Our living example of C. wallichii is as nearly as possible

51 inches, and is manifestly much smaller than any stag of the

Wapiti group, whether American or Asiatic, that I have seen.

It is, however, exactly the height assigned by Hardwicke to

the typical example of this species that was exhibited in the

menagerie at Barrackpore.

* T can see no reason for adopting Mr. Lydekker’s suggestion (‘ Field,’ May 11th,

1912) that the Stag he described from a pair of shed antlers picked up in Szechuen

as Cervus canadensis wardi (P.Z.S. 1910, pp. 987-989) is identical with Cervus

wallichit.

576 DR. F. E. BEDDARD ON

31. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea. By Frank BH. Bepparp, M.A.,

D.Sce., F.R.S., F.Z.8., Prosector to the Society.

[Received February 6, 1912: Read April 2, 1912.]

(Text-figures 72-83.)

TV. On A Species or JWERMICAPSIFER FROM THE HyRAX, AND

ON THE GENERA ZSCHOKKEELLA, T'HYSANOTANIA, AND

HVRACOTENIA,

INDEX.

Systematic ; Page

Inermicapsifer capensis, sp. n. ................4. 598

Ayracotania, gen. 0. 0.0.0... cece cee cee eee eee eee. 604

ET. capensis, Sp. We... 0.5 ..c.0c ee ceccceeeeeeeceetee eee 604

JED MORO A, SOs 06) lode bas bok oeso6b tocnacspoonacceoaa |» OOS

Thysanotenia (corrections and additions) ...... 606

ASChORBCEULG |e NN oro alts Seeeen eee Ee, OOF

I obtained during the month of October 1911 a large number

of fair-sized unarmed tapeworms from the gut and from the gall-

bladder of an example of Procavia capensis, and I referred them

provisionally to the genus Zschokkeella of Fuhrmann* in my

Report to the Prosectorial Committee, in spite of the fact that

the type species of that genus is a parasite of the Guinea-Fowl

Numida ptilorhyncha. I now refer them partly to the more

recently instituted genus Jnermicapsifer t, the aftinities of which

with Zschokkeella and my own genus Thysanotenia, I shall deal

with in the present communication. The author of the genus

Inermacapsifer—and in the paper referred to—assigns to that genus

with certainty or with more or less doubt eight species. I shall

compare these individually with the species I have found myself.

Dr. Janicki is certainly justified in saying that “the fact is of

interest that the genus Jnermicapsifer is parasitic in Procavia

with several nearly related species.”

From the intestine of Procavia capensis I have gathered

specimens of several distinct species, of which one was much

more abundantly represented than the others. The more

abundant species is represented in the accompanying text-figure

(text-fig. 72).

It is a long and rather slender worm, about 95 mm. in length

and of 2 mm. greatest diameter. There are fully 200 proglottids

in this example. The head is very conspicuous and half as wide

again as the part of the strobila which ensues. Anteriorly the

worm is more slender, of less diameter, than it is posteriorly ;

the increase is gradual, there being no sharp demarcation. The

body of these worms has a great tendency in specimens preserved

* Centralbl. f. Parasit. etc. Bd. xxxii. 1902.

+ Janicki, “ Die Cestoden aus Procavia,” in Schultze’s Zool. Ergebn. Forschungs-

reise in Stidafrika, Jena Denkschr. med. Ges. xvi., 1910.

NEW TAPEWORMS FROM THE HYRAX. 577

in alcohol to contract upwards or downwards, thus forming a

hollow or convex surface as the case may be. The characters

already given allow us to separate the present species from

I. hyracis*, in which the scolex is not of greater diameter than

the ensuing region of the strobila, and from J. settii t, where the

total number of proglottids is very small (i. e. 30-70).

Text-fig. 72.

A nearly complete example and two fragments of Inermicapsifer capensis.

X 2.

I do not at this stage of my description attempt to differentiate

between this species and the others dealt with in Janicki’s

* Janicki, loc. cit. pl. xii. fig. 1.

+ Janicki, loc. cit. pl. xii. fig. 10,

578 DR. F. E. BEDDARD ON

monograph, since there is some little variation in the characters

of the numerous examples which I have to consider here, which

variations do not militate against the exclusion of J. hyracis and

I. settii, though they render difficult the distinguishing of my

species from some of the others referred to by Janicki. Thus

two examples measured only 68 mm. and 75 mm. respectively,

and one of them was quite 3 mm. in diameter at the widest part.

The segments were never longer than broad, except perhaps a

trifle longer in the thin anterior region of the body. They

varied somewhat in relative dimensions, as is the case * with

those of J. hyracis. Occasionally the unilateral genital pores were

very visible on slight papille near to the hinder end of segments.

The scolex is quite unarmed and conspicuous by its size, as

already mentioned. It is usually more or less globular in form,

with a slightly prominent and somewhat pointed apex, as is

shown in Janicki’s figure of J. setéiti7, and which is, of course,

the rostellar region. Occasionally, however, this apical region

does not protrude, but is represented by a depression on the

surface. ‘The scolex is, moreover, sometimes flattened in its

entirety from above downwards, and presents a mushroom-like

appearance, the edges of the disc projecting round the neck,

which thus presents the appearance of the stalk of the

mushroom.

Longitudinal sections through the scolex confirm the total

absence of hooks or of any trace of a rostellum, save the slight

projection already mentioned. Nor could I detect anything

peculiar in the structure of the suckers, which, according to

Janicki and others, are remarkable for a funnel-like ingrowth

leading to the actual sucker, which thus lies at the bottom of

a depression. It is true that in the present species, as in many

tapeworms of the group Tetracotylea which I have examined,

the sucker does not lie externally on the scolex, but is covered by

an outer layer of body-wall which is only interrupted at the

orifice of the sucker. The free edge of this, when depressed

towards the interior of the sucker, is doubtless funnel-shaped

and would give rise to the appearances represented by Janicki.

Perhaps, however, there is some divergence from the normal

condition of the suckers in the members of the genus referred

to by him, which I certainly have not found in the species

with which I am at present concerned. The direction of the

suckers seems to me to vary somewhat; generally they are

lateral, but sometimes with a forward inclination.

It has been already observed that in this genus water-vessels

of unusual width extend far into the head. Such are figured by

Janicki, and referred to by Pagenstecher + (quoted by Janicki §)

as “einen mit Ringmuskeln umspinnenen Wasserbehilter.” In

* Janicki, loc. cit. pl. xii. figs. 2-5.

+ Loc. cit. pl. xii. fig. 10.

I Zeitschr. f. wiss. Zool. Bd. xxx. 1878.

§ Loc. cit. p. 389.

NEW TAPEWORMS FROM THE HYRAX. 579

the present species a good deal of the head lying between the

suckers is occupied by the coils of water-tubes, which approach

very nearly to the external surface at the apex of the head.

These tubes are not in any way irregularly dilated, but their

diameter throughout is rather greater than that of the tubes in

the immediately following neck-region of the body. Janicki is

not disposed to lay great weight upon the absence or presence of

a neck in these tapeworms. In the present form the strobili-

sations begin very close to the head, which is supported by a

segment broadening towards the head, which may be regarded as

the neck. In the accompanying text-figure (text-fig. 72) will be

seen the general characters of the strobila of this worm.

The body-wall of the worm is relatively thick, but here the

cortical layer is not so definitely marked off from the medullary

parenchyma as is so often the case. The absence of a strong de-

limitation is due to the very feeble development of the transverse

musculature, which is generally hardly, or not at all, recognisable

in transverse sections. Another reason which renders the two

layers more uniform in this species than in my Thysanotenia

gambiana and in the species of Hyracotenia described in the

present paper, is that there are stout muscular fibres in the

medullary parenchyma comparable in thickness to the longitu-

dinal muscular fibres of the cortical layer.

Water Vascular Tubes.—These tubes are, as usual, two on each

side. They lie, as a rule, almost parallel with each other, the

smaller dorsal tube having, however, an inclination to the dorsal

side which is more or less pronounced as the case may be. The

more medianly situate ventral tube is the wider, but the difference

between the two is not perhaps quite so well-marked as in

Thysanotenia gambiana. The two vessels are not united across the

proglottids by a regular series of transverse vessels, one to each

proglottid, but, as in other species of this genus and in Thysano-

tenia gambiana, are united by a network of tubes found all over

the segments. I did not follow out into any detail the precise

arrangement of the branching connecting-tubes.

The ovary, when fully mature, appears to be definitely a single

body lying anterior to and in contact with the vitelline gland.

The latter lies on a level with the group of testes of the pore side.

The ovary lies, when less fully mature, entirely to the median

side of the two water vascular tubes. Later on, it appears to

extend somewhat and is larger actually, and considerably larger

relatively, than I found it to be in Thysanotenia gambiana.

The ovary is clearly also larger in the present species than in

Inermicapsifer hyracis, according to Janicki’s figures. In the

later stages of the ovary, when it is gorged with ripe eggs and

these are beginning to spread through the proglottid, a peculiarity

of structure, perfectly plain in earlier stages, is not obvious.

One such example, representing the ovaries, is represented in

text-figs. 73 and 74. It will be there seen that the ovary really

consists of clumps of ripe and unripe eggs situated at the ends of

580 DR. F. E. BEDDARD ON

stalk-like processes of the oviduct, from which they radiate out.

There is some evidence that these are probably hollow outgrowths

of the oviduct, the main stem of which is continued on to form

the rudimentary (at any rate, short) uterus. There is also some

evidence that these outgrowths of the oviducal canal form here

and there anastomoses with each other. In the fully mature

ovary the ripe eggs are seen to be in groups surrounded bya thin

wall and with empty spaces surrounding eggs or groups of eggs.

Text-fig. 73. Text-fig. 74.

Text-fig. 73.—Ovary and vitelline gland of Inermicapsifer capensis in section.

o. Ovary.- V. Vitelline gland. ;

The masses of ova are seen at the end of stalk-like processes.

Text-fig. 74.—Another section through the same ovary to show the rudimentary

uterus (wz.).

It seems to me that we have here a later stage of the tubes

communicating with the oviduct. From this point of view the

ovary would appear to be a compound body, in reality consisting

of many ovaries; the indications of a network among these

ducts is particularly interesting in view of the network formed by

the testicular ducts, and brings the male and female organs into

closer agreement.

NEW TAPEWORMS FROM THE HYRAX. 581

The vagina opens on the exterior behind the aperture of the

cirrus sac and immediately contiguous to it. As will be men-

tioned in describing the cirrus sac there is no common cloaca.

The direction of the tube from its opening into the receptaculum

seminis is obliquely backward, the external pore lying at the side

of the segment posteriorly. The terminal portion of the vagina,

the length of which is not far from equal to that of the cirrus sac,

is somewhat dilated and with a wider lumen ; it is plainly ciliated.

The whole of the vagina is very muscular, the muscular layer

being as thick as that of the cirrus sac. The lining epithelium

is deeply stained. The course of the vagina is very straight until

it bends somewhat at its entrance into the large receptaculum

seminis. The latter is bent round the outermost of the two longi-

tudinal water vessels. From its opposite end arises the wide

common duct, which after a short course divides into the posterior

vitelline duct and the anterior oviduct. The two latter are in

the same straight line.

A closer examination, however, of the long muscular vagina

shows that it is really divisible into two regions. These are

inequisized, the larger section being that which is nearer to the

external pore. This section has a much wider lumen than the

ensuing section, which opens into the wide and thin-walled

oviduct. The latter has a narrow thread-like lumen, but still

thick and muscular walls. It is not so long as the distal region

of the vagina. This part corresponds to the very short and

suddenly abbreviated part of the vagina in Thysanotenia

gambiana ; but is obviously very much more developed. It is

not in the same straight line as the wider distai section of the

vagina, but bends back to meet the forwardly running oviduct.

This division of the vagina appears to be very characteristic of

the present species.

At the time that I was preparing my account of the anatomy of

the two species of my genus Thysanotenia I was not acquainted

with Janicki’s memoir upon his genus /nermicapsifer, and therefore

laid no stress upon some structural features which are of

importance in comparing these forms. I have therefore carefully

re-examined my sections of Thysanotenia gambiana, and made in

addition several series of fresh sections, in order the better to

accomplish this comparison. By these means I am in a position

to make a few corrections in, and additions to, my earlier

account.

I find that account, however, to be correct in all essentials,

excepting with regard to the presence of a vesicula seminalis

in the neighbourhood of the ovary. This was doubtfully

asserted, however, and I now find that the structure in question

is really a part of the dilated and thin-walled oviduct, which runs

straight, as I have said, from the thick-walled region near to the

external pore and gradually increases somewhat in width as it

approaches the ovary. I confirm my account of the variability

in the lateral extension of the ovary and vitelline gland, which

‘

582 DR. F. E. BEDDARD ON

are sometimes restricted in their extent to the space lying

between the dorsal and ventral vessels. In incompletely mature

proglottids this restriction appeared to be (at least usually) the

case, the gonadial tissue not extending outside or inside of the two

excretory tubes. I should also emphasize the fact that the ovary

and vitelline glands are, at least for part of their extent, situated

side by side. That is to say, the vitelline gland is not wholly

behind the ovary. In young proglottids the general lie of the

female gonad seemed to me to be rather oblique dorso-ventrally,

in accordance with the fact that the duct passes to the exterior

ventrally of the dorsal exeretory vessel and that the two excretory

vessels lie in a nearly parallel plane.

T have also a new fact of some little importance to add. I

have described the uterus in the sexually mature segments

as being a tubular structure running across the segments. I

have in younger specimens discovered the rudimentary uterus,

which consists of a solid chord of closely packed cells which

enclose no lumen. This is connected with the gonad tissue

lying between the two excretory tubes and passes towards, but not

far towards, the middle line of the body above the larger ventral

excretory tube, but below the sperm-duct. The rudimentary

uterus ends in a club-shaped blind extremity.

My former account of the testes of Thysanotenia gambiana

is confirmed by the examination of fresh material. I have,

however, two points to add: the first is that while the testes

are practically grouped as there stated into two groups, there

are, nevertheless, a very few testes lying between the two ventral

excretory tubes, and thus really constituting a bridge between

the two separate masses, and bringing the arrangement of the

testes more into line with that of the second species of the

genus Thysanotenia described in the paper referred to. These,

however, are indeed few; I counted but three in a single

proglottid. The general statement with regard to the testes

remains, therefore, unaffected. Moreover, each testis is rounded

or oval in form, but I could not find the distinctly pear-shaped

outline that I describe in the species of /nermicapsifer dealt with

in the present paper. I should, furthermore, add to my former

account the fact that the testes are mature before the ovaries,

and that in segments where the ovary is still immature the

vagina, as well as the vas deferens, is filled with sperm. The

sperm-duct is coiled and the coil hes principally between the

nerve-chord and the dorsal excretory tube, extending to a short

distance mediad of the latter. The coils have some regularity

and lie dorso-ventrally and parallel. I have found granular

tissue accompanying the sperm-duct. Cells lay here and there

between the coils, sometimes in numbers.

The testes of the worm from the Hyrax are very distinctly

arranged in two sets in each proglottid. The great bulk of these

organs are on the side furthest away from the external pore. I did

not find a fewscattered testes uniting these two groups such as occur

NEW TAPEWORMS FROM THE HYRAX. 583

in Thysanotenia gambiana. The point of difference, however, is

not a large one, since in other species of the present genus there

is no differentiation of the testes into two masses, but they form

a continuous mass reaching from side to side of the proglottid.

In the case of the group of testes on the pore side, they lie to the

pore side of the ovary and vitelline gland. These testes were

very conspicuously pear-shaped, the duct naturally arising from

the pointed end. In the case of the testes on the pore side, at

any rate, the narrower end faced forwards in the segment. In the

ease of each group the testes are closely packed together. They

are restricted to the medullary part of the parenchyma. The

efferent sperm-ducts from the testes which lie away from the

pore side form a widish trunk which breaks off and forms aslight

rete mirabile not so marked as in the species next to be described.

This opens evidently by one thick tube into the large vesicula

seminalis lying close to the ovary and to the receptaculum seminis,

The cirrus sac is not conspicuously large, and it lies close to

the lateral margin of each proglottid and thus well within the

cortical parenchyma. Its position is oblique, the end of the

sac being anterior to its aperture on to the exterior, and is

thus better shown in horizontal than transverse sections of the

proglottids. ‘The general outline of the cirrus sac, when displayed

in such sections, is, like that of many other species (e. g.

Otiditenia eupodotidis*), flask-shaped, there being a narrower

proximal neck and a wider distal region. It lies parallel to the

vagina and very close to it, but in a different plane, so that a

single horizontal section does not display both tubes lying side

by side. Such a section, however, does show the extreme

terminal part including the external orifice of both cirrus sac and

vagina; and it will be seen that they lie accurately side by side,

the male pore being, of course, anterior to the female pore. Such

a section also shows very plainly that there is no trace of a

cloaca genitalis ; the two pores are upon the surface of the body

upon a slightly raised papilla of not very great dimensions. The

papilla which bears the two orifices is clearly of less importance

than that which characterises the two worms for which I have

recently formed the genus Thysanoteniat. Ona re-examination

of my sections of the species Thysanotenia lemuris I find that

in that species, as well as in that which forms the subject of the

present communication, there is no sinus genitalis, but that both

cirrus sac and vagina open side by side on the summit of the

papilla. The case is otherwise with Thysanotenia gambiana,

where there is a distinct though not very large sinus or cloaca

genitalis, There are also corresponding differences in the vagina

and cirrus sac of the latter, which I shall deal with after describing

the arrangements found in the Anoplocephalid from the Hyrax.

The cirrus sac has a thickish muscular coat which does not

become at all markedly thicker in the “neck” region. In

* P. Z. 8. 1912, p. 206, text-fig. 27.

+ P.Z. 8. 1911, p. 1002,

584 DR. F. E. BEDDARD ON

longitudinal sections it appears that the fibres constituting

together this muscular wall run more or less along its longer

axis. It will be observed that in both of these characters the

cirrus sac of this species differs from that of a genus recently

described by myself as Otiditenia *, and, moreover, the cirrus sac

is very much larger in the latter genus. The cirrus itself is very

inconspicuous and occupies only the neck region of the sac; it is

thus necessarily very short. It is distinguishable from the

sperm-duct by reason of the fact that it is very darkly stained

with both carmine and hematoxylin. Between the cirrus and

the muscular walls of that part of the cirrus sac in which it lies

is a great accumulation of nuclei, which belong, as I presume, to

slender muscular fibres concerned with the retraction of the

cirrus.

As in other species the cirrus sac, where it swells out into the

rounded flask-like body, is filled with a delicate packing tissue

with abundant nuclei. Through this passes the sperm-duct in

two or three coils. The delicate sperm-duct takes up but little

stain and is thus very distinct from the cirrus. I found this

condition of the sperm-duct to exist in a segment posterior to

others in which the sperm-duct had undergone even further

modification. In the latter segments the sperm-duct lying within

the cirrus sac is dilated to form a vesicula seminalis. This

dilated duct is also coiled; but the two or at most three pieces

seen in an individual section completely fill the lumen of the

cirrus sac, with the exception of dividing lines filled with nuclei

belonging to the internal tissue of the cirrus sac. The fact

that an unaltered sperm-duct may le behind one which is

converted into a vesicula seminalis is important.

It is clear that, on the whole, the cirrus sac of this worm is

more like that of the two speeies which I have referred to a

new genus, Thysanotenia, than it is to that of, for example,

Otiditeniat or Anoploteniat. It differs, however, in details

from the cirrus sac of both of the two species which I have

temporarily placed in the genus Thysanotenia. In the con-

cluding part of the present communication it will be necessary

to go fully into the systematic position of this worm and to

compare it especially with the two species of the genus T’hysano-

tenia. It will be therefore convenient at the present moment

to compare the cirrus sac in these different forms. They agree

generally in the absence of a distal region, which I have termed

penis in Anoplotenia, the cirrus being rod-like up to its free

extremity and not lying at the bottom of an invaginated part of

the cloaca genitalis. Again, in all three species the general form

is the same, and the muscular layer runs along the longer diameter

of the sac and is not specially thickened at the “neck” end.

These features exhaust the general resemblances between the

* P.Z.S. 1912, p. 206, text-fig. 27.

+ P.Z.S, 1912, p. 194.

ft P.Z.S. 1911, p. 1016, text-fig. 215.

NEW TAPEWORMS FROM THE HYRAX, 585

cirrus sacs of the three tapeworms. That of Thysanotenia lemuris

differs from the other two by its larger size, the cirrus sac in the

two remaining species being of about the same size. The cirrus

sac of Thysanotenia lemuris is, indeed, fully twice the size that it

is in the two other species. In the species which forms the

subject of the present communication the cirrus is very small

and limited to the neck part of the cirrus sac, than which it is

no longer. The same characteristics apply to Thysanotcnia

lemuris, only that as the cirrus sac itself is larger, the cirrus also

is larger than that of the species with which I compare it. In

both of these species the rest of the cirrus sac is filled with the

sperm-duct, which is, in both, dilated to form a vesicula seminalis;

but in Thysanotenia lemuris the dilatation fills the whole sac,

while in the other species it is coiled and thus fills the sac in a

different way. In the species Thysanotenia gaumbiana there is no

such conspicuous vesicula seminalis, but just after the entry of

the sperm-duct into the cirrus sac it is dilated for a short space.

On the whole, it appears that the cirrus sac of the present species

is more like that of Thysanotenia lemuris than of Th. gambiana.

One of the principal differences which this species shows from

Thysanotenia gambiana is in the character of the uferus. As I

have already mentioned in the present paper, as well as in my

memoir dealing with that species, the uterus is very plain, first

as a solid cord and then a narrow tube with an obvious lumen.

An examination of a large number of sections of the species of

Inermicapsifer described here has shown nothing exactly like the

uterus of Thysanotenia gambiana. Nor can I reconcile what I

have seen with the figures of Janicki’s illustration of the uterus

of Inermicapsifer hyracis. In the latter species the uterus is

figured * in horizontal sections through the ripe proglottid as an

irregularly shaped sac with numerous projections and outpushings

of its lumen—as, for example, in the genus Jetrabothrium ft.

It is shown, however}, to commence as a sinuous tubular

structure, which appears to me to resemble very much the uterus

as I have described it in Thysanotenia gambiana. “Earlier still

than this a solid cord of cells issues from the generative mass

which again would appear to be exactly comparable to what I

have seen in Thysanoteenta gambiana. Janicki, however, com-

ments upon the remarkable fact that the formation of a lumen in

this cord begins, not where it would be expected to begin, at the

ovarian end, but towards the middle of the segment. It is here,

moreover, that in Thysanotenia gambiana the solid cord of cells,

which subsequently becomes the hollow uterus, widens out into a

club-shaped extremity ; but I am unable to confirm or differ from

Janicki in fixing the point at which the uterus begins to be

hollowed out to be coincident with this club-shaped extremity.

It seems, however, to be very likely.

* Loc. cit. pl. xii. fig. 13.

+ Spatlich, in Zool. Jahrb. Bd. 28 (1909).

[ Loe. cit. pl. xii. fig. 17, wz.

Proc. Zoou. Soc.—1912, No. XX XIX, 39

- 586 DR. F. E. BEDDARD ON

The examination of more than one series of sections has shown

me that, in the species of Jnermicapsifer which forms the subject

of the present communication, the commencement of the forma-

tion of the uterus is in many respects different from that of

TInermicapsifer hyracis as described by Janicki and of Zhysano-

tenia gambiana as described by myself. The earliest stages that

I have observed are perhaps best seen in horizontal sections,

though I have also observed them in transverse sections. In

horizontal sections, such as is represented in text-fig. 74, the

ovary lies in the middle of the segment from before behind and

extends some way towards the middle of the segment from right

to left. The oviduct with its numerous branches, which has been

Text-fig. 75. Text-fig. 76.

Text-fig. 75.—A portion of the medullary parenchyma of Inermicapsifer capensis,

showing centrally a condensation of tissue which is connected with the

rudimentary uterus.

Text-fig. 76.—A portion of the ovary of the same species with eggs (0.) detached

from the ovary and lying in the parenchyma, and a portion of the

network of tissue (a.) which may possibly represent the uterus.

already described, is seen to be prolonged into an extension not

distinguishable from it, which runs for a little way into the

medullary parenchyma towards the median point of the proglottid.

It is quite short, and ends more or less abruptly in a strand of

condensed parenchymal tissue, which is apt to be branched,

sending out shorter strands of a similar appearance in an oblique

direction. Some of these ceased after a short course. In neigh-

bouring regions of the proglottid (text-figs. 75 and 76) there were

patches or rather strand-like parts of the medullary parenchyma

NEW TAPEWORMS FROM THE HYRAX. 587

exactly similar to those which have been described as continuous

with the prolonged oviduct, but not always continuous with it.

They would seem to be produced independently of the generative

ducts, A network arrangement often characterised these regions

of the medullary parenchyma. This condition of the uterus

characterised proglottids in the same stage of evolution as those

which in Thysanotenia gambiana possessed an uterus with a distinct

lumen. Butif weare to compare them with earlier proglottids in

Thysanotenia, then it must be remarked that in the present

species of Jnermicapsifer there is no such regular band of nuclei,

marking out the future uterus, as exists in the former worm.

The appearance is, in fact, totally different. In Inermicapsifer

the suggestion is merely of a crowding together in a quite

irregular fashion and a local multiplication of the nuclei of the

cells of the medullary parenchyma. In Thysanotenia the nuclei

are in orderly arrangement and regular sequence, obviously

belonging to an organ which, as already pointed out, swells at

its medianly situate end into an oval region of greater diameter.

In Thysanotenia we have obviously a regular outgrowth of the

generative system; in my species of Inermicapsifer what would

appear to be a condensation of parenchymal tissue in contact

with and continuing on a very short, and in this species hollow,

outgrowth of the generative duct. I am not clear whether the

species described by Janicki agrees most nearly with the species

described in the present paper or with Thysanotenia gambiana.

Iam disposed to think that the branched strand in which the

slightly prolonged oviduct ends in this Inermicapsifer is not the

homologue of the solid mass of cells in Thysanotenia gambiana,

which afterwards becomes hollowed out to form the uterus

of that worm; in this case Inermicapsifer hyracis agrees more

closely with Thysanotenia gambiana than with the species

described here.

It might possibly be held that this network, which permeates

the segment when seen in horizontal section, is merely a stage

subsequent to that figured by Janicki in a proglottid of some-

where near the same stage of maturity as that which I am now

considering in the species of Znermicapsifer studied by myself—

that the lumen had, in fact, existed and had disappeared. That

this is not the case is clear from the fact that in this stage there

were not any ova contained in the meshwork, and in fact no

ripe ova anywhere outside of the ovary. The meshwork formed

by condensation of the parenchyma, in fact, precedes the extrusion

of ova from the ovary. Still, it may represent the imperfect

remains of a retiform uterus such as is characteristic of the genus

Dipylidium or, better perhaps, in relation to the present genus,

the Anoplocephalid genus Andrya.

I have also been able to approach the question of the identity

or non-identity of the cavity in which the eggs lie with the

cavity of the uterus from another side. Text-fig. 77 represents

portions of horizontal sections through a proglottid in which the

39*

588 DR. F. E. BEDDARD ON

eggs are leaving the ovary to be scattered through the medullary

parenchyma, though in this particular stage the whole paren-

chyma is not filled with eggs as it is later and at the end when

the whole medullary region is crammed with the paruterine

organs. As will be seen by a reference to the figures cited, the

eggs lie partly in round or oval cavities which might well be

Text-fig. 77.

Three sections through the medullary tissue in the neighbourhood of the

ovary of Inermicapsifer capensis.

In A a single ovum is shown to the left. In the centre is an oval area with

strongly marked walls, but filled with the ordinary medullary tissue.

B shows eggs (0.) lying each in a single vesicle of the medullary tissue and three

eggs lying in a cavity (o.c.).

C. Three of the cavities (0.c.) containing eggs and also showing remains of the

medullary tissue which is shown unaltered in A.

thought to be uterine cavities were no further information

forthcoming. I believe, however, that these cavities are not

remnants of a uterus, as is held by Janicki to be the case with

apparently similar cavities in.Inermicapsifer hyracis. A close

NEW TAPEWORMS FROM THE HYRAX. 589

examination of the cavities seems to show that they are lined by

a flattened epithelium, of which the nuclei are apparent and are

represented in the figure, and there is a slight accentuation of

the wall of the cavity, which is merely, as I think, the portion

of the parenchymatous network which abuts upon the cavity and

is not a special layer distinct from that; in this case the ap-

parent lining epithelium should be regarded merely as the nuclei

belonging to this part of the parenchymatous network. I believe

this to be the real explanation; for when we look carefully into

the cavity itself, in which eggs lie in various numbers, there is

nearly always, if not quite always, some granular detritus to be

seen and which is shown in the figures referred to. This formless

detritus (as it often, but not always, is) might, of course, be inter-

preted as simply coagulated fluid which it would not be surprising

to meet with in the interior ,of a uterus were this system of

cavities the remains of a uterus. But, as will be seen, this

detritus is susceptible of another explanation ; it is, as I think,

the remains of the delicate parenchymal network originally

present, as is shown in text-fig. 77 A, in these regions of the

parenchyma set apart for the development of the eggs. The

figure referred to shows plainly an area oval in section and with

a slightly accentuated wall, marking it off from the surrounding

tissue, which is filled with parenchyma network, but does not as

yet contain any eggs. If my contention is right, then it must

follow that the space containing developing eggs is at least not

always to be referred to a uterus. Nor is it in the least against

this view that it is possible to meet with these circular spaces in

which nothing is apparent but eggs—that is to say, no remains

of the originally present parenchymal network, for this may have

completely disappeared. I would furthermore point out that

the position of the eggs in the nearly empty cavity shown in

text-fig. 77 B suggests that they have only just forced themselves

into the cavity.

I have yet another argument to show reason against regarding

the cavities of the paruterine organs in which the embryos are

finally lodged as detached fragments of the pre-existing uterus ;

or, at any rate, to show that they cannot always be so regarded.

In text-fig. 77B and text-fig. 76 is represented an ovum lodged

in one of the meshes of the parenchymatous network of the

medullary region without any special relation to the larger egg-

containing cavities which have been looked upon as detached

fragments of a uterus. There are plenty of such examples to

be seen in sections of this age, and it is plain to me that eggs are

constantly lodged singly in the parenchymal network. I argue

this from the fact that in these cases the cavities lodging the

eggs are in every way indistinguishable in size and appear-

ance from the cavities of the parenchyma in which an egg is

not lodged. It may be, of course, that these are eggs which

got extruded from a uterine cavity and forced into the sur-

rounding parenchymatous network. But it is equally reasonable

590 DR. F. E. BEDDARD ON

to assume that they have got directly to their situation from the

ovary (see text-fig. 76). The lax parenchymal tissue, the mesh-

work of which is filled with a substance plainly visible as granular

matter after staining with Ehrlich’s hematoxylin, but not to be

seen after staining with borax carmine, can offer little obstacle

to the immigration of eggs; so that in any case some of the

paruterine organs are without vestiges of an uterine cavity.

I believe, as a matter of fact, that all are so, and that there is

no persistent uterus in this worm.

The paruterine organs of this species resemble those of the

species which I described as Thysanotenia*. At the time when

that description was written, I believe that Janicki’s careful

account of the genus Jnermicapsifer with similar egg-capsules

had not actually appeared. I had not realised from the descrip-

tions of Zschokkeella that the organs containing the ripe eggs

were doubtless of the same structure. I had considered that

those organs probably resembled the figure given by Ransom 7,

not entirely grasping the fact that that figure was intended rather

as a diagram to distinguish between those species of Davainea

which had several eggs enclosed in one capsule and those species

in which each capsule had within it but one egg. I was thus

misled, though not through Mr. Ransom’s fault. There was, of

course, no other genus with which I could directly compare

Thysanotenia gambiana.

In defining the genus Zschokkeella, Ransom speaks of the fate

of the uterus in the following words :—‘ Uterus early breaks

down into egg-capsules.” arlier in his résumé, Ransom defines

the subfamily Linstowine in the same way ; he remarks that the

“uterus breaks down into egg-capsules.” As the subfamily

Thysanosomine is defined by the presence (inter alia) of numerous

paruterine organs, I thought myself justified in placing my genus,

as I regarded it, in the latter subfamily and marked its affinities

by the use of the generic name Thysanotenia. I was indeed of

opinion that the uterus in Zschokkeella really persisted in separate

pieces, each containing so many eggs. It appeared to me, in fact,

after studying a tapeworm which I have lately described in the

‘Proceedings’ of this Society as Otiditeniat, that the fate of

the uterus in Zschokkeella might be like that of Otiditenia.

No figures are given by Fuhrmann in his account of Zschokkeella

linstowt§ which illustrate this particular point, and the only

reference to the matter is the assertion that the eggs are

surrounded by a “‘ Parenchymhiille.”

Janicki ||, however, is apparently of my earlier opinion ;

for, in distinguishing between the genus Zschokkeella (written,

as originally—Zschokkea) of Fuhrmann and his own genus

* P.Z.S. 1911, p. 1001.

+ Bull. U.S. Nat. Mus. No. 69, 1909, p. 17, fig. 8.

+ P.Z.S. 1912, p. 194.

§ Centralbl. f. Bakt. u. Paras. Bd. xxxii. (1902).

|| Loe. cit. p. 393.

NEW TAPEWORMS FROM THE HYRAX. 591

Inermicapsifer, he uses the differences between the egg-capsules

in the two genera. He considers that in Zschokkeella the eggs

lie ‘‘einzeln in einfache Bindegewebskapseln,” a difference which

also appeared to me to hold good. Ido not, however, feel confident

about this point, and in view of other points of likeness between

the genera am disposed to compare more nearly the paruterine

organs in the two. In my paper upon Ofiditenia just referred

to I have dealt to some extent with the ‘“egg-capsules” of

Davainea, of which the various figures published are not quite

in unison. I find a justification for this in the paruterine organs

of Inermicapsifer capensis, where the appearances vary slightly

among examples which I cannot refer to different species.

To begin with, I prefer the name of paruterine organs for the

structures in question, because they seem to me to be exactly

comparable to structures so named in other tapeworms. They

are, in fact, sacs formed out of the parenchymal tissue, whether

they have or have not ultimately a connection with the uterus.

I should prefer to term the egg-containing spaces in Otiditenia

“ego-sacs” which are formed in a different way, i.e. by a

breaking up of the uterus. When specimens of this Jnermi-

capsifer were examined alive in salt-solution, the individual

paruterine organs could easily be squeezed out of the body

by crushing it. They were then spherical or egg-shaped and

appeared to be surrounded by a thick, colourless, and almost

structureless membrane. This membrane exhibited only faint

strie in a longitudinal direction, being thus concentric in reference

to the whole body. The interior was filled with quite transparent

spherical embryos, between which were abundant cells with

granular contents.

In stained preparations (text-fig. 78) the outer layer was also

perfectly distinguishable. It was stained more lightly than the

inner by carmine and more deeply by logwood. A figure of the

mature organ in Jnermicapsifer hyracis is given by Janicki *, with

which may be compared my own figure of the same organ in the

species described here. The appearances shown by the organ in

the living condition are not borne out by preserved and stained

material. The outer layer is not fibrous but cellular, as shown

by Janicki and others. This layer is, however, as a rule, quite

distinct from the inner mass of cells immediately surrounding

the embryos. The distinction between the two partly depends,

as already mentioned, upon stains, and is not always obvious.

I do not see in my specimens of Jnermicapsifer capensis so great

a distinction between the outer and inner cells in point of size

and shape as does Dr. Janicki, which is perhaps rather remarkable

in consideration of the very different appearances they present

when living. The cells of both layers are, in fact, rounded and

nucleated, and not greatly different in size and shape. Those

of the inner layer are filled with larger spherules. They are

* Loe. cit. pl. xiv. fig. 28.

592 DR. F. E. BEDDARD ON

separated by a more deeply staining reticulum, which appears to

be similar to that of the general parenchyma. This forms a some-

what thicker layer enclosing the whole organ. The reticulum of

the inner mass of cells isthicker than that of the outer layer, and

the nuclei lying at intervals along its strands are very obvious.

The reticulum also forms a distinct layer separating the outer from

the inner mass of cells. In fact, the whole organ is of an appear-

ance more like that represented by Fuhrmann* for Davainea

than that by Janicki. This is so, at any rate, as regards the

outer layer. The interstitial substance of the inner mass of cells

forming the reticulum is represented by Janicki, but as of much

greater relative extent than | have found it.

Text-fig. 78.

—

Paruterine organ of Inermicapsifer capensis, showing differentiation between

outer and inner coat.

The species that has just been described cannot, as I think, be

referred to any of those enumerated by Janicki. It differs from

TI, hyracis in the form of the scolex, in the fact that the testes

are arranged in two separate masses in each proglottid instead

of forming a continuous row. From J. interpositus my species

differs also in the arrangement of the testes, and in the fact that

the genital pores of J. interpositus are anterior in position and

open into a well-marked cloaca, and also in the fact that the

sexual products are ripe earlier in the body in J. inéerpositus.

Nor ean J. setéii be confused with my species. For in the

former the body is very short and consists of not more than

70 proglottids. Moreover, the excretory vessels do not approach

* Rev. Zool. Suisse, iv. (1896).

NEW TAPEWORMS FROM THE HYRAX. 593

so nearly to the apex of the scolex, nor are they so coiled in that

region as in my species. On the other hand, the two species

agree in the posterior position of the genital pores and the

separation of the testes into two masses. There may be a

difference also in the lack of an excretory network in J. setézi,

of the existence of which Janicki is not certain.

There remain certain less-known species referred by Janicki to

the genus Jnermicapsifer with more or less certainty and of whose

characters he gives some account. Of these, “ Zenia paronai”

cannot be identical with my species, since it possesses hooks ;

“ Tenra spatula” of von Linstow is too imperfectly described to

admit of its definite inclusion in the genus Jnermicapsifer.

It cannot, however, be identical with my species, since the cirrus

sac is apparently much larger (‘Der Cirrusbeutel nimmt 1/7

des Querdurchmessers ein”) and the testes are scattered through

the greater part of the segment. Tenia ghondhorensis of

Klaptocz is very imperfectly known, but a pit upon the scolex

shows that it is not identical with my species.

From J. eriticus of Pagenstecher (which is perhaps identical

with another species described below), the present species can

be distinguished by the grouping of the testes into two masses.

I. pagenstecheri of Setti differs from my species by its few

proglottids (net more than 80); otherwise it seems to present

more resemblances to my species than any other form except

I. settw by virtue of the posterior position of the genital pores.

To a species termed by Nassonov Anoplocephala hyracis Rud.

var. hepatica, and by Janicki “ Jnermicapsifer spec.?”’ I shall

recur in considering some worms from the gall-bladder of Procavia

capensis. In the meantime my own species may be thus defined

and named :—

Inermicapsifer capensis, sp. n.

Length about 95 mm., breadth 2mm.; number of proglottids 200.

Scolex wider than the neck. Proglottids at end of body nearly as

long as wide. Genital pores unilateral near to posterior end of

proglottids, not borne upon a conspicuous projection. Testes in

two separate groups, one on pore side and one on opposite side of

proglotiid. Vas deferens forms a network; «a large vesicula

seminalis present ; cirrus sac small, filled with slightly coiled and

dilated sperm-duct. Uterus short and not persistent. Many

paruterine organs, each containing several embryos.

Hab. Procavia capensis.

On Species of the Genus Hyracotzenia, gen. nov.

Along with the numerous examples of the tapeworm which I

have described here as Znermicapsifer capensis, I found in the gut

of the Hyrax two complete or nearly complete examples of a tape-

worm which has quite a different external appearance, and whose

594 DR. F, E. BEDDARD ON

internal structure is different from that of the former species.

Before examining it anatomically it was, of course, impossible to

say whether or no these worms belonged to the genus referred to,

and even now it is possible that they may be of the same species as

one or more of those which Janicki has—provisionally, at any rate,

—assigned to the genus Jnermicapsifer. They are, however,

Text-fig. 79.

Hyracotenia hyracis. X 2.

clearly not members of that genus, as the following account of

their structure will prove. Nor can I, with any confidence, refer

them to any other known genus. HenceI propose the above name,

which is indicative of their habitat. I shall enquire later as to

NEW TAPEWORMS FROM THE HYRAX. 595

the possibility of their identity with any other species known from

the Hyrax. The two worms belong, as I think, to two different

species, but are referable to the same genus without any doubt.

I shall consider the anatomy of both of them together.

The larger specimen (text-fig. 79) measured about 90 mm. in

length by a greatest diameter of 5-6 mm. The proglottids are

very short in an antero-posterior direction, but, at any rate some

way back in the body, rather thick. If there is a neck present at

all it is very short. The scolex is unarmed and distinctly marked

off from the strobila, but not much wider than the ensuing body.

The latter increases gradually in width up to the widest point,

and towards the end of the body again decreases. Of the smaller

specimen I cannot give such precise details, since, believing at first

that both specimens were of the same species, I investigated this

individual anatomically without making full notes on its external

characters. It was, however, rather shorter and of less breadth,

while the anterior narrower region of the body widened out to

the full dimensions rather sooner than in the larger specimen.

I have investigated the scolex by means of longitudinal sections

only in the smaller specimen last referred to. The four suckers

look directly upwards, their orifice being terminal in such sections.

There is nothing remarkable that I could detect about their

structure. They do not bulge to any extent from the sides of

the scolex, and these sections show that the scolex is hardly, if

indeed at all, wider than the immediately following strobila. The

rudimentary rostellum is merely a hemispherical elevation lying

between the suckers. There is no terminal pit of any kind and

no hooks discoverable. The water vascular tubes extend into the

rostellum. Of the larger specimen I only examined the scolex

without destroying it. It is clear that the structure is the same

in all the points mentioned above, but I am not able to report

upon the water vascular tubes in this region.

The structure of the body-wall (text-fig. 80) also differentiates

these two species from the /nermicapsifer whose anatomy has been

described above. The principal difference lies in the much more

marked layer of transversely running fibres which bound the

cortical layer internally and the medullary parenchyma externally.

This layer is very much the same—I think exactly the same—in

both of the two individuals of this genus which I refer later to

different species. It is composed of very delicate fibres; but the

layer, as a whole, is rendered more conspicuous by the fact that large

fibres belonging to the longitudinal layer occur between the trans-

versely running fibres. The cortical parenchyma is nearly as thick

as the medullary. The stout longitudinal fibres which run in the

former are to be found in the greatest numbers at about the

middle of the cortical layer, but they occur elsewhere. They are

not massed into large bundles, but two or three are here and there

closely associated. This massing of the longitudinal fibres is not

obvious in the larger specimen.

596 DR. F. E. BEDDARD ON

The excretory vessels consist of the usual dorsal and ventral tubes

running continuously through the strobila. Anteriorly there is

less difference in the calibre of these tubes than posteriorly. The

position in relation to each other also becomes altered. Inimmature

segments (where I have studied the excretory tubes) the rather

smaller dorsal vessel lies obliquely above the ventral vessel to the

Text-fig. 80.

Hyracotenia hyracis.

Transverse section through a proglottid.

dv. Dorsal excretory tube. JZ. Row of testes. V. Vitelline gland.

v.v. Ventral excretory tube.

pore side of the same. In maturer proglottids the two vessels are

practically superposed, and the dorsal vessel is at times so minute

as to escape observation. Both of these vessels are connected

with a network of larger and smaller water vascular capillaries

which traverse the medullary region of the body. I have seen

branches of these ending in a testis, and it becomes a matter of ,

NEW TAPEWORMS FROM THE HYRAX. 597

great interest to enquire if this network is directly connected with

the network of vasa efferentia. But I have at present no further

facts to offer.

The ovary of this tapeworm is unquestionably single. It lies

on the pore side of each proglottid in the neighbourhood of the two

longitudinal water vessels of that side. The ovary is apparently

sometimes quite anterior to the testes, and sometimes is sur-

rounded laterally by them. The vitelline gland is posterior to the

ovary, which is thus the most anterior of the generative organs.

Text-fig. 81. Text-fig. 82.

d.v.

sp.d. &-S.- Vg.

Text-fig. 81—Part of a transverse section through a proglottid of Hyracotenia

hyracis, to illustrate the-immature ovary.

dw. Dorsal excretory tube. o. Ovary. v.v. Ventral excretory tube.

Text-fig. 82.—Transverse section of one-half of a proglottid of the same species.

d.v. Dorsal excretory tube. o. Part of ovary. 7.s. Receptaculum seminis.

sp.d. Sperm-duct. ¢. Testes. vg. Vitelline gland. v.v. Ventral excretory tube.

Its exact position with reference to the two water vascular vessels

is not always identical. It is more dorsal or more ventral, as the

case may be, and is sometimes entirely to the pore side of the

ventral tube of the excretory system, and sometimes entirely to

the median side of that tube ; sometimes it extends to both sides.

In any case it lies more or less between the dorsal and ventral

excretory tubes. The ovary is not solid and compact, but arranged

in a series of finger-like outgrowths radiating outwards and

598 DR. F. E. BEDDARD ON

dorsally. In young ovaries, such as the one figured in text-

fig. 81, the riper eggs lie in straight lines connected by delicate

threads with the central mass, and suggestive almost of the

pseudopodia of a Rhizopod with the eggs carried along them.

This radiating appearance of the ovary is retained until the organ

is quite mature, when it consists of a group of sacs with, for the

most part, definite walls, enclosed in which lie the ripe ova. I

refer to this appearance of the ovary in considering the uterus on

a later page. It seems possible that this condition of the ovary

is to be compared to the testes, and that there are really several

separate ovaries, which, however, are more closely adpressed than

are the testes. In any case we have instances, like Stilesia,

where the single ovary consists simply of a single mass of egg-

cells, a condition which is to be compared with one of the sub-

divisions of the ovary in the present species.

The vitelline gland in the immature segments lies exactly

opposite to the ovary, the vagina dividing, as described later, into

two ducts, which end respectively in the ovary and vitelline gland.

It is large and conspicuous in the mature proglottid. The shell-

gland is also quite conspicuous in this tapeworm.

The vagina is wider and with thicker walls for a short space

after its orifice on to the exterior. It then narrows and runs a

very straight course towards the interior of the proglottid. It

then becomes again wider, and opens gradually or abruptly into

a dilated receptaculum seminis, which lies beside the vesicula

seminalis. This region of the female duct is thin-walled. It is

on a level with the ventral water-vessel. From the median end

of the swollen receptaculum arise two tubes, one of them being

the vitelline duct and the other the duct leading to the shell-gland

and to the ovary. These two ducts are very much narrower than

the receptaculum, into which they suddenly open. In immature

proglottids the receptaculum is rather wider than the vagina, and

gradually widens towards the internally situated end, there

diverging into two horns which are respectively the vitelline

duct and the ovarian duct. These ducts are in these immature

segments of hardly less calibre than the end of the receptaculum

into which they open. These and the sperm-duct pass towards

the exterior between the dorsal and ventral water vascular tubes.

The terminal region of the vagina, 7.e. that part nearest to the

external orifice of the tube, has a lining which is very deeply

stained by logwood, and so has the narrow region which imme-

diately ensues; the rest of the vagina is not deeply stained in this

way. I noticed in the larger of the two specimens which I report

upon in the present communication that the narrow region of the

vagina lying between the terminal part and the portion which

may be termed receptaculum was much shorter than in the other

example. I am not certain, however, whether there may not be

some variation in this matter from segment to segment, an ex-

pansion of the lumen accounting for the different appearance, It

NEW TAPEWORMS FROM THE HYRAX. 599

is certain, moreover, that the wide internal region of the vagina,

before it divides into vitelline duct and ovarian duct, is susceptible

to variation ; for in some cases it was an abruptly formed spherical

sac, at other times merely a wider tube than the immediately pre-

ceding region of the vagina. I mention later that the end of the

receptaculum contains ripe ova in the mature proglottids. I have

never seen spermatozoa therein.

Uterus.—tIn both of the two examples of this species the last few

segments became somewhat shorter from side to side than those in

front, and were also longer in the antero-posterior direction. One

would naturally associate these changes in the facies of the pro-

glottids with complete maturity and the existence in those segments

ofembryos. As I find in these segments completely ripe eggs quite

detached from the ovary and associated together in small masses,

I shall assume that the anatomical structure of these proglottids

is that of complete maturity. In this case the present species

differs from those which we have been considering by the entire

absence of numerous paruterine organs like those of Jnermi-

capsifer, etc. Kven were these terminal segments not so fully

mature as I presume them to be, there would be, I should

imagine, at least some preparation for the formation of the

paruterine organs. But there is none.

The ripe eggs were massed into more or less spherical groups

surrounded by a membrane. These were not to be distinguished

from the lobes of the ripe ovary, and I imagine that they were

merely the persistent ovary. In addition to these masses of eggs,

the end of the vagina, 7. e. the dilated portion which I have termed

receptaculum seminis, was found in many cases to be full or nearly

full of ripe eggs unaccompanied by any interstitial cells. This

was not only the case witha receptaculum which was swollen into

a spherical contour at its base, but also in cases where the re-

ceptaculum ended merely as a slightly wider tubular sac. I have

no reason whatever to doubt these facts, as the receptaculum is

quite easy of identification. I am therefore disposed to think

that there is no uterus as a distinct and separate structure, but

that the eggs are partly voided into the receptaculum and partly

remain in situ awaiting the loosening and perhaps disintegration

of the proglottids. It may be that one of the rounded sacs which I

regard as an ovary is really a uterus. Of this I cannot be positive,

especially in view of the very few completely mature proglottids

in both specimens. That both specimens were identical looks as

if the conditions above described are to be regarded as normal.

The éestes extend through a good part of each proglottid, and,

as seen in sagittal sections, there are five or six rows of them

laterally, though not so many in the median region. In transverse

sections they are seen to extend from edge to edge of each segment,

being nowhere interrupted save where they meet with the female

reproductive organ. This row is mostly one deep, but in places two

or even three deep. Icounted from 40 to 50 or so of separate testes

600 DR, F. E. BEDDARD ON

in a single complete transverse series. There are thus altogether

two or three hundred of these gonads to each mature proglottid.

In more immature segments the testes did not appear to extend

to the pore side of the longitudinal water-vessels but to stop

short before quite reaching the median side of those vessels. In

a mature segment I found that 44 out of the 50 sections which

displayed it in its entirety were occupied by the testes, which thus

fill up most of the segment, though the proportions were not always

exactly as stated in the above instance. In proglottids from the

other example of this species 3 sections without testes were fol-

lowed by 15 sections showing testes, and these again by 5 without

testes, and thereafter 14 with testes. Itis therefore obviously the

case that the testes occupy a great deal of the segments.

It will be observed that there is no grouping of the testes into

two masses such as I have described in Jnermicapsifer capensis.

They lie mainly behind the ovary and vitelline gland, and in some

proglottids the ovary lay rather more distinctly in front of the male

gonads. The testes are more or less spherical or egg-shaped, and

when ripe are seen to be surrounded by a layer of spermatozoa,

which lie therefore, as I take it, in a cavity surrounding the

testis, a coelomic cavity. I never found the testes of this tape-

worm to be pear-shaped, like those of Jmermicapsifer. Further-

more, the testes, all of them, lie dispersed in quite unaltered

parenchyma. As is very generally the case among the members

of this group, the testes were mature much more anteriorly in the

body to the ovary. It is, indeed, a striking feature of the present

species, and one in which it contrasts, for example, with the

species of Jnermicapsifer that has just been described, that the .

mature testes occupy so many segments of the body, while the

mature ovaries are so exceedingly limited in the number of seg-

ments in which they are found. The efferent tubules which

collect the sperm form a very definite network (see text-fig. 83),

which is copious and formed often of unequally sized vessels. A

similar network has been described in other tapeworms, for

example in Chapmannia*.

The vas deferens of this species (text-fig. 82, p. 597) is quite dif-

ferent from that of the species which we have already considered.

The reticulate efferent ducts finally find their way into a large sac,

which in the mature segments is stuffed with sperm, and which lies

in the female generative mass alongside of the receptaculum

ovorum. ‘This large vesicula seminalis is flask-shaped, and there-

fore gradually narrows and emerges from the female generative

mass as it passes towards the genital orifice. It is impossible to

draw a hard-and-fast line between the vesicula seminalis and the

sperm-duct proper with which it is continuous, for the gradual

diminution in calibre of the entire tube forbids such a delimitation

The tube pursues a winding course, narrowing gradually and but

slightly ; it never forms an actual coil like the sperm-duct of so

* See Fuhrmann, Swedish Zool. Exped. Egypt, 1909, pt. iii,

NEW TAPEWORMS FROM THE HYRAX. 601

many other—indeed the majority of —Tetracotylea, but is at most

once or twice bent upon itself. It becomes very narrow for a

Text-fig. 83.

Portion of medullary region of a proglottid of Hyracotenia hyracis, to illustrate

network formed by vasa efferentia (v.d.).

T. Testis. tr. Transverse muscular layer. v.v. Ventral excretory tube.

Proc. Zoo. Soc.—1912, No. XL. 40

602 DR. F. E, BEDDARD ON

short distance in front of its opening into the cirrus sac. The

course of the sperm-duct is roughly parallel to that of the vagina,

with which it might be sometimes confused in those cases where

the vagina has not so abrupt a transition into the receptaculum.

Along the course of the sperm-duct, which in ripe segments is

gorged with sperm almost throughout, there lie masses of what

appear to be prostatic cells, similar in the fact of their existence

to those of Jnermicapsifer, but different in appearance. In the

present species these cells are of a clear, almost hyaline, ap-

pearance, which is possibly due to the state of their activity at

the time when the worm was killed. In Jnermicapsifer capensis

and in the species which I originally named Thysanotenia gam-

biana, the prostatic cells were darkly staining and granular.

Nevertheless, they appear to be equivalent structures in the two

tapeworms. In sections where the sperm-duct appears in trans-

verse section, these cells present the appearance of a winding duct

cut transversely. This appearance is due to the clear cells

clustered round the actual sperm-duct, which, as already said, is

narrow of calibre close to where it opens into the cirrus sac, and

thus not obvious in such sections. There can be no mistake,

however, in transverse sections of proglottids, where the course

of the sperm-duct is easily to be followed owing to its being filled

with sperm.

In the second and larger individual there are certain definite

differences in the form of the sperm-duct. The tube has no such

great dilatation into a vesicula seminalis, and it is very much more

coiled as it approaches the cirrus sac. It has, in fact, the large

and close coil which is so typical of tapeworms. There is certainly

nothing of the kind in the other individual. The clear cells

already spoken of form a complete layer one cell thick round

the mass of sperm in the sperm-duct, and are therefore, I take it,

simply the epithelial wall of the sperm-duct. As the sperm-duct

was in parts full of sperm, this difference cannot be owing, I

believe, to the different stages of the maturity of the proglottids

in this tapeworm as compared with those already described. It

must, I think, be a specific difference, with which also, it will be

observed, go differences in the position of the ovary and vitelline

land.

: The cirrus sac of this worm is not at all large as in the allied

forms comprised in the genera Jnermicapsifer and Zschokkeella.

It can be seen in sagittal sections to lie straight in front of the

vagina close to the external aperture, and I have not noticed any

genital cloaca. There is certainly nothing of any size, and in one

section the penis was seen to protrude on to the exterior directly

from the cirrus sac without any intermediate and common

chamber. The cirrus sac is oval in form and surrounded, as is

usual, by a strong layer of muscles. I could not see any indications

of a flask shape such as is so common in tapeworms. In the in-

terior of the sac are the usual nuclei belonging, it is to be presumed,

NEW TAPEWORMS FROM THE HYRAX. 603

to delicate muscles which retract the cirrus. The latter was

relatively wide and short and the sperm-duct within the cirrus

sac not coiled. The protruded male copulatory organ reminded

one rather of the penis of Anoplotenia* than of a cirrus, for it

was wider at the free end than just within the cirrus sac.

It is evident that this genus presents many resemblances

to the genera Zschokheella and Inermicapsifert. It agrees with

those genera in the following assemblage of characters :—The head

is unarmed and the excretory tubules form a plexus, or at least a

coil, at the very extremity of the rostellum, as in the species which

has just been described; the segments are narrow and the genital

pores are unilateral. The excretory tubules furthermore form a

plexus within the medullary parenchyma throughout the body.

The ovary lies to the pore side of the segments and is distinctly

not a double organ; the vagina dilates into a wide receptaculum

seminis. The cirrus sac, moreover, is small as contrasted with that

of many other tapeworms. On the other hand, there are certain

characters which argue against this placing of the worm whose

anatomy has just been described. These are as follows :—The

sperm-duct in our species is short and almost immediately dilates

into a large and long seminal vesicle, a state of affairs which is not

met with in the species of Jnernicapsifer known at present.

Finally, the network formed by the vasa efferentia is a feature

hitherto unknown in the genus Zschokkeella, though it occurs in

Inermicapsifer. Inasmuch as a reticular disposition of the vasa

efferentia is not necessarily diagnostic of a given genus as far as

we know, for it occurs in Chapmannia lapica and Hymenolepis

reticulata and not in other (at any rate in some other) species of that

genus, this fact alone would not perhaps necessitate the removal

of the present species from the genus Zschokkeella to which other

important characters would appear to assign it. But there is a

negative character which may be of very great importance. In

neither of the two individuals which I have studied was there the

least trace of the formation of the characteristic ‘‘ egg-capsules,”

which I prefer for reasons already given to call paruterine

organs.

In all the species of Zschokkeella and Inermicapsifer examined

from this point of view, the formation of these capsules began

perhaps rather far back in the body, but still a long way before

the actual termination. Now both examples of the present genus

in my possession ended posteriorly in a few segments which

were rather longer than those preceding them and at the same

time rather narrower, suggesting, in fact, the end of the body.

They were, moreover, thicker than unripe segments. If this be

not the completely mature end of the body, the worm would be

very exceptional in the deferring of the egg-reservoirs to a point

so very far behind the scolex. Besides, two specimens selected at

* See Beddard, P. Z.S. 1911, p. 1015, text-fig. 215.

+ For the generic distinctions see below, p. 607.

40*

604 DR. F. E. BEDDARD ON

random from the same host would hardly be likely to prove both

abnormal in any way. Finally, we have in these segments

rounded sacs with ripe eggs, though it must be admitted that

these were eggs and not embryos. Some riper proglottids may,

however, be missing; but even then the commencement of the

paruterine organs would surely be visible.

It is therefore, as I think, impossible to include these worms

in either of the genera with which I have just compared them.

Of the remaining Anoplocephalide (to which family I think

that these worms must be referred) there are only the genera

belonging to the subfamily Anoplocephaline. Of these Cittotenia

and Moniezia need not be considered, since their generative

apparatus is double in each segment. Of the remaining genera

none agree with the two worms under consideration in all of the

following points, viz., uterus at most inconspicuous, cirrus sac

small, genital pores unilateral, ovary to pore side of proglottids,

genital ducts pass between excretory vessels, testes posterior. I

believe, therefore, that they must be referred to a new genus.

This new genus may be thus defined :-—

Hyracotenia.

Scolex unarmed, with four unarmed suckers. Proglottids wide

and very short, a little longer at extreme end of body, but always

much wider than long. Genital pores unilateral, not borne wpon

papille. Cortical parenchyma thick, separated from medullary by

a thin layer of circular fibres. Water vascular tubes four, dorsal

and ventral, the latter larger, connected by a network of capil-

laries. Testes numerous, dorsal in position, lying behind and to

sides of ovary; vasa efferentia forming a network ; sperm-duct

wide und sinuous or coiled ; cirrus sac small; a short blunt wide

penis protrusible. Ovary near water-tubes of pore side, single, in

front of vitelline gland ; dilated receptaculum seminis and very

narrow vagina. Uterus small and sac-like ; paruterine organs

absent.

Hab. Procavia capensis.

It is not possible for me to distinguish definitely at present

between generic and specific characters. The above embody

characters usually considered in generic definitions. The two

species may be, for the present at least, defined as follows :—

(1) Hyracotenia procavie, sp. n.

Length about 90 mm. ; greatest diameter 5-6 mm. Body attains

its greatest width about 25 mm. from anterior end. Testes very

numerous. Sperm-duct rather dilated posteriorly, much coiled

anteriorly. Ovary ventral, on outer side only or both sides of

ventral vessel ; vitelline glands dorsal. Vagina not greatly dilated

posteriorly.

NEW TAPEWORMS FROM THE HYRAX. 605

(2) Hyracotznia hyracis, sp. n.

Length about 70 mm. ; greatest diameter 45 mm. Body attains

its greatest width about 6 mm. from anterior end. Testes less

numerous. Sperm-duct much dilated posteriorly, sinuous but not

coiled anteriorly. Ovary more dorsal, to median side of water-

vessels ; vitelline glands ventral. Vagina usually much dilated

posteriorly.

We now come to the consideration of the question of the

possible identity of either or both of the above species with any of

those enumerated from the Hyrax by Janicki. The only species

of that series that can be considered (if, that is to say, there are

really no paruterine organs in the forms described by myself)

are Tenia (Anoplocephala?) gondokorensis of Klaptocz*, Tenia

(Anoplocephala) spatula of v. Linstowt, and Anoplocephala hyracis,

var. hepatica of Nassonow £, termed /nernicapsifer spec. ? by Janicki.

Of these Klaptocz’s species has a small scolex like the species

described by me, but also an apical depression (?a rudimentary

rostellum) which my species have not. Furthermore, the pro-

glottids appear to be much smaller. In the species of v. Linstow

we find too great a breadth, and the cirrus sac is too large for

comparison with my species. The shape and proportions of

Nassonow’s species are like mine, but the scolex has a conical

process. Im all these species the details are insufficient.

There are thus in African animals—chiefly in Mammals (mostly

in the Hyrax (Procavia), but also in Rodents and Lemurs), but

extending to Birds (Wumida ptilorhyncha)—a group of worms

which show at least specific differences, but all of which have the

following characters in common, viz., head unarmed and no neck,

proglottids wider than long and as a rule very much so, genital

pores unilateral, cirrus sac not very large, testes numerous, ovary

not double. To these characters may possibly be added, if we

exclude the species described above as Hyracotenia spp., or are

led by further material to interpret their anatomy differently, the

formation of numerous paruterine organs—or egg-capsules as they

have been termed by others.

These characters (excluding the fate of the uterus) seem to me

to necessitate the inclusion of this group of worms in the family

Anoplocephalide.

The various species which agree in the foregoing characters

cannot, however, on these alone be massed into one and the same

genus without further consideration. If we subtract from the

assemblage the species which I have described as Hyracotcenia

_ procavice, and which, as I think, must in any case be withdrawn

from the group, the reasons for uniting the rest under a single

generic name become more striking. Yor in this case all of the

species possess paruterine organs of the same kind, unless, indeed,

* $.B. Wien. Ak. 1906. + Jen. Zeitschr. Naturw. xxxv. 1901.

{ Arb. Zool. Lab. Univ. Warschau, 1897.

606 ON NEW TAPEWORMS FROM THE HYRAX.

Janicki be right in inferring that Zschokkeella really differs, a

point which I have already gone into above (see p. 590). The

description of a second species of Zschokkeella* from a Cerco-

pithecus does not throw any further light upon this particular

matter. If it were not for the fact that Davainea seems in

some of its species to possess paruterine organs of the same type,

the African worms referred to might well be regarded as all

congeneric, in which case, of course, Zschokkeella would have to be

the name.

Janicki appears to me to be rather hard put to it to separate

his Znermicapsifer from Zschokkeella. The differences are certainly

smal]. As already stated, he relies upon supposed differences in

the paruterine organs of which I am disposed to doubt the

existence. He also mentions the thickness of the muscular walls

in Zschokkeella as compared with Inermicapsifer, and a few other

points which seem to me to be of minor importance and not even

collectively as of generic rank. Janicki’s comparisons are based

chiefly upon his own account of Inermicapsifer hyracis, which was

the only species investigated by him in a detailed fashion. I do

not think that a further examination of other species referred to

by Janicki will necessarily prove the identity of /nermicapsifer and

Zschokkeella throughout. I would point out that my own account

in the present paper of J. capensis shows some differences between

that species and J. hyracis. These differences are mainly the

posterior position of the genital pore, the existence of a vesicula

seminalis, the complete separation of two groups of testes, and the

presence of a rete mirabile along the course of the sperm-duct.

Finally, the uterus is much more rudimentary in J. capensis than

in I. hyracis. In some of these characters it would appear that

I. settti agrees with my species and differs from J. hyracis. A

further examination of these species may show that they agree

in other characters not referred to by Janicki in his résumé of

these forms.

I would reserve the generic name /nermicapsifer for these forms

and refer ‘“‘ /nermicapsifer” hyracis to Zschokkeella. 'There now

remains my genus Zhysanotenia. Of that genus I have described

two species which show many differences of structure. Thysano-

tenia gambiana is, as I now think, undoubtedly to be referred to

Zschokkeella, with which it agrees in all points, if we may assume

that the paruterine bodies are identical in the two. On the other

hand, it will be, as I think, advisable to retain the name Thysano-

icenta for the second species of the genus (Z’. lemuris), which differs

mainly in the following points :—There is no plexus of excretory

tubes and the ventral vessel is very large, the dorsal being appa-

rently absent in mature segments; the ventral vessels are connected

in each segment by the usual transverse trunks; the receptaculum

seminis is quite different from that of the other forms; the uterus

* Z. remota, see v. Linstow, Zeitschr. wiss. Zool. Ixxxi. 1905.

ON THE AUSTRALIAN LUNG-FISH. 607

is more rudimentary than in Zschokkeella. I should distinguish

the various genera thus :—

I. Excretory system forms a network in each segment. Recep-

taculum seminis ong and forming end of vagina. Cirrus sac

small.

A. Genital pores median on edge of segment. Testes forming a

continuous row. No vesicula seminalis. Uterus well

developed at firss . . . . . . . . 4Sschokkeella.

B. Genital pores posterior on edge of segment. Testes in two

groups. Large vesicula seminalis present. Seminal ducts

form a network. Uterus never well developed.

Inermicapsifer.

II. No excretory network. Receptaculum short and globular

along the course of vagina. Cirrus sac rather large. :

A. Genital pores on conspicuous papilla. Testes forming

continuous row. Uterus never well developed.

Thysanotenia.

This arrangement is naturally only tentative, since we are at

present in need of more information concerning the majority of

the species already known from the Hyrax and enumerated by

Janicki in the paper which has been so often referred to. There

are also points in the structure of the species referred to the genus

Zschokkeella which demand further investigation.

32. Additional Notes on the Living Specimens of the

Australian Lung-fish (Ceratodus forsteri)in the Collection

of the Zoological Society of London. By Basnrorp

Dean.*

[Received February 9, 1912: Read April 2, 1912.]

(Text-figures 84 & 85.)

InpDEX.

Page

Development and Coloration ........................ 608

Regeneration of injured portion ..................... 611

The two specimens of the Australian Lung-fish in the Zoolo-

gical Society’s collection have been living under unchanged

conditions since 1898, 7. e. about fourteen years. In this time

they have been observed repeatedly by zoologists, whose interest

in these important and rare batrachian-like fishes has led them,

in several instances, to publish their notes in detail. There

is still, however, much to learn about the habits of these

fishes, and it is to be hoped that the opportunity will be

seized generally to observe the present specimens, especially

* Communicated by the SecRETARY.

608 DR, BASHFORD DEAN ON

since it is fair to assume that they are living under fairly

normal conditions. In this connection it may be mentioned

that the fishes have been subjected to no changes in their

aquarial habitat ; indeed, they have remained practically undis-

turbed for over a decade.

Among the notes dealing with these specimens are those of

the present writer, published six years ago in the ‘ Proceedings’ of

the Zoological Society of London (1906, vol. i. pp. 168-178), a con-

tribution which gave also data about Ceratodus summarized from

earlier literature. In this paper details were recorded regarding

the movements of the fishes, their mode of breathing, both with

gills and lungs, their manner of feeding, their nocturnal activity,

and in general their salamander-like habits.

The writer had again the opportunity of observing these

specimens during June 1911; his supplemental notes are as

follows :—

Colour.—One of the fishes, the larger one, remains notably

darker than the other. ‘This distinction in colour, therefore,

is neither abnormal nor seasonal. Norcan there be vast adaptive

colour changes in Ceratodus, for the reaction to similar sur-

roundings would then be the same in the two fishes. Is the

darker specimen a male ?

So far as could be learned, the fishes have shown no evidence

of sexual activity. The colours have remained constant, and

there have been no signs of the brilliant tones noted by Schmeltz

(1876, J. Mus. Godeffy., vol. viii. p. 138). According to his

account, the ventral side of Ceratodus is of a deep orange-red,

and several scales on the sides are margined with red;

nothing is said, however, of the relation of these colours to the

season, From the characters of the present specimens, and by

analogy with Ama, we can safely conclude that the tones of

orange and red appear only at the time of spawning. In Amia,

aquarium-kept fishes show no bright colours, but under native

conditions the male fish develops wonderful brilliancy; the spot

at the base of the tail is conspicuous, red scale-margins appear,

and the hinder abdomen glows with tones of orange.

It was noted (1911) that the paired fins were margined with

a narrow white band. Could this have been an indication of

a breeding colour? No coloration of this kind was seen on the

edges of the unpaired fins.

Size.—At present the darker specimen measures 334 inches,

the lighter 294 inches, having grown but a very few inches during

the past seven years. They have reached, accordingly, nearly

their greatest length, Macleay leading us to infer that 3 feet is

about their maximum (Cat. Austr. Fishes, p. 284). Exception-

ally, however, a specimen may measure 45 inches, such a case

having been cited by O'Connor (1897).

Age.—The present specimens give us an idea of the age to which

Ceratodus may attain. We have in the first place data that they

have grown, broadly speaking, at the rate of three quarters of an

HE AUSTRALIAN LUNG-FISH. 609

inch annually for the last thirteen years. At this rate the present

fishes are over forty years old, and a fish of the record size

(45 inches) would be over sixty. But this assumes that the rate

of growth in length is approximately uniform in fishes of different

ages. In point of fact it is known that fishes in all groups grow

quickly when young, and slowly, if at all, when old: it is also

known that under favourable conditions a fish may grow with

far greater rapidity. In the case of Ceratodus the rate of growth

of the young is as follows :-—

Length of specimen. Age.

inches

E85 i ‘Notes from stages

soa sal nian «am cite At hatching. Se iria ialaneed

42 CDG a AAR ci = month. aquaria in Gayndah,

9 13 th Queensland, by Mr.

TE coccescsse esses 4 montn. Thomas Illidge.)

PS sch ater? a 2 months.

Se sage teat cele 24 months.

ira ecb Mab cdecnetise 6 months.

(ee saben cercges ae 7 months.

3 1 =

at ne pen reac ne user 74 months.

Dien ease sae menee eae 8 months.

In stages lately hatched the growth is seen to be rapid, its

rate suggesting that a specimen 12 inches in length might be

not older than a year. The rate, however, changes notably when

the young fish no longer subsists on its yolk. In fact, for a

period of about two months it actually decreases in size, a state

of affairs, however, probably abnormal and due to the lack of

proper food in the aquarium. Young an inch long are nine

weeks old; young two inches long are over seven months old.

At the end of the first year the young Ceratodus measures, we

infer, about five inches, a rate of growth which would be not

unlike that in young Amia, gar-pikes, or in a number of teleostean

fishes. If the analogy with known ganoids continues, a two-

years’ Ceratodus would measure 8-9 inches; and a specimen

25 inches in length, approximately the size of one of the present

fishes when it appeared in London, is estimated to be from eight

to ten years old; this added to the thirteen years of their

living in the Society’s aquarium, makes the total age of the

present specimens between, roundly, twenty and twenty-five

years. Older and larger specimens it is fair to credit with

great age, probably fifty years.

Breathing.—The aquatic respiration of Ceratodus varies con-

siderably according to the temperature of the water, but its range

has not been observed. In September (1904), on a cool day,

with water temperature not far from 65° Fahrenheit, the respl-

ratory movements were “slow and regular ; the opercular cavity

filled and emptied about twelve times a minute.” In late

June, when the water temperature was nearly 75°, the successive

610 DR. BASHFORD DEAN ON

movements of the gills varied from twenty-two to thirty-one

a minute. Both fish had long been quiet; if their movements

had been active there is no doubt that this rate would have been

notably exceeded.

Rhythmic movements in breathing are well shown in the

opercular membrane. In the early stage of breathing the cheek

in the subopercular region is seen to dilate slowly and strongly ;

this dilated region is then passed (rather slowly) backward, and

Text-fig. 84.

Opercular movement in breathing of Ceratodus forsteri.

The crest of the undulatory wave is indicated by the asterisk.

its enclosed water is discharged (text-fig. 84). During this

process the dilated part becomes more and more conspicuous

until the final discharge, and at that moment the free rim of the

gill-opening is thrown outward and drawn forward, exposing the

lighter coloured hinder border of the gill-slit. The free border up

to that time has been closely apposed to the head.

It was earlier noted that Ceratodus is a “ nostril breather.”

THE AUSTRALIAN LUNG-FISH. 611

The mouth itself showed no noticeable movement of opening

or closing ; it was indeed hardly open, the gape scarcely more than

4 inch. During the later observations, the mouth was seen

slightly to open and close; its maximum gape noted (measured

close to the glass) was 3 inch, its minimum } inch. The opening

of the mouth was here doubtless correlated with the higher

temperature of the water and the more rapid respiratory

movements. In general, however, the mouth margin was

almost motionless, the fish breathing through the nostrils.

In the matter of breathing air at the surface, Ceratodus shows

greater variability than earlier noted. On one occasion over

seventy minutes elapsed without either fish coming to the surface.

Feeding.—Little was added to the former notes. The only

detail suggests that minute food, in the form of alge together

with vegetable débris, forms a part of the normal diet. It was

observed that the fishes would “nose” about in corners and suck

in this finer material. In the process little pebbles would some-

times be taken in and retained for afew moments, then rejected—

the process several times repeated, in a fashion which suggested

that the stones thus mouthed yielded food sought for by the

fish. It was noted in this connection that the stones in the

aquarium were in many cases well coloured with alge.

Text-fig. 85.

Pectoral fin of Ceratodus forsteri, showing regenerating margin.

Movements.—The writer’s preceding paper gave a number of

figures showing characteristic movements of Ceratodus. The

only addition to this series would be a variant of the fig. 11

there given; the right hand _ pectoral remained in its resting

position (as in fig. 14); the left, however, was suddenly twitched

up over the back several times (as in fig. 11), but in this case

brushed close to the body, giving the observer the amusing

impression that the fish was thoughtfully rubbing its head. The

habit was curiously unlike that of a fish; it suggested rather

the movement of a tetrapod; and a very similar movement is

known in urodeles.

Regeneration.—The larger (dark coloured) specimen had suffered

an injury to the left ventral fin; a portion of it near the tip

612 MR. G. H. O'DONOGHUE ON THE

had been lost and the margin was regenerating. Text-fig. 85 illus-

trates how far this process had extended. It will be seen that

the restitutive proliferations were most active along the free distal

margin of the fin. Here several (four) eminences were present, each

suggesting the pointed tip of the fin; there can, however, be no

doubt as to which of these is the terminal one, since the skeleton

of the fin can be followed into the lowest of these lappets. The

case is evidently akin to one known to teratologists, for when

certain areas in injured limbs of batrachians are stimulated, there

appears polydactyly or polypody. It may therefore be worthy of

record that a similar condition occurs in the lung-fish Ceratodus.

33. The Circulatory System of the Common Grass-Snake

(Tropidonotus natria). By Cnas. H. O’ DonocHvE,

B.Sc., F.Z.S., Assistant to the Jodrell Professor of

Zoology, University College, London.

[Received April 1, 1912: Read April 23, 1912.]

(Plates LX X.-LX XII. and Text-figures 86-91.)

INDEX.

Page

DP intbroduehionts sac. sncn ces ee Ree eto ce eece ae orner msec OLS

II. The Heart.

JAW Developmientt:— -<.)SRe Oh camer esos siete dicen snore OL

BPA ult Bormaaee vies co eee enews cose a cee VOLT

III. The Arterial System.

mA Development ec es cn scedenneaee Secmeaese cae) OLD)

BewAta alle B orm ee Wants io hee te aca. Ses Wea eeeeen Dail

(@) AnterioraViesselis: (eco ee eee neeeee ail:

(6) *Bosterior Vessels Vieni. ss) .cosceduanceeen cae scenes) ae)

IV. The Venous System.

JAC Mevelopmient:: 22) .curoress sao eee rene ORG

I Ba ANG 0 Hal os eer meen tA EAI cine taain em antes hecobae (61s)

(a) Anterior Vessels ...........0..0ccc ccc eee cee eeeeeeseeees 630

(b) Posterior Vessels 2.0.00... ...cccseeceeceeeeeereeeeeeeee, 631

V. The Vessels of the Head.

PAM AM TOES ih in cncties case ak elect eRe a even. tek OGD)

DBs. Wiens crc teadecstic net sic coe eee eee oe ere eee OAO.

Wil, Tuistiof References) vince. acces eu reeeee eee cee nner O48

VII. Explanation of Plates ................cc ccc ceecee cee cseceeeseaneeee se, 645

I. INTRODUCTION.

Our knowledge of the circulatory system in snakes is far from

exhaustive ; indeed, we have only a complete account of the

vessels in the Python by Hopkinson and Pancoat (25), and a

later and a more full one by Jaquart (26), and in Pelophilus

madagascariensis by Gadow*. Although Zropidonotus natrix is

* This is incorporated in the account given by Hoffmann (28).

1S2, TE Lee

Paes):

fi oss

riper

aa ge eet

SOMA.

AAV.

ERR,

IMA

HRV.

Ouch

FA.

OusR

ECR

MV.

AVA

Oey

Thy CL

AVP

AAR

RLA

RLV.

PCV.

ja]

Vives

i \

INS

IMAd.

IMAS.

> = SS

a: a

= = oo

O4.AP_

fel.

AAV.

CN.

Bale & Danielsson, It? amp.

Chas. H. O’D onoghue, del.

CIRCULATORY SYSTEM OF THE CRASS SNAKE.

Chas. H. O'Donoghue, del.

i | =A —f—____OavL.P

L |

}—_/—_iRP

DA.

SS; 7

Sy == PV.

0a AR

RPL.

H.PV.O

CaA.

CIRCULATORY SYSTEM OF THE GRASS SNAKE.

Bale & Danielsson, L'? imp

12 Ze Sy IIA. Jl IDear

Op AL

SVAy

Sup.La b. G E

OLA.

Orb,

Ret-A.

/¥ EMA.

RECA.

EMA fit. /os

oy '

he farGl

Sa MA.

A] Oc.

CM.

ISA.

LCM.

: Cer A.

Chas.H. O'Donoghue, del. Bale & Danielsson, L'¢ imp.

CIRCULATORY SYSTEM OF THE GRASS SNAKE.

Ton. dh.

ECA. On,

Ton. Mus,

. MY.

MV.

MA.

ECR. sos

Gils.

OR.

GAN,

SNA.

MxA RY.

Extn oS = ONY,

Sup Lin Gt ; Olf. lo.

OSA Mx.

OPV.

A _CVSA.

Br.

ar Gl.

far. V.

Cep.V. D,

CV,P

Sp.V.

Cer, __

CJL, - e e CIR.

Chas. H O'Donoghue, del. % Bale & Danielsson, Lt4 imp.

CIRCULATORY SYSTEM OF THE GRASS SNAKE.

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 613

the snake most commonly dissected in the laboratories of this

country, no description of its vascular system has appeared since

the anatomical account of the blood-vessels in snakes written by

Schlemm (35) in 1827, which was based largely on Coluber

(i. e. Tropidonotus) natriz and Trigonocephalus mutus. This

account by Schlemm, although excellent in many respects, is by

no means complete, and, owing to the overlooking of the cerebral

_ carotid artery, the remaining arteries of the head are misinter-

preted. A great deal of work has been done, however, on

different parts of the circulatory system of this animal by various

authors. Weare indebted to Rathke for a valuable account of

its development (30) and also of the arteries of the head and

neck (31); the last is the best general account of these vessels in

snakes that has been written as yet. Hochstetter has dealt with

the development of the posterior veins (20) and of the blood-

vessels in general (22), Grosser and Brezina (19) with the

development of the veins in the head and neck, and Bruner (12)

with the veins and sinuses in the head of the adult.

In addition to these works bearing directly on 7’. natria,

Beddard (1-6) has added considerably to our knowledge of the

bleod-vessels of other snakes, and the intracranial circulation has

been dealt with in the vertebrate series in general by De Vriese

(14) and Hofmann (24) and in reptiles by Dendy (13).

The blood-vessels of the Grass-Snake were investigated by

means of the dissection of a number of injected specimens. The

injection fluid used for the main vessels was that recommended

by Kingsley (28), 2. e.,

Corn starch and 2 per cent. chloral hydrate (each) 400 vols.

95 per cent. alcohol 100 vols. and Colour and glycerine (equal

parts) 100 vols.

For the finer vessels a gelatine mixture advised by Tandler (36)

was used, 2. é.,

5 gms. of gelatine in 100 c.c. of distilled water coloured with

Berlin blue or carmine.

5-6 gms. of potassium iodide added slowly while warming

_ gently.

These are two very good mixtures, as the first will keep almost

indefinitely, and the second, with the addition of a few crystals

of thymol, will keep in a stoppered bottle for months, and, in

addition to being useable when almost cold, will withstand acids.

By mixing a quantity of the gelatine mass with about one-third

or less of its volume of the solid residue that settles to the

bottom of the starch mixture, an extremely useful general in-

jection mass is obtained. The latter mass, which flows very

readily if only slightly warmed, and sets firmly and fairly quickly

in 70 per cent. alcohol or in 4-5 per cent. formalin, was the one

most frequently employed in making the preparations for this

investigation,

614 MR. C. H. O DONOGHUE ON THE

For the sake of clearness in description, the account of the

blood-vessels of the head is not included in the general description

of the vascular system of the whole animal, but is dealt with

separately later. A brief account of the development of the

heart and of the arterial and venous systems has been introduced

in order to throw some light on the condition that obtains in the

adult.

Two or three features of general interest in connection with

the elongation of the body and the loss of limbs in the Ophidia

are clearly brought out in dealing with the vascular system of the

Grass-Snake. The first is the marked asymmetry of the viscera

and their blood-supply ; not only are the organs of the right side

anterior to those of the left, but they are also considerably larger.

Thus the right ovary, supra-renal body, and the kidney are in

front of and larger than the corresponding organs on the left,

and, as is well known, in the case of the lungs the left one is

entirely suppressed.

Secondly, the tendency to form longittidinal systems of vessels,

common to all Ophidia, as Beddard (1) pointed out, is well

marked. The various arteries supplying the intestine and the

fat-bodies are in each instance indirectly connected into one long

system. The ovarian artery forms a longitudinal trunk along the

corresponding supra-renal body. Among the veins also we find

that the hepatic portal vein runs from one end of the intestine

to the other, and that each oviduct possesses a sinus running

beside it for the greater part of its length. This oviducal sinus

is very conspicuous in 7’. natrix, although it does not appear to

have been described previously in other snakes. The liver, too,

is greatly elongated, and the post-caval vein and the hepatic

portal vein pass along its opposite faces from one end to the

other.

Lastly, the blood-vessels of the adult, with the exception of a

small pair of veins in the cloacal region, which may represent the

pelvic veins of Lacertilia, give no indication of their derivation

from those of a limb-bearing ancestor.

In conclusion I should like to express my sincere thanks to

Professor J. P. Hill of this College for the kindly assistance and

advice he has given me throughout the work.

II. Tae Hearr. (Pl. LXX.)

(A) Development.

The development of the heart has been very fully dealt with

by Rathke (30), and as this account differs but slightly from that

of Lacerta given by Greil (18) and Hochstetter (20), it does not

appear necessary to give more than a brief outline here.

The primitive heart is in the form of a simple tube stretching

in an antero-posterior direction in the region of the gill-slits.

Its posterior end is formed by the union of the two omphalo-

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 615

mesenteric veins, and the anterior is continued as the short

common stem (truncus arteriosus) of the first pair of branchial

arches. It soon bends towards the right, and as the two ends

remain in approximately the same position, while the tube itself

grows longer, it is forced to take on a curved form. In this

twisted condition three portions can be distinguished: first, a

posterior part running from the union of the omphalo-mesenteric

veins, close to which now open the paired ductus cuvieri,

ventrally and towards the left; secondly, a median part situated

ventrally and running obliquely from left to right ; and thirdly,

an anterior part running from the right to the median line, where

it bends sharply dorsally before passing over into the truncus

arteriosus, from which, by this time, two pairs of branchial arches

are given off.

Grooves appear on the posterior part of the tube which indicate

the divisions between sinus venosus and atrium and between

atrium and ventricle. Into the sinus venosus now open the

paired umbilical veins, The middle part, afterwards to become

the ventricle, becomes dilated ventrally, and as the anterior part

also dilates, the two parts are separated by a deep furrow.

According to Hochstetter (20) and Langer (29) this anterior

part is homologous with the bulbus cordis of the Batrachia.

The ventricle expands still further and moves caudally, so that

the atrium, which has also become dilated, comes to lie close to

the bulbus cordis. The further dilatation of the atrium takes

place cranially and towards the left, causing it to take up a

position on the left side of the bulbus. The constriction between

the atrium and ventricle, corresponding to the auricular canal,

becomes mcre marked as these two structures swell out. At this

stage, too, the sinus venosus is sharply constricted off from the

atrium and the truncus arteriosus gives off the six pairs of

branchialarches. Rathke erroneously described only five branchial

arches, but this will be referred to again later.

After this the ventricle gradually assumes its adult shape

The base of the bulbus cordis, originally joining the ventricle on

the left, moves into an almost mid-ventral position. Spirally

twisted grooves appear between the branchial arches, now reduced

to three in number, and extend downwards over the bulbus.

The atrium now gives off another dilatation, but this time to the

right, and consequently the truncus arteriosus and bulbus cordis

lie in a deep groove between the outgrowths of the atrium.

These two dilatations are the definitive auricles, and already

the sinus venosus, which lies in the atrio-ventricular sulcus on the

dorsal side of the heart, opens into the one on the right.

The groove between the bulbus and the ventricle gradually

disappears, and ultimately the proximal part of the bulbus

becomes incorporated with the latter, while its distal portion

becomes assimilated to the truncus arteriosus. The spiral

grooves finally extend over the whole of the so-formed truncus,

and they indicate its internal division into three parts by the

616 MR, C. H. O'DONOGHUE ON THE

‘backward growth of two septa. One, the septum aortico-

pulmonale, arises from the edge of the pulmonary artery, and as

it grows it divides the truncus cavity into two tubes, an aortic and

a pulmonary. The other, the septum aorticum, arises between

the two aortic arches, and so subdivides the aortic cavity of

the truncus into two, a right and a left. Of the three tubes

formed in this way, one lies to the left, ventrally, and leads to the

pulmonary arch; another lies to the right, ventrally, and leads

to the left aortic arch ; and the third lies dorsally and leads to

the right aortic arch.

Text-fig. 86.

Ventral view of the heart and adjoining vessels.

Az. Azygos vein. C.A. Left common carotid artery. L.A. Left auricle.

L.C.A. Left coronary artery. L.J. Left common jugular vein. L.S. Left systemic

arch. P.A. Pulmonary artery. P.C. Primary carotid. P.V. Pulmonary vein.

P.V.C. Post-caval vein. R.A. Right auricle. R.C.A. Right coronary artery.

R.J. Right common jugular vein. R.S. Right systemic arch. Th.A. Thyroid

artery. V. Ventricle. V.A. Vertebral artery.

Lastly, the sinus venosus also assumes its definitive form.

After the disappearance of the umbilical and omphalo-mesenteric

veins, it has opening into it, on the right, the right ductus

Cuvieri and the post-caval, while on the left it has the left ductus

Cuvieri. Thus we have practically the adult condition of the

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 617

heart, which, however, in the course of the further development

moves caudally, and ultimately comes to lie a long way from its

primitive position.

Text-fig. 87.

2 RI

iG eae

ThA.

PC. RS.

3) Ag,

FAS

LA : RRC.

LPC. oe

By Be PA.

Zs 4 DV.

L \

4 Oy

PV RVC.

Dorsal view of the heart and adjoining vessels.

C.V. Coronary vein. L.P.C. Left pre-caval vein (left common jugular).

R.P.C. Right pre-caval vein (right common jugular). §.V. Major part of sinus

yenosus. §.V.1. Minor part of sinus venosus. Other letters as in text-fig. 86.

(B) Adult Form.

Tropidonotus in common with all the reptiles, except the

Crocodilia, possesses a three-chambered heart. This is situated a

considerable distance behind the head and slightly towards the

right. It is enclosed in a pericardium in which it lies freely,

not being attached to it by a gubernaculum cordis as is the case

in the heart of the Lacertilia and Crocodilia. Beddard (2) has

pointed out that although a gubernaculum cordis is generally

absent in snakes, it is not completely so, as a homologous structure

occurs in some species*. The pericardium on the right side lies

* BH. g. Coronella getula, Celopeltis monspessulana, and Ophiophagus bungarus,

Beddard (loc. cit.).

Proc, Zoo, Soc.—1912, No, XLI. 4]

618 MR. C. H. O DONOGHUE ON THE

against the body-wall, while on the left side it is separated from

it by the intervention of the cesophagus. In conjunction with

the elongated form of the body, we find that the heart also is

long and narrow.

The Sinus Venosus is situated on the dorsal surface of the

heart, and appears as a saccular structure divided into two parts

and formed by the swollen extremities of the common jugular

veins and the post-caval veins (text-fig. 87). It is hardly dis-

tinguishable externally from the right auricle, although internally

the two cavities are separated by the bicuspid sinu-auricular valve.

The right common jugular vein from the anterior end of the body

and the post-caval from the posterior end join together to form

the major part of the sinus venosus, which lies on the right of

the dorsal surface of the heart. The left common jugular vein

runs down along the outer edge of the left auricle and then across

in the groove between the left auricle and ventricle (text-fig. 87).

Its mouth opens into the smaller part of the sinus venosus%*,

which is partially separated from the major part by a valvular

septum. The efficacy of this septum is seen when injecting, for

while the right common jugular and post-caval veins may be easily

injected from the major part of the sinus, it is almost impossible

to inject the left common jugular from it.

The thin-walled Awricles (text-fig. 86) are unequal in size, the

right, of an elongated oval form, being much larger than the left,

which is shorter and more rectangular. Into the right auricle

opens the sinus venosus and into the left the single pulmonary

vein (text-fig. 87). The opening of the pulmonary vein is not

guarded by a valve as Fritsch (16) pointed out, but it seems

highly probable, as Sabatier (33) suggested, that during systole

a fold of the auricle in this region functions as a valve and so

prevents regurgitation. The auricles are completely separated

by an imperforate inter-auricular septum which is continued

caudally so as to divide the auriculo-ventricular aperture into two.

The internal surfaces of the auricles possess a network of raised

muscular ridges, the musculi pectinati.

The Venéricle is somewhat oval in shape, but very asymmetrical.

The posterior end forms a bluntly conical apex, and the base,

although more or less transverse on the right side, is produced

anteriorly on the left side into a conical process, so that the left

side of the ventricle is nearly as long again as the right. It is

extremely thick-walled, and its cavity contains a large number of

muscular trabecule, some of which interlace in such a way as to

form an oblique, incomplete ventricular septum. This partial

septum keeps the aerated blood brought in by the left auricle

more or less completely separated from the non-aerated blood

from the right auricle. Two valves, a right and a left, similar in

arrangement to those in Lacertilia, guard the auriculo-ventricular

apertures.

* They do not open separately into the auricle as stated in Rolleston (82).

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 619

The Bulbus cordis, as has been pointed out above, is not to be

found as a separate structure in the adult, and so the three aortic

arches arise directly from the ventricle. The base of each of

these is guarded by two semilunar valves, which Langer (29) has

shown to be homologous with the distal row of valves in the

amphibian heart.

II. Toe Artertan System. (Pl. LXX.)

(A) Development.

The development of the aortic arches in Z'ropidonotus is very

similar to that of other Reptilia*. It was first described by

Rathke (30), whose general account has been confirmed since by

Van Bemmelen (7 & 8) except in one particular. Rathke

Text-fig. 88.

Diagram of embryonic arterial arches. It represents the condition after the

disappearance of arches 1, 2, and 5, and shows also the division of the truncus

arteriosus into three arterial roots.

C.A. Common carotid. D.A. Dorsal aorta. H.C. External carotid. I.C. Internal

carotid. L.S.A. Left systemic arch. P.A. Pulmonary artery. P.Ar. Pulmonary

arch. R.S.A. Right systemic arch.

Adapted from Hochstetter (22).

describes the development of only five visceral arches on each

side, which he numbered from the anterior end 1-5. Van Bem-

melen, however, showed that there was another arch, which has

however, a somewhat transient existence, between arches 4 and 5

of Rathke, so that the latter’s fifth arch is in reality the sixth of

the series, and thus the snake is brought into line with other

Amniota.

* For a general account of this see Hochstetter ae i

620 MR. C. H. O'DONOGHUE ON THE

These arches soon become reduced to three on each side, viz.

3, 4, and 6, by the disappearance of arches 1, 2,and 5. Of the

remaining arches, 3 is the carotid, 4 the systemic, and 6 the

pulmonary. By the separation of the truncus arteriosus into

three tubes the two carotids and the right systemic have a common

opening into the ventricle; the left systemic opens separately,

and the two pulmonaries open by a common vessel (text-fig. 88).

The most remarkable change in the development is the enormous

lengthening of the carotids, brought about partly by the elonga-

tion of the neck but largely by the caudal shifting of the heart.

Thus it happens that in the adult condition the 3rd arch is far

removed from the 4th and 6th arches.

Text-fig. 89.

Diagram to show changes in embryonic arterial arches. It shows the change

trom the condition in text-fig. 88 to the definitive state. The filled-in portions

yepresent the vessels left in the adult, and those indicated by dotted lines the

vessels that disappear in the course of development.

A.B. Basilar artery. C.C. Left common carotid. P.C. Primary carotid.

Th.A. Thyroid. Other letters as in text-fig. 88.

Adapted from Hochstetter (22).

The Carotid Arch goes through considerable changes in the

course of its development. The two common carotids arise from

the systemic by one root, the primary carotid (carotis primaria

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 621

of Rathke), which remains short as in most snakes*. After the

common carotids have split into internal and external carotids,

each internal vessel gives off a branch which enters the neural

canal with the first spinal nerve and joins the basilar artery on

the ventral side of the nerve-cord. These branches quickly

widen out and, forming as they do an anastomosis between the

two internal carotids, make it possible for the right common

carotid to degenerate. This it does completely f from the point

where it divides into internal and external branches down to

close to its union with the left common carotid, but the last part

of it remains and is to be found in the adult as a small artery

supplying the Thyroid gland (text-fig. 89).

The Systemic Arch undergoes very little change during

development.

The Pulmonary Arch degenerates almost completely on the

left side. In conjunction with the suppression of the left lung

in Tropidonotus, we find that the pulmonary branch of the 6th

arch is only developed on the right side. In the adult only one

pulmonary artery is to be found.

(B) Adult Form.

(a) Anterior Vessels.

The Left Aorta bends dorsally around the esophagus and

trachea and then posteriorly to unite with the right aorta in the

mid-dorsal line. During this course it gives off two very small

branches to the cesophagus, but none whatever to the parietes.

The Right Aorta takes a corresponding course on the other side,

during which it gives off the following branches :—

J. The Left and Right Coronary Arteries arise behind the two

semi-lunar valves which guard the base of the aorta. The right

coronary artery runs in the groove between the auricle and

ventricle, and is the chief supply of the dorsal surface of the

heart. The left passes around the base of the pulmonary artery

and spreads out over the ventral side of the heart.

* In some snakes it is absent altogether, so that the common carotids come off

separately from the systemic arch, e. g. Boa.

+ It is interesting to note, however, that in some variations of 7. natrix this

does not occur, and so the two common carotids persist in the adult. The first

specimen of 7. natria that I examined was in this condition, although in all other

respects if appeared a perfectly normal adult male. The two common carotids, left

and right, sprang from a common stem, the primary carotid, and were about equal

in calibre. On the left side the carotid pursued a normal course. The abnormal

right carotid passed ventral to the esophagus, just behind the thyroid gland, to

which it sent a small branch, over to the right side of the neck. From here up to

the posterior end of the skull it followed a similar course to its fellow on the left.

Unfortunately the vessels of this specimen were not injected, so that the relation of

the persistent right carotid to the basilar artery could not be ascertained. However,

this apparent anomaly in the arterial system is quite readily understood in the light

of the developmental history of these vessels. Only one other example of this

peculiar abnormality seems to have been described before, and that by Van Bemmelen

(Q), but in this case the right carotid was only a fine tube.

622 MR. C. H. O DONOGHUE ON THE

II. The Primary Carotid (Carotis primaria, Rathke) is a short

trunk which quickly divides into two unequal branches :—

i, The Thyroid Artery is the smaller of the two, and, in addition

to supplying the thyroid gland, it sends a twig to the

right thymus glands. This is the sole remnant of the

right common carotid.

ii. The Left Common Carotid (Arteria carotis communis,

Rathke, Arteria cephalica, Schlemm) runs along the left

side of the cesophagus and trachea, to which it sends three

or four slender branches, until it reaches the posterior

region of the head. Here it divides into internal and

external branches, and supplies the whole of both sides of

the head. (The distribution of this vessel in the head will

be dealt with later.)

Ill. The Vertebral Artery (Arteria vertebralis, Rathke, Arteria

collaris, Schlemm) arises from the anterior dorsal part of the right

carotid arch and runs forward, a little to the right of the vertebral

column about half-way to the head. It gives from three to

seven branches to the parietes and one or more to the cesophagus

before disappearing into the vertebral musculature in the mid-

dorsal line.

IV. Five Parietal Arteries are then given off. The first three

are very slender and close together, while the remaining two are

of the same size as the regular parietal arteries.

V. One or two small Wsophageal Arteries run to the cesophagus.

After this the two systemic arches unite to form a single

vessel, the dorsal aorta.

The Left Ductus Botalli is not completely closed up in the

course of development, and its proximal portion is to be found in

the adult animal as a cul-de-sac running cranially from a point

low down on the root of the right carotid arch*. This remnant

varies somewhat in size in different individuals, and although

it is always more or less short, is of nearly the same calibre as

the right carotid. It is completely hidden by the left auricle, but

is readily seen if that body be carefully removed. From its

somewhat bluntly conical end comes off a thin strand of tissue

which runs forward into the left systemic arch at the point where

it bends over to run backwards. This represents the closed part

of the left pulmonary arch, and is therefore the left ligamentum

Botalli, such as has been described by Brenner (11) and Hoch-

stetter (21). I find myself in agreement with the former author

also when he states that he was unable to find a right ligamentum

Botalli in Tropidonotus natria.

According to Brandt (10), quoted also by Hoffman (23), there

* Such a saccular appendage appears to be present generally in those snakes with

one lung suppressed, and has been recorded by Hochstetter (21) in Tropidonotus

natrix, T. tesselatus, Coluber cesculapii, Coronella levis, Vipera berus, and

Cerastes vipera.

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 623

is present in the Grass-Snake a solid strand of tissue joining the

primary carotid to the transverse part of the left aortic arch and

called by him the ligamentum caroticum. This, he states, in

exceptional cases may remain open and may then be described as

a ductus caroticum, and is a vestigeal structure somewhat similar

to the ductus Botalli. In the hearts I have examined no trace of

this vessel or cord could be found, and, indeed, no such connection

exists in the course of embryonic development, as a glance at text-

figs. 88 and 89 will show. A connection between the carotid and

left aortic arches is present in the embryo, but in the adult it

would run from the dorsal part of the left systemic arch along the

whole length of the neck up to the point of origin of the internal

carotid (text-fig. 89). This, however, does not fit in with Brandt's

description of the ligamentum caroticum.

(5) Posterior Vessels.

The Right Pulmonary Artery (Arteria pulmonalis, Schlemm)

arises separately from the ventricle and leaves the heart the most

dorsal of the three arterial roots. It runs backwards alongside

the cesophagus almost parallel with the right systemic over which,

however, it passes ventrally, and then runs dorsal of the post-

caval vein to the anterior end of the lung. As it passes along

the right border of that organ it gradually gets smaller and

smaller until it disappears as a distinct vessel at the level of the

posterior end of the liver, although the lung is continued on for

some distance.

In correlation with the suppression of the left lung no left

pulmonary artery is found at any time.

The Right and Left Aortic Arches unite posterior to the heart to

form the dorsal aorta.

The Dorsal Aorta runs in the body-cavity just ventral to the

vertebral column, back to the level of the cloaca. Just posterior

to this it leaves the body-cavity and enters the hemal canal, and

in this is continued along the tail as the Caudal Artery. During

its course through the ccelom it gives off a number of branches.

The Parietal Arteries form a numerous and more or less regular

series of branches going to the body-wall, of which there are

about twelve up to the point of origin of the superior mesenteric

artery. These arteries enter the body-wall in the mid-dorsal line,

a characteristic of most colubrine snakes, and do not split into

two before so doing, as in the pythonine snakes (cf. Beddard,

4 and 1).

The Gsophageal and Hepatic Arteries.—In front of the liver

the dorsal aorta gives off two or three slender branches to the

esophagus. After these come a series of common trunks, about

fifteen in number, which divide into two branches, one going to

the liver and one to the cesophagus or posteriorly to the stomach.

The last of this series is considerably larger than the others and

has more branches, some of which go to the anterior end of the

stomach.

624 MR, GC. H. 0 DONOGHUE ON THE

The following vessels then come off from the dorsal aorta in

order :—

1. The /zeno-gastric Artery is the first of these. Its gastric

branch is the main artery supplying the stomach, and it also sends

a branch to the spleen, and yet a third, the cystic artery, to the

gall-bladder. No branch of it goes to the pancreas, nor does

the superior mesenteric artery send twigs to the spleen and gall-

bladder as Beddard (1) has described in Zropidonotus fasciatus.

2. The Superior Mesenteric Artery is the largest vessel arising

from the dorsal aorta. Soon after its origin at about the level

of the pancreas it divides into two branches ; a smaller one, the

duodenal, running anteriorly supplies the part of the intestine

immediately after the pylorus and also the pancreas; a much

larger one running posteriorly supplies the many coils of the

intestine as far back as the posterior end of the right ovary.

Small branches from it also supply the anterior part of the fat-

body. I have been unable to find any branch of this artery

running to the right ovary such as Beddard (1) recorded in

Tropidonotus fasciatus.

3. The Right Ovarian Artery, a moderate-sized vessel, runs to

the right supra-renal body and, dividing into anterior and pos-

terior branches, forms a longitudinal vessel along it. From this

longitudinal trunk are given off :—

(a) Six small ovarian arteries of equal size.

(6) Three somewhat larger oviducal arteries. One of these

arises from the anterior end of the longitudinal vessel

and supplies the fimbriated opening of the oviduct, and

the other two arise from the posterior end.

a posterior.

(d) A number of fine twigs to the supra-renal body.

The posterior of the three fat-body arteries in some cases has

an independent origin from the aorta.

4. The first Inferior Mesenteric Artery supplies the coils of the

intestine just posterior to the right ovary. Its point of origin

varies, however, in different individuals and in the two sexes, as

will be pointed out below.

5. The Anterior Right Renal Artery conveys blood to the

anterior half of the right kidney, and also gives off a branch a

the right oviduct.

6. The Left Ovarian Artery is distributed in a very similar way

to the right, and from the longitudinal trunk it forms along the

supra-renal body come off :—

(a) Five equisized ovarian arteries.

(6) Three oviducal arteries. The anterior again supplies the

oviducal funnel.

(¢Z) A number of fine twigs to the supra-renal body.

As on the right side the posterior of the fat-body arteries may

arise separately from the aorta.

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 625

7. The Anterior Left Renal Artery divides soon after its origin

into two branches, one going to the fat-body and the other to

the kidney. This latter branch supplies the anterior half of the

kidney and sends a twig to the oviduct.

8. The Second Inferior Mesenteric Artery arises about the level

of the anterior end of the left kidney, and supplies the intestine

in the region of the median part of the right kidney.

9. The Median Right Renal Artery, in addition to taking blood

to the posterior median part of the kidney, sends a branch to the

right oviduct.

10. The Posterior Right Renal Artery feeds the posterior end of

the kidney and gives off two branches to the oviduct.

11. The Third Inferior Mesenteric Artery, arising near the level

of the posterior end of the right kidney, supplies the intestine in

the region of the posterior part of the left kidney.

12. The Median Left Renal Artery is distributed very similarly

to the corresponding vessel on the right.

13. The Fourth Inferior Mesenteric Artery, arising at the level

of the posterior end of the left kidney, supplies the last part of

the intestine.

14. The Posterior Left Renal Artery resembles its fellow on the

right, save that it does not send a branch to the oviduct.

15. The Posterior Oviducal Arteries are a pair of arteries

running to the posterior ends of the oviducts. In addition to

this there may be one or two small twigs going to the rectum.

16. The Rectal Artery is a small vessel supplying the last part

of the rectum.

An Epigastrie Artery is present, and it appears to be similar to

that deseribed by Beddard (1) in Ophiophagus bungaris, that is,

it runs along the body-wall in the mid-ventral line in close con-

nection with the epigastric vein, and is fed by branches from

the carotid anteriorly and the fat-body posteriorly. Its precise

relations are somewhat difficult to make out, for it is a vessel too

- gmall for individual injection, and in order to get a satisfactory

injection of the arterial system it is necessary to open the snake

from end to end, dissect away the skin, and free the gut to some

extent. This, however, necessitates cutting either the epigastric

artery itself or some of its smail tributaries.

The arrangement of the vessels in the male snake is very

similar to that just described for the female. All the arteries

anterior to and just posterior to the heart are precisely the same,

and it is not until the region of the urino-genital organs is

reached that we find any difference.

Each testis receives one spermatic artery as is general in snakes,

which first runs to the supra-renal body, whence it sends branches

to the testis and also to the vas deferens. These two spermatic

arteries are each followed by another vessel that runs to the pos-

terior part of the supra-renal body and also supplies the vas

deferens for a considerable distance, that on the right side also

sends a branch to the fat-body.

626 MR. C. H. O DONOGHUE ON THE

The kidneys each possess an anterior and a median renal artery

as before, but instead of one posterior trunk there are at least

two, but most often three *. These vessels also send small twigs

to their respective vasa deferentia, and the left anterior renal

supplies the fat-body with a large artery.

The supply to the alimentary canal is also slightly different.

The first of the inferior mesenteric arteries arises posterior to the

anterior left renal artery, whereas in the female it is anterior to

the anterior right renal t. This is followed by a series of about

four smaller inferior mesenteric arteries. The various mesenteric

arteries are connected by their small branches and so form more

or less of a longitudinal system along the gut.

The origin of the arteries supplying the fat-bodies in both sexes

is fairly similar and is somewhat interesting. The anterior part

is supplied by branches from the superior mesenteric artery, the

next portion by vessels from the right genital artery, and the

posterior end is fed by branches from the left genital and left

renal supply. In addition to which the posterior branch from the

genital artery on each side may in some cases arise independently

from the dorsal aorta. All these branches are joined one to

the other by small twigs into a longitudinal system running the

whole length of the fat-body, but there does not appear to be one

unbroken artery, an arteria epiploica, traversing the whole length

of the fat-body.

IV. Tue Venous System. (PI. LXX.)

(A) Development. |

The general course of the development of the venous system

in Tropidonotus is similar to that of other reptiles (vide Hoch-

stetter, 22), but it has some points peculiar to itself (vide Rathke,

30, and Hochstetter, 29).

The first veins to appear are the two omphalo-mesenterics, of

which the right is somewhat stouter than the left (the reverse is

the case in Lacerta), and they open into the sinus venosus. Soon

after their appearance the anterior and posterior cardinal veins

arise on each side, and their common stem unites with the

umbilical vein on each side to form the ductus Cuvieri, which

becomes associated with the omphalo-mesenteric veins at the

* The number of renal arteries varies in different species, being only one in Python

spilotes and eight in Coronella catenifer, Beddard (1).

+ This differs from the account of 7. fasciatus in Beddard (1), where “it springs

from the aorta in both sexes close to the second (i. e. left) gonad artery, in front of it

in the male, behind it in the female.” Some variation is to be found, however, in the

position of this artery in different individuals, for in the females of ZT. natrix that I

have examined, although it has generally been in front of the right anterior renal and

left ovarian arteries, it may be behind these vessels. Inthe male it is usually behind

the left anterior renal artery, but it may be in front of it. It has not occurred

anterior to the right anterior renal artery in any male snake that I have examined.

Again, we find considerable variation in the number of these gut-arteries in various

snakes. Lachesis gramineus has only one, while in the genus Coluber there may be

ten or eleven (Beddard, loc. cié.).

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 627

point where they open into the sinus venosus. An anastomosis

between the two omphalo-mesenteric veins forms on the dorsal

side of the gut just posterior to the pancreas rudiment, and the

portion of the left vein between this point and the sinus venosus

disappears. A similar anastomosis between the two veins now

forms on the ventral side of the gut, and thus a complete ring is

formed. In a short time, however, the right half of this ring

disappears, leaving a single vein which runs in a spiral manner

round the gut. While these latter changes are taking place, the

middle part of the right omphalo-mesenteric vein between the

sinus venosus and the dorsal anastomosis spreads out and forms a

venous network in the liver. The portion of the right omphalo-

mesenteric vein in front of the hepatic network persists as the

hepatic vein, and the part behind it always remains as the anterior

end of the hepatic portal vein (text-fig. 90).

Text-fig. 90.

Diagram of posterior veins in the embryo. It shows the change from an early to

a late embryonic condition. The shaded vessels being the first to disappear.

A.C. Alimentary canal. D.C. Ductus Cuvieri. H.N. Hepatic network. L.O.

Left omphalo-mesenteric (soon disappears). L.U. Left umbilical. O. Omphalo-

mesenteric. P.C. Post-caval. R.O. Right omphalo-mesenteric (disappears later

than L.O.). R.U. Right umbilical. §.V. Sinus venosus.

Adapted from Hochstetter (20).

The posterior cardinal vein originates at the caudal end of the

mesonephros, and runs forward along the dorso-lateral edge of

that body. After leaving the kidney, however, it breaks into

several branches, which soon reunite and then it runs into the

ductus Cuvieril. When the caudal vein develops it divides into

two branches at its anterior end, each of which runs to the

extremity of the corresponding cardinal vein (not along the mesial

wall of the kidney as in Lacerta). The post-caval vein springs

from the union of the right umbilical and omphalo-mesenteric

628 MR. C. H. O DONOGHUE ON THE

veins and runs backwards in the mesentery to the kidneys.

Between them it breaks up into two branches, which run pos-

teriorly along their mesial borders. These branches meet the

kidneys at a point some distance from their front end, and there

each receives a branch from the anterior part of the kidneys.

Subsequently the parts of the posterior cardinal veins in front of

each mesonephros disappear, and so all the blood from the on ndal

veins has to pass through those organs.

Still later the adult kidney arises caudally to the Boke hiros

and then the end part of each posterior cardinal vein comes to lie

on its ventral and external border, while a continuation of one of

the posterior mesonephric branches of the post-caval vein extends

along the inner border of the permanent kidney.

The left umbilical vein loses its opening into the ductus Cuvieri,

and all its blood is taken to the liver network. The right

umbilical vein, on the other hand, retains its original opening

into the ductus for some time (text- fig. 90). In the subsequent

changes the post-caval vein increases in size, and the part of the

omphalo-mesenteric vein joining it breaks up into a venous net-

work with the caudal extension of the liver. As the two umbilical

veins are joined by an anastomosis at the navel, it is possible for

the right to disappear, which it does, leaving the left, which,

however, disappears soon after birth*. Ultimately the omphalo-

mesenteric vein, lying on the dorsal ‘side of the liver, can only

communicate with the sinus venosus wid the hepatic network and

the post-caval vein, which is on the ventral side of the liver.

The anterior cardinal veins originate in a similar way to those

of Selachians and Amphibians, but the parts of these veins in the

head are completely replaced in an interesting way (vide Grosser

and Brezina, 19). The original cardinal vein runs backwards

from the infraorbital and the anterior cerebral veins ventrally to

the cranial nerves into the ductus Cuvieri. Three venous rings

are now formed in succession; the first around the root of the

facial nerve and the auditory vesicle, the second around the root

of the glossopharyngeal nerve, and the third around the vagus

root. Their median portions afterwards disappear, and the external

ones unite to form one lateral trunk. In the meantime two new

vessels come to open into the anterior cardinal: one, the median

cerebral vein, coming from the cerebellum, opens just posterior to

the trigeminal nerve; and the other, the posterior cerebral vein,

coming from the medulla, opens posterior to the vagus, leaving

the skull by the foramen magnum.

Still later the lateral trunk opens anteriorly into the infra-

orbital vein by means of an extension by the side of the second

and third branches of the trigeminal nerve, and posteriorly past

the hypoglossus into the anterior cardinal opposite to the pos-

terior cerebral vein. In this way is formed a new, complete

* Generally the umbilical vein disappears in the adult snake, but remains of it

persist in some species, e.g. Boa constrictor, B. diviniloqua, Python regius, Bunectes

murinus, Corallus cookii. (Bedard. 4& 5).

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 629

lateral trunk, the lateral cephalic vein (V. capitis lateralis, Grosser

and Brezina). At the same time the three cerebral veins become

connected by a median longitudinal vessel (text-fig. 91). Further,

two new anastomoses arise from the median cerebral vein, one

goes to the anterior cerebral vein and the other, the secondary

median cerebral vein, leaving the skull with the trigeminus, goes

to the lateral cephalic vein. This becomes the main vein leading

from the anterior part of the brain.

Diagram of ves in the region of the head in the embryo. It shows the original

veins indicated by the shaded portions and the definitive vessels indicated in

black.

A.C.V. Anterior cerebral. A.V. Auditory vesicle. A.V.C. Anterior cardinal.

I.0.V. Infra-orbital. J.V. Right common jugular. L.C.V. Lateral cephalic.

M.C.V. Median cerebral. P.C.V. Posterior cerebral. P.V. Prosencephalic.

S.M.C.V. Secondary median cerebral. II, V, VII, 1X, X, XII, Primordia of cranial

nerves.

Adapted from Grosser and Brezina (19).

The adult condition is reached by the regression of the dorsal

part of the anterior cerebral vein, the complete disappearance of

the portion of the anterior cardinal vein in the head, leaving its

cervical part, however, as the common jugular vein, and the

formation of a vein bringing back blood from the upper and

lower jaws.

630 MR. C. H. O'DONOGHUE ON THE

(B) Adult Form.

(a) Anterior Vessels.

The whole of the blood from the head is brought back to the

heart by the two common jugular veins. The distribution of the

veins in the head and their union to form the common jugulars

will be dealt with later.

The Left Common Jugular Vein * (V. jugularis sinister, Schlemm)

runs from the anterior end of the neck parallel with and close to

the left common carotid artery to the heart. It passes ventral to

the left systemic arch to the dorsal side of the left auricle, and it

runs along the dorso-lateral edge of this structure to the auriculo-

ventricular sulcus. Here it bends sharply to the right and runs

into a part of the sinus venosus somewhat sharply marked off

from the remainder. It receives :—

i. In its anterior part a few small tributaries from the anterior

end of the cesophagus.

ii. The Coronary Veins (text-fig. 87), a series of small vessels

which join it in its course along the auriculo-ventricular

sulcus. .

The Right Common Jugular Vein* (V. jugularis dextra,

Schlemm) is similar in disposition to the left, but at the level of

the anterior end of the right auricle it receives a large tributary,

the azygos vein. After this it runs straight on and unites with

the post-caval vein to form the major part of the sinus venosus.

I. The Azygos Vein (V. azygos, Schlemm) is a short trunk

leading into the right common jugular vein from the union of the

anterior and posterior azygos veins.

A. The Anterior Azygos Vein (V. azygos anterior, Schlemm) lies

in front of the heart between the cesophagus and the

backbone, and extends from the beginning of the neck

back to the heart. It collects blood from the dorsal body-

wall along this region by means of a fairly regular series

of intercostal veins, all of which come off to the right of

the vertebral column. Just before it unites with the

posterior azygos vein it receives a branch from the

cesophagus.

(a) The Esophageal Vein is formed at the level of the front

end of the thyroid gland by the union of two branches.

One of these branches comes from the anterior dorsal

part of the esophagus, and the other from the posterior

dorsal part. ‘Their common stem runs almost straight

to the right, ventral to the vertebral artery and dorsal

to the right common jugular vein, and opens into the

anterior azygos vein.

* It is to be noted that the proximal parts of these two veins are homologous

with the precaval veins of Lacertilia. As, however, there is no representative of the

subclavian veins to mark the beginning of the precaval portion, it is convenient to

apply the one name to the whole vessel.

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 631

B. The Posterior Azygos Vein (V. Azygos posterior, Schlemm)

is a much shorter vessel than the anterior, and also than

the homologous vein in Lacerta. It originates at about

the level of the middle of the ventricle, and runs forward

to meet the anterior azygos just anterior to the right

auricle. In its course it receives three or four intercostal

veins, all of which come from the body-wall to the right of

the vertebral column.

(0) Posterior Vessels.

The Right Pulmonary Vein (V. pulmonalis, Schlemm) first

becomes noticeable as a definite vessel on the left side of the lung

at the level of the posterior end of the liver. It runs forward

along that organ to its anterior end, receiving numerous branches

and, leaving the lung, it runs parallel to the post-caval vein and

ventral to the right systemic arch to open into the right auricle.

A left pulmonary vein never exists at any time.

The Caudal Vein (V. caudalis, Schlemm) arises far back in the

tail and runs forward in the hemal canal, together with the caudal

artery. It leaves this canal and divides into two branches, the

renal portal veins, a short distance before reaching the cloaca.

The Renal Portal Vein (V. venalis advehens, Schlemm) on each

side runs forward from the bifurcation of the caudal vein over

the cloaca and along the outer side of each kidney. It receives

soon after its origin a lateral tributary, which from its position

and distribution appears to correspond to a pelvic vein*. If this

be the case, it is interesting as being the only indication in the

circulatory system of the derivation of the snake from a limb-

bearing ancestry. Each renal portal vein receives several small

tributaries from the dorsal body-wall, the cloaca, and the lateral

part of the tail. It then passes over the cloaca and lies between

the oviduct which is on its outer side and the ureter which is on

its inner side. Shortly after passing the cloaca each vein gives

off a well-marked branch, the right being anterior to the left,

which passes dorsally around the ureter and then runs ventrally

to unite with its fellow on the dorsal wall of the gut. The vein

so formed, the ‘“Veine mésentérique postérieure” of Jourdain

(27), runs forward along the gut and is continued as the hepatic

portal vein. In its course from the cloaca to the kidney each

renal portal vein receives a number of small tributaries from both

ureter and oviduct, one of which, the Posterior Oviducal Vein, is

well marked, and also one or more from the dorsal parietes. On

reaching the kidney it runs along its lateral border closely accom-

panying the ureter, and, gradually getting smaller, disappears at

the anterior end of that organ. It is not continued anterior to

the kidney as in some snakes, e. g. the Boide and also in Zamenis

gemonensis (Beddard, 4 & 2), a feature recalling the condition in

the Lacertilia. During its course along the posterior part of the

kidney this vein receives a well-marked tributary from the

oviduct.

* A similar pair of veins is to be found in Hunectes murinus (Beddard, 2).

632 MR. C. H. O'DONOGHUE ON THE

The Right Lfferent Renal Vein (V. venalis revehens, Schlemm)

originates along the inner margin of the kidney at its posterior

end, runs to the anterior end, and after a short independent

course in front of the kidney unites with its fellow of the opposite

side to give rise to the post-caval vein.

The Left Hfferent Renal Vein also has its origin along the inner

margin of the kidney. But, unlike the right, it receives branches

from the supra-renal body, the ovary, and the left oviduct in its

course from the kidney to the point where it unites with its

fellow. Thus :—

1. The Left Ovarian Veins are numerous small tributaries

flowing into the left efferent renal veins.

il. The Left Supra-Renal Veins are a large number of small

branches running from the supra-renal body into the

efferent renal vein, to which it is closely attached. They

return the blood brought to that body by the supra-renal

portal veins,

ill. The Left Oviducal Vein is a large vein leaving the oviduct

at the level of the fimbriated funnel and flowing into the

efferent renal vein just in front of the supra-renal body.

It returns blood from the oviducal sinus.

A. The Left Oviducal Sinus is a wide thin-walled vessel

which runs from the extreme anterior end of the ovi-

ducal funnel backwards to about the level of the middle

of the corresponding kidney. It is very conspicuous in

injected specimens and was present in all the female

examples of 7’. natrivx that I have examined, although it

does not appear to have been recorded in any other

Ophidian,

The Post-Caval Vein (V. cava posterior, Schlemm) is formed,

as has been stated above, by the union of the two efferent renal

veins. It passes forward, dorsal to the gut, freely in the mesen-

tery to the posterior extremity of the liver, and then along in a

groove in the ventral surface of that organ to its anterior end.

After leaving the liver it runs almost straight forward, ventral to

the right systemic arch and unites with the right pre-caval to

form the larger division of the sinus venosus, Soon after its

origin it receives veins from the right ovary, the supra-renal

body, and the oviduct. Thus :—

i. The Right Ovarian Veins are similar to those of the opposite

side. ;

ii, The Right Supra-Renal Veins also resemble those on the

left and return the blood gathered by the supra-renal

portal veins.

iii. The Right Oviducal Vein is like its fellow on the left and

comes from an oviducal sinus.

A. The Right Oviducal Sinus extends along the oviduct from

the anterior end of the funnel to about the level of the

middle of the right kidney,

CIRCULATORY SYSLEM OF THE GRASS-SNAKE, 633

~The Supra-renal Portal System.—Each supra-renal body has

a portal supply, an arrangement of vessels which is universally

present in snakes according to Beddard (1), who, however, attributes

the discovery of this system to Gratiolet in 1853 (17), whereas it

had been described seven years previously by Ecker (15). The

supply consists of two, but in more rare cases of three, intercostal

veins which arise from the corresponding side of the dorsal body-

wall and also of a vessel from the oviducal sinus in the female,

all of these pour their blood into the supra-renal network, The

last of these vessels is rather small and runs at the level of the

middle of the ovary on each side,

The Hepatic Portal Vein (Vena porte, Schlemm) arises by two

roots, from the renal portal veins, which unite to form a single

vessel on the dorsal wall of the rectum in a way that has already

been described *, It passes forward through the gut mesentery,

receiving on its way numerous branches from the many coils of

the small intestine and also from the fat-body. ‘Towards its

anterior end a tributary joins it bringing blood from the front

part of the intestine, the pancreas, and the spleen, This is shortly

followed by another vein coming from the gall-bladder. In the

same region it js joined by the anterior abdominal vein which

runs down from the fat-body, From this point instead of being

on the ventral side of the post-caval vein it passes dorsally to the

left of this vessel and enters a furrow on the dorsal side of the

liver. Between the entrance of the vein from the pancreas and

the spleen and the posterior end of the liver, 7. ¢. just anterior to

the superior mesenteric artery, the hepatic portal vein receives

three large intercostal veins from the parietes of the right side.

The vessel runs in the groove of the liver right to its anterior end,

gradually diminishing in calibre, and it receives in this part of its

course a more or less regular series of intereostal veins arising to

the left of the vertebral column and a greater number of small

veins from the stomach and cesophagus. A very similar condition

obtains in 7’. fasciatus (Beddard, 1).

I. The Anterior Abdominal Vein fT, corresponding to the simi-

larly-named vein in Lacertilia, is a single small vein arising at

* According to Schlemm (85) the hepatic portal vein has only one root, and that

arises from the right renal portal vein. “This statement is also made in Hoffmann

(23), but it should be noted that this author quotes nearly the whole of Schlemm’s

account of the venous system almost verbatim without indicating in any way that

he is so doing. My own investigations confirm those of Jourdain (27) aud Hoch-

stetter (20), who describe a double root for this vein in Tvopidonotus, one part

arising from each renal portal vein. The latter author makes a similar statement’

with regard’ to Coluber esculapii. Beddard (4 & 3) has also recorded the same

arrangement in Coluber corais, Zamenis gemongnsis, and Ancistrodon piscivorus.

According to Jaguart (26) there are a number of anastomoses between the hepatic

portal vein and the right renal portal vein in Python. In Gadow’s account of

Pelophilus madagascariensis, quoted by Hoffmann (23), it is stated that there is no

connection hetween the hepatic portal and renal portal veins.

+ This vein is especially interesting, as it is subject to considerable variation among

the Ophidia. In Lizards, as is well known, it arises by two roots from the renal

portal veins, and a similar condition is to be found in some snakes, viz., Hry« jaculus,

E. johni, Python seb@, and Boa diviniloqua, Beddard (2,3, & 4). In other snakes

Proc. Zoou. Soc.—1912, No. XLII, 42

634 MR. C. H. 0 DONOGHUE ON THE

the posterior end of the fat-body and running forward in it

to the level of the spleen, where it passes dorsally and opens into

the hepatic portal vein. It is only connected in an indirect way

by small anastomosing branches with the renal portal veins.

Along its course it receives little twigs from the epigastric vein.

The E'pigastric Vein lies beside the epigastric artery in the mid-

ventral line of the abdominal wall. In the region of the liver it

gives off five or six small branches, all of which enter directly

into the left side of that body and are not connected with the

hepatic portal vein. Behind the liver the epigastric vein is

connected by a number of small venules with the anterior

abdominal vein. As Beddard (4) has pointed out, this is one

of the most constant veins in Snakes, and is single save in

Lioheterodon madagascariensis, where it is alternately single and

double.

The veins in the male are, like the arteries, on the whole very

similar to those in the female. Those in front of the heart are

precisely similar in both sexes.

The caudal vein bifureates to form the renal portal veins which,

at the level of the cloaca, receive the paired pelvic veins, and in

addition, in the male a vein from each corpus cavernosum. The

renal portals give rise to the two branches which unite above the

gut to form the beginning of the hepatic portal vein and then

pass forwards to the kidneys between the vasa deferentia and the

ureters. On the kidney they receive no specially marked tribu-

tary from the vas deferens to correspond with the one from the

oviduct in the female.

Each testis gives off one spermatic vein, whereas in the female

there are a number of small ovarian veins, just in front of the

corresponding supra-renal body, that of the left side opening into

the left efferent renal-vein, and that of the right into the post-

caval vein.

There is no vessel in the male to correspond with the oviducal

sinus, and consequently no branch from it to the supra-renal

body. The supra-renal portal supply consists of two intercostal

veins, one at each end of that body, which arise from the corre-

sponding side of the vertebral column.

Some variation is to be met with in the position of the union

of the two efferent renal veins with regard to the kidneys. The

junction may be as much as an inch in Front of the right kidney,

or, on the other hand, this kidney may overlap the point of union,

in which case sever: al small veins bring back the blood from the

it has only a single origin from the left renal portal vein, viz., Hiya conieus, Hunectes

murinus, and H. noteus, Beddard (8). Lastly, it may have no direct connection

with the renal portal veins, but only indirect ones by means of anastomosing twigs,

viz., Zamenis gemonensis, Beddard (2), Coluber escula, and Tropidonotus ee.

Hochstetter (20)! This last observation I have been able to confirm. Further, the

anterior abdominal vein may be partly double throughout, as in Boa constrictor,

B. diviniloqua, Eryx jaculus, Beddard (4), and Python sebe, Jaquart (26), or

single as in Zamenis gemonensis, Causus rhombeatus, Eryx johni, Beddard (2), and

Tropidonotus natrix.

CIRCULATORY SYSTEM OF TILE GRASS-SNAKE. 635

anterior end of the kidney into the right efferent renal vein, On

the whole this distance appears to be greater in the female than

in the male.

The remaining vessels of the male, the hepatic portal factors,

the anterior abdominal vein, and the epigastric vein correspond

in all respects to those of the female.

V. Tue Vessets or tHE Heap. (Pls. LXXI. & LXXII.)

(A) Arteries.

It has been pointed out above that the whole of the blood

is brought to the head by the left common carotid, the right

common carotid having disappeared early in the course of

development. To compensate for this absence of an artery on

the right ride we find developed three arterial anastomoses

between the two sides of the head. ‘The first lies beneath the

medulla oblongata and joins the two internal carotids ; the second

is situated beneath the fore-brain just in front of the optic

chiasma and unites the anterior cerebral and facial carotids ; and

the third is behind the symphysis of the lower jaw and joins the

two external carotids.

The Left Hxternal Carotid (Carotis externa, Rathke; Arteria

inframaxillaris, Schlemm) arises from the common carotid internal

to the articulation of the lower jaw and the quadrate bone. It

runs forward between the floor of the pharynx and the broad

mylohyoideus * muscle, first inwards towards the tongue sheath

and then outwards to the inner side of the mandible, being

accompanied throughout the greater part of its course by its

corresponding vein, the glossopharyngeal nerve, and the cutaneous

branch of the hypoglossal nerve. At the anterior end of the

lower jaw, just a short distance behind the symphysis, the left

external carotid anastomoses with its fellow by a well-marked

vessel.

The hight External Carotid is similar to the left, save that the

common carotid from which if originated has disappeared and is

represented only by a small branch vessel, It receives its blood-

supply partly from the anastomosis just mentioned, and partly

from the anastomosis between the internal carotids.

The distribution of the arteries in the dorsal part of the head

is the same on both sides, so that the one description will apply

equally well to either side.

The Jnternal Carotid (Carotis interna, Rathke; Art. cephalica

and Art. carotis communis, Schlemm) starts from the origin of

the external carotid and bends in a sharp curve dorsally round

the angle of the lower jaw on the inner side of the vagus and

hypoglossal nerves. It then passes forward under the columella,

and along the inner side of the quadrate to a point behind the

orbit and above the posterior pterygo-sphenoidalis muscle, where

* ‘The nomenclature of the muscles is that adopted by Hoffmann ( 23).

636 MR. C. 1. O'DONOGHUE ON THE

it divides into two branches, the cerebral carotid and facial carotid

arteries,

During this course it gives off the following branches :—

I. One well marked and sometimes also a smaller artery that

supply the cesophagus.

Il. A branch that suppliés a part of the cervico-mandibularis

muscle, the lateral and ventral walls of the pharynx and skin in

this region.

TIL. A branch (Ramus ee caters Schlemm) which supplies

the transverso-maxillo-pterygo-mandibularis, and the cervico-

mandibularis muscles and the skin near them.

IV. The First Spinal Artery (Ramus spinalis, Rathke; Art.

nervi spinalis I, Hofmann) which arises at the same level or

slightly in front of the preceding. The vessel passes through the

atlanto-occipital membrane between the axis and the basi-occipital

bone and, running into the loop of the basilar artery beneath the

medulla oblongata, forms an anastomosis with its fellow of the

opposite side, a relationship first described by Schlemm (35), who

called this vessel the Truncus anonymus. By means of this

anastomosis the blood can pass from the left to the right internal

carotid. Shortly before piercing the atlanto-occipital membrane

this artery gives off two branches :—

A. A small branch (Ramus pterygoideus, Schlemm) to the

Pterygo-sphenoidalis muscles.

B. A fairly large Occipital artery (Art. occipitalis, Schlemm),

which however soon divides into two vessels, a small and a

large.

The smaller is the occipital branch (Ramus occipitalis, Schlemm),

supplying the muscles and skin of the occipital region.

The larger is the Cervical Artery (Art. cervicalis, Schlemm), a

long and fairly well-marked vessel. It runs backwards

near the hypapophyses of the anterior vertebra, covered

by the rectus capitus anticus muscles to which it sends

small branches, to the fourteenth vertebra. Here it joins

with a small branch from the bifurcated extremity of the

vertebral artery. During its course it gives off a series of

branches that lie beside the spinal nerves, and each of

these sends branches not only to the skin and muscles,

but also a small one, a spinal artery, that enters the

neural canal through the intervertebral foramen.

V. The Mawillary Artery (Art. maxillaris, or dentalis inferior,

Rathke; Art. alveolaris inferior, Schlemm) arises about midway

between the spinal artery and the point just behind the orbit

where the internal carotid divides. It runs outwards, accompany-

ing the inferior maxillary branch of the trigeminal nerve between

the second and third parts of the par jetali-quadrato- -mandibularis

muscle, to which it gives branches. After giving off several twigs

also to the transverso- -maxillo- -pterygo-mandibularis muscle, it

passes on through the posterior maxillary foramen into the lower

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 677

jaw. It soon gives off a branch to the inferior labial gland *, and

then runs on in the lower jaw to its anterior end. Here it comes

out through the anterior maxillary foramen as the Mentalis Artery

(Art. mentalis, Rathke) and supplies the anterior part of the

inferior labial gland and the skin in the region of the chin.

Just anterior to this branch the internal carotid splits into two

branches, the cerebral carotid and the facial carotid arteries.

The Cerebral Carotid (Art. carotis cerebralis, Rathke and Hof-

mann) is a vessel of moderate calibre running downwards just

behind the infra-maxillary branch of the trigeminal nerve.

It then turns inwards under the skull and enters the cranial

cavity through a foramen in the basisphenoid. This vessel was

overlooked by Schlemm. In its course it gives off a small artery

just outside the skull going to the posterior pterygo-sphenoidalis

muscle. Inside the skull near the hinder end of the pituitary

fossa it splits into three fairly equal vessels, the posterior, median,

and anterior cerebral carotids,

I. The posterior branch (Ramus caudalis, Hofmann) runs back-

wards round the outside of the pituitary fossa, giving off branches

to the cerebellum and, about half-way to the foramen magnum,

unites with the similar vessel of the other side to form the median,

basilar artery.

The Basilar Artery (Art. basilaris, Rathke and Hofmann)

passes backwards until just before the foramen magnum, where.

it splits up into a loop in the form of an isosceles triangle. Into

the corners of the base of this loop open the right and left first

spinal arteries, and thus it forms the anastomosis between these

two vessels. Before it divides to form the loop, the basilar artery

gives off on either side a well-marked internal auditory artery

(Art. auditiva interna, Rathke and Hofmann) that enters the

ear with the auditory nerve, and also a series of smaller branches,

some of which supply the medulla, and some run on to the small

choroid plexus of the fourth ventricle.

II. The median branch f runs outwards behind the cerebral

hemisphere, to which it gives some twigs, and a short distance

from its origin divides into two.

A. The Anterior Choroid Artery (Art. choroides anterior,

Hofmann) is the anterior branch. It passes around the

hemispheres, supplying them with small twigs, to the

dorsal side of the brain near the pineal stalk, where it

breaks up in the choroid plexus of the third and lateral

ventricles and anastomoses with a branch from the olfactory

artery.

B. The posterior division passes behind the optic lobes and

spreads itself out over their dorsal surface.

* The nomenclature of these glands in the head of the snake is that given by West

(37 & 38).

+ In some examples this median branch does not arise at the point where the

cerebral carotid splits into anterior and posterior branches, but a little way down the

latter ; consequently it appears as a branch of the posterior cerebral carotid and is

described as such by Hofmann (24).

638 MR. C. H. O DONOGHUE ON THE

III. The anterior branch (Ramus cranialis, Hofmann) runs

forward along the side of the pituitary fossa and unites with the

similar vessel from the other side under the bases of the optic

nerves immediately in front of the chiasma. By the union of

the two anterior and the two posterior branches of the cerebral

carotids a complete arterial ring, the circle of Willis (circulus

arteriosus cerebralis, Hofmann) is formed around the pituitary

fossa. During its course this anterior branch gives off :—

A. A Median Cerebral Artery (Art. cerebri media, Hofmann),

which is a large vessel running round to the dorsal side

of the brain and supplying the anterior end of the hemi-

spheres and the olfactory lobes.

B. An Ophthalmic Artery (Art. ophthalmica, Hofmann), which

is given off immediately before the two anterior branches

of the cerebral carotids unite to form the circle of Willis.

This passes out of the skull with the optic nerve, and

immediately on reaching the orbit anastomoses with a

branch of the facial carotid.

The Olfactory Artery * (Art. olfactoria, Schlemm) arises from

the mid-point of the anastomosis of the two anterior branches of

the cerebral carotids and runs forward in the skull in the furrow

between the two olfactory lobes, to which it sends branches. At

the anterior end of the lobes it gives off two symmetrical branches.

Each of these again divides into two (Aa. ethmoidales, Rathke ;

Aa. bulbi olfactorii mediales, Hofmann), which pass out of the

skull with the olfactory nerve to ramify over the olfactory

membrane. The main trunk then recurves dorsally and runs back

in the fissure between the two hemispheres, to which it sends

numerous small branches, and near the pineal body anastomoses

on each side with a branch of the anterior division of the median

branch of the cerebral artery.

T.e Inferior Spinal Artery (Art. spinalis inferior, Rathke ;

Tractus spinalis ventralis, Hofmann) is situated just below the

ventral fissure of the spinal cord, and runs in a fairly straight line

caudally from the anastomosis between the right and left first

spinal arteries at the posterior end of the basilar artery. On its

course it gives off branches to the spinal cord, some of which pass

around to the dorsal side and others enter the ventral fissure, and

it also receives the paired spinal arteries which come in through

the vertebral column at the points of exit of the spinal nerves.

In some places where a pair of such arteries enter it, the inferior

spinal artery splits into a diamond-shaped loop.

The Facial Carotid (Carotis facialis, Rathke) takes a fairly

siraight course forward through the temporal fossa close to the

infra-maxillary branch of the trigeminal nerve to the orbit.

Here it passes under the post-frontal bone into the orbit and

* This artery is double for the greater part of its length in Python molurus,

Beddard (6).

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 639

immediately divides into two fairly equisized terminal branches.

On its way it gives off :—

I. A branch (Ramus glandule maxille superioris posterior,

Schlemm) which arises a short distance behind the orbit and runs

outwards to the posterior part of the parotid gland, @. ¢., the gland

corresponding ‘to the poison-gland of other snakes. It ramifies

in the gland, and one of its branches anastomoses with a branch

of the artery supplying the anterior part of the gland. During

its course it gives off a palatine branch and branches to several

muscles.

II. Several small branches and one fairly well-marked one that

supply the Harderian gland.

ILI. 'wo or three slender branches to the skin overlying the

skull and one or two to the under side of the skull.

One terminal branch (Carotis facialis, Schlemm) of the facial

carotid passes along the back of the orbit downwards to its floor,

where it runs forward (Ramus palatinus anterior, Schlemm),

accompanying the infra-orbital branch of the trigeminal nerve.

It goes on forward out of the orbit, close to the outer wall of the

internal nares, and finally spreads out over the skin and muscles

at the front end of the snout. During its course it gives off :—

T. A well-marked vessel (Ramus glandule maxillaris superioris

anterior, Schlemm) that accompanies the supra-maxillary branch

of the trigeminal nerve outwards in the posterior wall of the

orbit and along the upper jaw. It supplies the teeth in the pos-

terior part of the upper jaw, the superior labial gland, the

anterior part of the parotid gland, within which one of its small

twigs anastomoses with a similar twig from the artery supplying

the posterior part of this gland, and finally it gives off small

vessels to the skin in this region.

II. A posterior palatine branch (Ramus palatinus recurrens,

Schlemm) that goes to the posterior palatine teeth.

III. Small branches to the anterior palatine teeth.

IV. Branches to the teeth in the anterior end of the jaw.

V. Branches to the nasal gland.

The other slightly stouter terminal branch of the facial carotid

artery (Art. carotis cerebralis, Schlemm) runs in an irregular arch

along the upper, inner, and hinder sides of the orbit to the optic

foramen, where it anastomoses with the ophthalmic artery. In

this way the blood from the left facial carotid can pass vid the

ophthalmic artery into the circle of Willis, and thence to the

encephalic arteries and also over into the right facial carotid. In

its course this artery sends out the following branches :——

T. Several well-marked twigs to the part of the Harderian

gland within the orbit.

TI. Branches to the muscles of the eye (Aa. musculares oculi,

Rathke).

ILI. Two short ciliary branches (Aa. ciliares postice breves,

Rathke).

640 ‘ MR. C. H. O DONOGHUE ON THE

IV. Two longer ciliary branches (Aa. ciliares postice longa,

Rathke), one of which runs forward on the inner side of the eye

and the other outward on its hinder side.

V. A Retinal Artery (Art. centralis retine, Rathke) which

enters the eyeball with the optic nerve and spreads out over the

retina.

(B) Veins.

The veins of the head of TVropidonotus natrix have recently

been described in great detail by Bruner (12), who also describes

a muscular mechanism in the head whereby the blood-pressure in

its veins and sinuses may be considerably increased. It is not

possible by ordinary dissection to make out all the small vessels

given by that author, and, as my own results agree closely with’

his, it will only be necessary, for the sake of completeness, to give

a brief description of the cephalic veins.

The Mandibular Vein (V. inframaxillaris, Schlemm ; V. mandi-

bularis, Bruner) arises from a small sinus at the anterior end of

the lower jaw and runs backwards close to and on the outer side of

the external carotid artery. On its way it receives veins from the

trachea, tongue-sheath, muscles of the floor of the mouth, and:

the pharynx. I¢ runs into the maxillary vein immediately before

the latter joins with the lateral cephalic vein to form the common

jugular vein. The bases of the mandibular, maxillary, and lateral

cephalic veins and the anterior end of the common jugular vein

are surrounded by constrictor muscles whose morphology and

function are described by Bruner (Joc. cit.).

The Maxillary Vein (V. palatina, Schlemm; V. maxillaris,

Bruner) also commences in a small sinus which is situated just

behind the premaxilla. This sinus has a double anastomosis with

the similar one on the other side. From this point it runs back-

wards beneath the nasal cavity along the floor of the orbit and

then above the palate to join the mandibular vein at the angle of

the lower jaw. During its course it receives :—

I. The Rostral Vein (V. rostralis, Bruner), which enters at the

level of the anterior anastomosis and brings the blood from a

venous network at the front end of the snout. It also receives

nasal veins (V. nasales externe, dorsalis, and ventralis, Bruner)

from the nasal gland.

Il. The Subnasal Vein (Sinus subnasalis, Bruner), which enters

at the level of the posterior anastomosis. It drains a somewhat

complex system of subnasal sinuses, which anastomose with one

another at the posterior end of the nasal cavity and which receive

also the palato-pterygoid vein.

The Palato-pterygoid Vein (Sinus palato-pterygoideus, Bruner)

runs from near the anterior to near the posterior end of

the skull on the inner side of the palatine and pterygoid

bones, and flows into the anastomosis between the sub-

nasal sinuses.

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 641

- TIT. A vein that joins it at the anterior end’ of the orbit and

forms an anastomosis between it and the orbital sinus.

IV. An Inferior Palpebral Vein (V. palpebralis inferior, Bruner)

that also enters at the front end of the orbit and runs backwards

. in the lower eyelid to its posterior end, where it enters the orbital

sinus together with the superior palpebral vein.

V. Several small veins from the orbital sinus just posterior to

the orbit.

VI. An Oblique Palatine Vein (V. palatina obliqua, Bruner)

that runs obliquely forward beneath the skull at the level of the

hypophysis to join the similar vessel of the opposite side and form

a median palatine sinus which runs forward for a short distance.

Before reaching the middle line it gives off a palato-cerebral

vein.

The Palato-cerebral Vein (V. palato-cerebrales, Bruner) runs

dorsally around the skull and enters the median cerebral

vein soon after this leaves the cranium.

The Orbital Sinus (Sinus orbitalis, Bruner) is a fairly large well-

defined sinus occupying the inner and hinder parts of the orbit.

At the outer end of the hinder part it is prolonged outside the

orbit beneath the Harderian gland. It receives the following

branches :—

T. A small vein at its anterior end which comes from the nasal

gland.

IL. The vein joining it to the maxillary vein which also enters

at the anterior end.

III, The Superior Palpebral Vein (V. palpebralis superior,

Bruner), which arises at the anterior end of the sinus and runs

backwards in the upper eyelid, re-entering the sinus at its

posterior end and receiving just before it does so the inferior

palpebral vein.

IV. The Secondary Anterior Cerebral Vein (Sekundire Verbind-

ung der v. cerebralis media mit der v, cerebralis anterior, Grosser

und Brezina) which runs from the posterior internal corner of the

orbital sinus backwards inside the skull into the median cerebral

vein just as the latter is leaving the cranial cavity. The anterior

segment of this vessel is formed by a part of the original anterior

cerebral vein.

The Lateral Cephalic Vein (V. capitis lateralis, Grosser und

Brezina; V. jugularis interna, Bruner) arises from the posterior

prolongation of the orbital sinus and runs inwards and backwards

to the side of the internal carotid artery. It passes backward

closely accompanying this artery to the posterior end of the head,

where it bends round to the ventral side and unites with the

maxillary and mandibular veins to form the common jugular vein.

On its course it receives :—

I. A vein from the Harderian gland.

II. The Median Cerebral Vein (V. cerebralis media, Bruner).—

This runs from the longitudinal cerebral vein on the dorsal side

642 MR. C. H. O'DONOGHUE ON THE

of the brain, outwards, and around the posterior face of the optic

lobes to the ventral side of the brain. Here it goes forward and

leaves the skull by the foramen for the trigeminal nerve. Out-

side the skull it bends sharply backwards and joins the lateral

cephalic vein as the latter reaches the internal carotid artery.

The last part of this vessel outside the skull is a secondary con-

nection (V. cerebralis media secundaria, Grosser and Brezina)

developed between the median cerebral vein, which originally

opened into the internal jugular, and the lateral cephalic vein.

During its course it receives :—

A. The Dorsal Cephalic Vein (V. capitis dorsalis, Bruner), which

arises from its dorsal side within the skull and passes

outward through a special foramen. It runs backwards

between the pro-otic and squamosal bones, receiving one

or two cutaneous veins, and then bends laterally and enters

the lateral cephalic vein.

B. The Secondary Anterior Cerebral Vein, which runs on the

floor of the cranium and joins it to the orbital sinus (vide

supra).

C. The Palato-cerebral Vein, which connects it with the oblique

palatine vein (vide supra), According to Bruner there

is also an external secondary anastomosis with the anterior

cerebral vein, as well as the internal one described above.

I have been unable to find this vein by dissection.

III. A large vein from the parotid gland and the muscles of

the head, which closely accompanies the maxillary artery and

enters the lateral cephalic vein close to the place where the

maxillary artery leaves the internal carotid.

IV. A Dorsal Cephalic Vein (vide supra), which joins it to the

median cerebral vein.

V. The Posterior Cerebral Vein (V.cerebralis posterior, Bruner),

which runs from the end of the longitudinal cerebral vein a little

behind the posterior end of the optic lobes obliquely outwards

over the medulla oblongata and leaves the skull by the foramen

magnum. Just before leaving the cranium it gives off a Spinal

vein Which runs caudally on the ventral side of the spinal cord,

where it unites with the similar vessel from the other side.

VI. One well-marked and several smaller veins from the

muscles of the posterior end of the skull.

VII. A Cervical vein which returns blood from the muscles of

the neck.

The Longitudinal Cerebral Vein (V.longitudinis cerebri, Bruner)

is a vessel running backwards along the mid-dorsal aspect of the

brain from between the olfactory lobes. At the posterior end of

the optic lobes it gives off the median cerebral veins, and a short

distance further back divides to form the posterior cerebral

veins,

CIRCULATORY SYSTEM OF THE GRASS-SNAKE. 643

VI. List or REFERENCES.

(1) Bepparp, F. E.—Contributions to our Knowledge of the

Circulatory System in the Ophidia. Proce. Zool. Soc.

ie 1904, vol. i. p. 331.

(2) Bepparp, F. E.—Notes upon the Anatomy of Certain Snakes

of the Family Boide. Proc. Zool. Soc. 1904, vol. ii. p. 107.

(3) Bepparp, F. E.—Contributions to the Anatomy of the

Ophidia. Proc. Zool. Soc. 1906, p. 12.

(4) Bepparp, F. E.—Contributions to the Knowledge of the

Vascular and Respiratory Systems in the Ophidia, and to

the Anatomy of the Genera Goa and Corallus. Proc.

Zool. Soc. 1906, p. 499.

(5) Bepparp, F. E.—A Comparison of the Neotropical Species

of Corallus, C. cookii, with C. madagascariensis ; and on

some Points in the Anatomy of Corallus caninus. Proc.

Zool. Soc. 1908, p. 135.

(6) Bepparp, F. E.—A Contribution to the Knowledge of the

Encephalic Arterial System in Sauropsida. Proc. Zool.

Soc. 1905, vol. i. p. 59.

(7) Bemmecen, J. F. van.— Entwikkelung en metamorphose der

kieuw of veseralspalten en der aortenbogen bij Embryonen

van Tropidonctus natria en Lacerta muralis. Kon. Akad.

v. Wet. Amsterdam, Afd. Natuurk., 1885.

(8) Bemmecen, J. F. van.—Die Visceraltaschen und Aortenbogen

bei Reptilien und Voégeln. Zool. Anz., 1886.

(9) Bemmeten, J. F. yvan.—Beitriige zur Kenntnis der Halsgegend

bei Reptilien. 1. Anatomischer Teil. Mededeelingen tot

de Dierkunde. ‘Tijdschrift van het Genootschap Natura

artis magistra te Amsterdam, 1887.

(10) Branpr, F.—Ueber einen eigenthumlichen, spater meist

obliterirenden Ductus Caroticus der gemeinen Kreuzotter

(Pelias berus). Mélange biologique, Bd. v., 1865.

(11) Brenner, A.—Uber das Verhiiltnisse des N. laryngeus

inferior vagi zur einigen Aortenvarietiiten des Menschen

und zu dem Aortensystem der durch Lungen athmenden

Wirbelthiere iiberhaupt. Archiv fiir Anat. und Physiol.,

Anat.-Abt., 1883.

(12) Bruner, H. L.—On the Cephalic Veins and Sinuses of

Reptiles, with a description of a mechanism for raising

the venous blood pressure in the head. American Jour.

of Anat., 1907.

(13) Denpy, A.—The Intracranial Vascular System of Sphenodon.

Phil. Trans., 1909.

(14) De Vriesz, B.—Sur la Signification morphologique des

Arteres cérébrales. Archives de Biologie, xxi., 1905.

(15) Ecker, A.—Der feinere Bau der Nebennieren beim Menschen

und den vier Wirbelthierklassen. Braunschweig, 1846.

(16) Frirscu, G.—Zur vergleichenden Anatomie der Amphibien

herzen. Archiv fiir Anat. und Physiol., 1869.

644 MR. CG. H. O'DONOGHUE ON THE

(17) Gratrorer, P.—Note sur le systéme veineux des Reptiles.

L’Institut xxi., 1853.

(18) Gretu, A.—Beitriige zur vergleichenden’ Anatomie und

Entwicklungsgeschichte des Herzen und des Truncus

arteriosus der Wirbelthiere. Morph. Jahr., Bd. 31, 1903.

(19) GrossEr, O., und Brezina, E.—Ueber die Entwicklung der

Venen des Kopfes und Halzes bei Reptilien. Morph.

Jahr., Bd. 23, 1895.

(20) Hocusterrer, F.— Beitriige zur Kntwicklungsgeschichte der

Amnioten, 2. Reptilien (Zropidonotus, Lacerta). Morph.

Jahr: Bd, VO. Tsos.

(21) Hocusterrer, F.—Ueber Varietiiten der Aortenbogen,

Aorten wur. zeln und der von ihnen entspringenden Ar ter ien

bei Reptilien. Morph. Jahr., Bd. 29, 1902.

(22) Hocusrerrer, F.—Die Entwicklung des Bin taebicksyaten.

In Hertwig’s Handbuch der Entwicklungslehre der Wir-

beltiere, 1901— 06.

(23) HOFFMANN, C. K.—Schlangen und Entwicklungsgeschichte

der Schlangen. Bronn’s Thier-Reich, 1890.

(24) Hormann, M.—Zur vergleichenden Anatomie der Gehirn-

und Riickenmarksarterien der Vertebraten. Zeitschrift

fiir Morph., Bd. ii., 1900.

(25) Hopxinson, J. P., and Pancoat, J.—On the Visceral Anatomy

of the Python (Cuvier) described by Daudin as the Bow

reticulata. ‘Trans. American Phil. Soc., 1837.

(26) Jaquart, H.—Mémoire sur les Organes de la Circulation

chez le serpent Python. Ann. des Sci. Nat., Tome iv.,

1855.

(27) Jourpatn, 8.—Recherches sur la vein Porte Rénale. Ann.

des Sci. Nat., 1859.

(28) Kinestny.—Guides for Vertebrate Dissection. New York,’

1907.

(29) Lancer, A.—Uber die Entwicklungsgeschichte des Bulbus

cordis bei Amphibien und Reptilien. Morph. Jahr., ,

Bd. 21, 1894,

(30) Ratuxe, H.—Entwicklungsgeschichte der Natter. Kdénigs-

berg, 1839.

(31) Rarnxe, H.— Ueber die Carotiden der Schlangen.

Denkschr. Wien. Akad. Wiss, , Math.-nat. K1., Bd. xi. 1856.

(32) Rotueston, G., and JACKSON, W. H.—Forms of Animal

Life, Ond Ed., p. 70. Oxford 1888.

(33) Sasatier, A.—Etudes sur le cceur et la circulation centrale

dans la série des Vertébrés. Ann. des Sci. Nat., Tome 18,

1873.

(34) Saparrer, A.—Observations sur les transformations du

Systeme aortique dans la série des Vertébrés. Ann. des

Sci. Nat., Tome 19, 1874.

(35) Scutemm, F.—Anatomischer Beschreibung des Blutgefiiss-

system der Schlangen. ‘Treviranus’s Zeitschrift fiir

Physiologie, Bd, 3, 1827.

. CIRCULATORY SYSTEM OF THE GRASS-SNAKE, 645

(36) Tanpuer, J.—Mikroskopische Injectionen mit kaltflussiger

Gelatin. Zeitsch. f. wiss. Mikros. v. f. Mikros. Tech.,

1901.

(37) West, G. $.—On the Buccal Glands and Teeth of certain

+ Poisonous Snakes. Proc. Zool. Soe. 1895, p. 812.

(38) West, G. S.—On two little-known Opisthoglyphous Snakes.

Jour. Linn. Soc., Zool. 1896.

VII. EXPLANATION OF PLATES LXX.-LXXII,

Lettering.

A.A.L. Left Aortic Arch. G.A, Gastric Artery,

A.A.R. Right Aortic Arch. Gall. Bl. Gall-bladder.

A.A.V. Anterior Abdominal Vein. H.A. Hepatic Artery.

A.C.A. Anterior Choroid Artery. Har. Gl. Harderian Gland.

A.V. Azygos Vein. H.G.A. Harderian Gland Artery,

A.V.A. Anterior Azygos Vein. H.P.V. Hepatic Portal Vein.

A,V.P. Posterior Azygos Vein. H.P.V.’ The posterior part of the

B.A. Basilar Artery. Hepatic Portal Vein, the

B.A.L. Loop of Basilar Artery. “Veme mésentérique

C.A. Left Common Carotid postérieure”” of Jourdain

Artery. : (27).

Ca. A.

C,C.A

Caudal Artery.

. Cerebral Carotid Artery.

.O. Origin of the Hepatic Portal

Vien trom the Renal Portal

C.C.Ant. Anterior branch of the Vein,

Cerebral Carotid Artery. J.A.A. Internal Auditory Artery.

C.C.M. Median branch of the 1.C.A. Internal Carotid Artery.

Cerebral Carotid Artery. 1.C.M. Muscular branch of In-

C.C.P. Posterior branch of the ternal Carotid Artery.

Cerebral Carotid Artery. I.C.O. @sophageal branch of In-

Cep. V.D. Dorsal Cephalic Vein. ternal Carotid Artery.

Cep. V.L. Longitudinal Cephalic I.M.A. First Inferior Mesenteric

Vein. Artery.

Cer. A. Cervical Artery. I.M.A.2. Second Inferior Mesenteric

Cer. V. Cervical Vein. Artery.

Cer. Hem. Cerebral Hemispheres. 1.M.A.3. Third Inferior Mesenteric

C.W. Circle of Willis. Artery.

C.J.L. Left Common Jugular I.M.A.4. Fourth Inferior Mesenteric

Vein. Artery.

C.J.R. Right Common Jugular Inf. Infundibulum.

Vein. LP.V. Inferior Palpebral Vein.

C.V. Caudal Vein. I.S.A. Inferior Spinal Artery.

C.V.L. Longitudinal Cerebral Kid. L. Lett Kidney.

Vein. Kid. R. Right Kidney.

C.V.M. Median Cerebra! Vein. L.A. Left Auricle.

C,V.P. Posterior Cerebral Vein. L.C.A. Long Ciliary Artery.

C.V.S.A. Secondary Anterior Cere- L.G.A. Lieno-gastric Artery.

bral Vein. L.R.A. Anterior Left Renal

D.A. Dorsal Aorta. Artery.

H.C.A. Anastomosis of the two Ex- L.R.M, Median Left Renal Artery,

ternal Carotid Arteries, L.R.P. Posterior Left Renal

E.C.L. Leff External Carotid Artery.

Artery. M.A. Merle Artery,

E.C.R. Right External Carotid M.C.A, Median Goebel ‘Artery.

Artery. M.C.P. Posterior branch of the

H.M.A. Eye Muscle Artery. Median branch of Cere-

E.R.L. Left Efterent Renal Vein. bral Carotid Artery,

a: E.R.R. Right Efferent Renal Vein. Med. Medulla oblongata,

But. Nar. Exter nal Nares. M.V. Mandibular Vein.

FA. Fat-body Artery. Mx. Maxillary Vein,

¥.C.A. Facial Carotid Artery. Mx.A. The double anastomosis of

F.C.P. Palatine branch of the the Maxillary Vein.

Facial Carotid Artery. O,A,L. Left Ovarian Artery.

646

ON THE CIRCULATORY SYSTEM OF THE GRASS-SNAKE,

O.A.R. Right Ovarian Artery. Pel. V. Vein probably corres-

Oc. A. Occipital Artery. ponding to the Pelvic

Od. A. Oviducal Artery. Vein ot Lacertilia.

Od. A.P. Posterior Oviducal Artery. Pit. Fos. Pituitary Fossa.

Od.S.L. Lett Oviducal Sinus. Pl.C.V. Palato-cerebral Vein.

Od.S.R. Right Oviducal Sinus. Pl. P.V. Palato-pterygoid Vein.

Od. V.L. Lett Oviducal Vein. Pul. A. Pulmonary Artery.

Od. V.L.P. Left Posterior Oviducal Pul. V. Pulmonary Vein.

Vein. P.V. Parietal Vein.

Od. V.R. Right Oviducal Vein. R.A. Right Auricle.

Od. V.R.R. Right Posterior Oviducal Ret. A. Retinal Artery.

Vein. R.P.L. Left Renal Portal Vein.

Oe. A. Gsophageal Artery. R.P.R. Right Renal Portal Vein.

Oe. V. Csophageal Vein. R.R.A. Anterior Right Renal

Ol. A. Olfactory Artery. Artery.

O/f. lo. Olfactory Lobe. R.BR.M. Median Right Renal

Op. A. Ophthalmic Artery. Artery.

Opt.lo. Optie Lobe. R.R.P. Posterior Right Renal

Opt. Ner. Optic Nerve. Artery.

O.P.V. Oblique Palatine Vein. Rt. A. Rectal Artery.

Orb. Orbit. R.V. Rostral Vein.

O.S. Orbital Sinus. S.C.A. Short Ciliary Artery.

O.S.A. Anastomosis between the S.M.A. Superior Mesenteric

Orbital Sinus and the Artery.

Maxillary Vein. S.N.A. Anastomosis of the Sub-

Oy. A. Smaller Ovarian Artery. nasal Sinuses.

Ovar. L. Left Ovary. S.N.V. Sub-nasal Vein.

Ovar. R. Right Ovary. Sp. A. First Spinal Artery.

P.A. Parietal Artery. Sp. V. Spinal Vein.

Pan. Pancreas. Spin. Cor. Spinal Cord.

Par. Parotid Gland. Spl. Spleen.

Par. A. Anastomosis between a S.P.L. Lett Supra-renal Portal

branch of the Anterior Vein.

and a branch of the S.P.R. Right Supra-renal Portal

Posterior Parotid Gland Vein.

Arteries. S.R.L. Left Supra-renal Vein.

Par. A.A. Anterior Parotid Gland S.R.R. Right Supra-renal Vein.

Artery. Sup. Lab. Gl. Superior Labial Gland.

Par. P.A. Posterior Parotid Gland Sup. Ren. L. Lett Supra-renal body.

and Superior Maxillary | Sup, Ren. R. Right Supra-renal body.

Artery. Th. A. Thyroid Artery.

Par. V. Vein from Parotid Gland Thy. Gl. Thyroid Gland.

and also from neigh- Ton. Mus. Tongue Muscle.

bouring muscles. Ton. Sh. Tongue Sheath.

P.C. Primary Carotid Artery. V. Ventricle.

P.C.V. Post-Caval Vein. V.A. Vertebral Artery.

PuatE LXX.

Fig. 1. Diagram of the Heart and Blood-vesseis in the anterior part of Tropidonotus

natrix. ‘The two main arteries and veins on the ventral side of the head

are indicated, but their precise distribution is dealt with on Plate LX XII.

Between the lines X.X. and Y.Y.a portion of the circulatory system is

omitted, as the relations of the Hepatic and Pulmonary Vessels and the

Dorsal Aorta are similar throughout the whole length of the liver. The

part omitted is about the same length as the distance from the thyroid

gland to the line X.X. For the purposes of diagram the liver is repre-

sented as pulled out to the right of the animal. ‘The positions of the Liver

and Lungs are indicated by simple outline and those of the Gall-bladder,

Spleen, and Pancreas by shaded areas.

Fig. 2. Diagram of the posterior Blood-vessels in a female Tropidonotus natria.

The positions of the kidneys and ovaries are indicated by simple outline

and those of the supra-renal bodies by shaded areas. For the purposes of

diagram the intestine is represented as pulled out to the right of the

animal, the fat-bodies as external to the oviducts, and the oviducts as

external to the ureters. Owing to the fact that the anterior end of the

Fat-body is supplied from theright ovarian artery, while a little further

ON THE COURTSHIP OF THE REDSHANK. 647

back it is fed from the left ovarian artery, it is necessary to divide the

Fat-body between the two gonads. Thus it comes that the anterior part

of the Fat-body is represented on the right side of the animal and the

posterior part on the left. As the Anterior Abdominal Vein runs in the

Fat-body it is also divided and is so represented in the diagram, in which

its two ends are joined by a dotted line.

Prate LXXI.

Diagrams of the Blood-vessels in the head of Tropidonotus natrix.

Fig. 3. Diagram of the Cranial Arteries seen on the ventral surface of the brain.

The brain is represented as removed from the skull leaving behind,

however, the Pituitary Body.

Fig. 4. Diagram of the arteries of the head seen from the dorsal side. The brain is

removed, but some of the cranial arteries are left behind. The positions of

the glands of the head are indicated by shaded areas.

Prate LXXII.

Fig. 5. Diagram of the vessels on the ventral side of the lower jaw after the removal

of the superficial muscles.

Fig. 6. Diagram of the Veins and Sinuses in the head seen from the dorsal side.

The deeper vessels are represented in lighter shading and with a dotted

outline. The position of the brain is indicated in simple outline, while the

positions of the glands are indicated by shaded areas. For the purposes of

diagram the Superior Palpebral Vein is omitted. It runs from the upper

anterior part ot the Orbital Sinus above the Maxillary Vein and joins the

Interior Palpebral Vein as the latter enters the Orbital Sinus. In dis-

secting out the vessels of the orbit it is almost always removed with the

upper eyelid.

34. A First Account of the Courtship of the Redshank

(Votanus calidris L.). By Juytan 8. Huxuey, Lecturer

of Balliol College, Oxford.

[Received February 2, 1912: Read April 23, 1912. ]

INDEX.

Page

AbpInitroduetvone eee ee a eta s Ue eastern, Ok Sek Rn ST

DREW OCAi tiyuy sseteeo rae een an ahi Rahat. WNT OR. ne ON BAR

Sumlres Courtship pro periater steel ac. oak te peices scl. cee 649

4. Other habits of the Pairing-Season :

(@) pen owe site ih ae aes ered wee ei os gals NE G5ilh

(6) The Combats of the Males .................. ee ea 652

(ec) Calling from a conspicuous perch ................ 652

Sse DISCUSSIONYS rare t eave seme epee tak 8 Cae eae (GOO

1. InrrRopuctTion.

While staying last spring in a lonely corner of North Wales

it was my good fortune to come across a number of rare and

interesting birds. But great as was the pleasure of seeing, for

the first time, such comparatively uncommon species as the Gre

Plover and Black-tailed Godwit, it was far surpassed by that of

being able to study, under the most favourable conditions, the

natural behaviour and home life of some of the commoner shore-

birds. Of these I was particularly fortunate with the Redshank,

648 MR, J, 8. HUXLEY ON TUE

and was able to see the whole of the courtship and pairing.

When I say the whole, I do not mean that I saw every detail,

nor that every detail I saw is clear to me, But I mean that

I know what is the general course of events, and can interpret

the birds’ behaviour more or less consistently.

On returning to civilization and libraries, to my surprise I

could find very little on the subject: the observations recorded

were either fragmentary or inaccurate. It was not until most

of this paper was written that I discovered a fairly e complete

account by Selous *, This had remained undiscovered owing to

the absence of any reference to Redshanks in the title of the

paper or in the index of the volume.

I have thought it worth while to publish my observations,

however, since they differ in several points from those of Selous.

Meanwhile, I fully realize their incompleteness, and recognize

that they cannot as yet be properly used in any general discussion

of the theory of sexual selection. I hope to continue my own

observations when opportunity offers, but venture to publish

this general outline at once as a stimulus to other bird-watchers

and naturalists.

Locartry, ete,

Before passing on to the birds’ actions I must first just

mention the theatre where I saw them played. This was part of

a.small estuary in the northern half of Cardigan Bay: an arm

runs out on one side at right angles to the river, thus giving

during high spring tides a land-locked sheet of water nearly <

mile long and half a mile wide; during neaps, even high tide

failed to cover it. Numbers of Redshanks and other birds fre-

quented this expanse, and especially its head or most landward

end, where they were close to a thick bed of reeds and tussocks ;

the mud here was scarcely ever covered by the tide, though kept

always moist by a little stream.

At one side of the head. was a low ridge of grass-covered

dunes, about five feet high and thirty or forty feet long, with level

ground behind them. Thus, by crawling over the flat, I could

get up to the dunes into an excellent position for viewing the

whole top of the bay ; every bird was easily seen against the wet

mud. Sol kept watch with the naked eye until some disturb-

ance or unusual behaviour attracted attention ; then, being armed

with a telescope magnifying 30 diameters (for the loan of which

I have to thank my brother, Mr. N. T. Huxley), I focussed this

on the spot, and could see the minutest details of attitude and

behaviour in the nearer birds, and even in those on the far side

of the bay could quite well interpret what I saw. I made a

number of notes on the spot, and usually within twenty-four

hours embodied what I had seen the day before in a letter to

an ornithological friend.

* BE. Selous, “ Obs yes tending to throw light on the Question of Sexual

Selection in Birds,” ete., Part 1; Zoologist (4) x., 1906, pp."201-218.

COURTSHIP OF THE REDSHANK 649

3. THE COURTSHIP PROPER.

I will begin with an account of the typical course of the

courtship and pairing, such as I have seen repeated, with but

slight variations, a considerable number of times.

Among the forty or fifty birds that usually would be quietly

feeding on the flats, walking or running in short starts from

mouthful to mouthful, a disturbance would every now and then

be visible—two birds running, one pursuing the other. ‘These

two are cock and hen. A cock takes a fancy to one of the

hens, leaves his feeding, and starts running towards her. She at

once runs away from him, and there ensues a regular game of

follow-my-leader. The hen never goes far in a straight line;

she usually runs in a series of curves, often doubling sharply

back, and sometimes describing a complete circle or even a figure

of eight. Where she goes the cock goes after her, following

exactly, but some yards behind. The couple would be ridieulous

enough in their devious course, with heads somewhat down and

quick-moving legs—rather the action of a fast-trotting horse—

but the attitude of the cock adds to the effect: his eyes being set

on the sides of his head, his neck has to be stretched stiff out and

markedly sideways (at an angle of at least twenty degrees with the

line of his body), in order to keep the hen in sight. In addition

he spreads his pure white tail, so that you see half the fan of it on

either side of the tips of the folded wings; but whether the hen

so far ahead can see anything of this I do not know.

This pursuit goes on often for quite a long time, the birds

covering maybe a quarter of a mile. The hen usually flies away,

leaving the cock disconsolate, but sometimes she will consent to

inaugurate the second stage of the courtship, in which she is

able to inspect the suitor more closely. This she does by suddenly

coming toa dead stop. The cock then perhaps runs a yard or

two further, but soon he too stops, and begins his part of the

second stage of the courtship.

The first stage was almost mere pursuit: the second is pure

display. He first unfolds his wings and raises them right above

his back, so as to expose their conspicuous under-surface of pure

white somewhat clouded or barred with grey. Then, fluttering

them tremulously, but keeping them raised all the time, he

advances very, very slowly towards the hen, lifting his feet high

in the air, and often putting them down scarcely in advance of

where they were before. To the human eye the whole action

seems the expression of eager excitement tempered by uncer-

tainty, and that, presumabiy, is what the bird is actually feeling.

Meanwhile, as he steps on, he stretches his neck a little forward,

opens his mouth, and gives utterance to a single continuous note,

which is changed into a long roll or rattle by the quick vibration

of the lower mandible. ‘The sound is quite like that of a Night-

jar, but higher, and without any of the little breaks in the pitch

of the note. So he advances closer and closer, the hen usually

Proc. Zoou, Soc.—1912, No. XLII. 43

650 MR. J. 8. HUXLEY ON THE

remaining motionless. Again at any time during this stage she

may reject his suit by flying off, but if she is going to accept him,

she simply stays still, often without moving a muscle the whole

time.

As the cock gets closer, he gets more and more excited, vibrates

his wings more and more rapidly, at length so fast that almost

his whole weight is supported by them, though he still continues

to execute the high-stepping movements with his feet. At last,

when just behind the hen, he abandons the ground, and flutters up

on to her back, on which he half alights. The period while he

is thus on her back is the third and last stage of the courtship:

it is very short, and is of course in a sense nothing more than

getting into the proper position for the actual pairing. But it

should still be called part of the courtship, for even now the cock

is not assured of his desire. Sometimes the hen, suddenly repug-

nant, or annoyed by the series of shriller, less continuous cries

which the cock is now uttering, gives a violent jerk or sideways

twist, and shakes him forcibly off on to the ground, herself

running or flying away.

Occasionally, however, she apparently is satisfied; she spreads

her tail diagonally, and the cock, with a quick and wonderfully

graceful motion, half supported all the time by his fluttering

wings, accomplishes the act of pairing. ‘Then the hen gives

the same violent twist that I have just mentioned, he gets shaken

off, and they both begin quietly feeding, often side by side, but

now no longer taking the least interest in each other,

That the course of true love should run so smooth was, how-

ever, quite the exception. Though I did not keep a record of

the number of unsuccessful cocks that came under my observa-

tion, there must have been at the lowest estimate fifty of them,

while but three times did I see the courtship consummated.

Thus in something well over 90 per cent. of the cases I saw,

pairing did not take place; and this was always due to the rejec-

tion of the male by the female, as the cock, once he had started a

pursuit, never of his own accord abandoned it. Thus, though the

hen does not actively select her mate from among a bevy of

competing cocks, yet, like the modern European woman, she has

the power of saying yes or no to each individual male who

may choose (here literally, there metaphorically) to run after

her.

The hen may reject her suitor at any time during the whole

proceeding. On the one hand, I have seen her break off the

courtship after a few seconds, while on the other I have seen her

stop still after running, stand and watch the cock’s display

and gradual approach, even let him fly up on to her back, and

only then with a sudden jerk throw him off and take to flight.

Between these extremes there were, of course, all intermediates.

During the first or pursuit stage, rejection was accomplished by

the hen simply taking wing and flying off some fifty or a hundred

COURTSHIP OF THE REDSHANK. 651

yards. The cock sometimes (though not usually) flew after her,

settled, and began his running once more; but then she would

very soon fly off again, and I never saw a cock persevere after a

second such repulse. ‘The first stage may be very short, as I

have said, owing to her rejecting her suitor at once; and it may

be short for the reverse reason, the hen stopping almost at once

to let the cock come up for inspection. Usually, however, it is of

considerable length, and in well over half the number of cases 16

ends in the rejection of the cock and the cutting short of the

courtship before the second stage is reached.

Of the minority who survived thus far, a still larger percentage

were rejected before getting to the third stage. This was most

often accomplished by the hen simply flying off, leaving her

suitor to fold his wings and pretend nothing had happened.

Sometimes, however, if more undecided, she would behave ina

curiously human way. As the cock got close she would as it

were lean away from him, and at last, giving a little quiver all

over, break into a short quick run of about four or five steps,

like a frightened horse shying across the road. Then she stood

still again, and, when the cock advanced again, very likely repeated

the action. Of the five or six hens I saw act thus, all as a matter

of fact at length flew off; but I have no doubt that occasionally

they make up their minds in the opposite way.

4. OTHER HABITS OF THE PAIRING-SEASON.

There are other habits of the pairing-season which call for

special remark. J do not understand their relations to the court-

ship proper, and shall merely mention them here. They are

introduced partly for the sake of completeness and partly to show

what a number of unsolved problems still exist concerning the

habits of common birds.

(a) The Love-flight.—This is a well-known habit. A Redshank

(presumably a cock) rises up into the air and there flies in a

series of switchbacks. JI will quote from my own notes on a

particular bird: “Just before the bottom of each switchback he

gave very quick wing-flaps, almost fluttering, one would call it ;

this made him start up again. He went on fluttering or flapping

till he was about half-way up, and for the rest of the up-stroke

of the switchback he soared up with the impetus he had gained.

His wings now were set back and down, his neck and head

thrown up in a beautiful proud attitude, his tail spread out.

Then he turned the angle of his wings and glided down, still in

the same attitude.” While flying thus, he gives vent to what

one may well call a song—a series of pure sweet single notes,

never uttered on other occasions. The flight may be quite short,

or may go on and on for several minutes. It is usually, I think,

gone through by single birds, but I have fairly often seen it done

43%

652 "MR. J. S. HUXLEY ON THE

by one of two or one of three birds. If so, however, it was

hardly ever repeated more than a very few times. I have several

times seen it take place when I have frightened a bird up trom

feeding.

The meaning of this habit is hard to discover, and its relation

(if any) to the courtship proper is equally obscure. It is per-

formed, I believe, only in the spring, and would certainly seem

to be of the same nature as the drumming of the Snipe or the

short soaring flight of the Wood-Pigeon.

(b) The Combats of the Males.—These have been well described

by Selous (loc. cit.). I agree with his opinion that the combats

of many birds are now at least merely formal. This was well

shown in the Redshank; the birds scarcely ever touched each

other, and often did not even seem fierce. It was mostly a mere

running up and down of two birds facing each other, often with

a very formal-looking character about it. Further observation

alone can reveal its significance.

various writers, and is a very noticeable custom. A single bird

will settle on a gate-post, railing, notice-board, or other prominent

perch, and will sit there, moving its head from side to side, and

uttering a single note many times repeated. Usually after every

few notes there comes a short pause. Long pausesarerare. One

bird continued calling thus from the same perch for 45 minutes,

and was still going on when I had to stop watching. I have at

present no idea as to the sex of the calling bird, or as to the

purpose of the whole proceeding.

5. Discussion.

T will confine myself in this paper to a short discussion of the

courtship proper, and will begin by comparing my observations

with those of Selous (oc. cit.); all other descriptions can here be

left out of account, owing to their vague and fragmentary

character.

Selous, who made his observations in Holland, seems, to start

with, to have seen a greater proportion of successful courtships

than I did. His description (often hard to follow, as there is no

arrangement—only notes in chronological order) differs in the

following chief points from mine :—

1. He seems to have seen only two examples of the first stage

‘of the complete courtship (pp. 212, 213), which is curious, since

the follow-my-leader evolutions of my birds were most con-

spicuous.

2. His first stage is described as follows :—‘ The male, ap-

proaching the female, ran about her twice or thrice, in so many

half-circles, fanning his tail as he did so, and inclining his body

towards her.” .... That is to say, the~hen was stationary,

COURTSHIP OF THE REDSHANK. 653

instead of leading the cock a long chase, as in the Welsh

birds.

3. Sometimes directly after pairing there was a curious set

run or antic of the couple, e.g. (p. 206) :—‘‘Two Redshanks,

after pairing, run, in an excited and curious-looking manner,

over the sand, following one another.” Another time (p. 204),

(in perhaps slightly different circumstances) :—“ the birds walked,

for a little, about and very near each other, fanning out their

tails, whilst bending them inwards, so that, had their Jegs been

short, they would have swept the ground at intervals, as does

that of the courting pigeon.”

4. He sometimes saw a male which had been definitely rejected

begin courting the same female again after a short time, once or

twice successfully ; I never sawthishappen. (This is presumably

an error of omission on my part.)

These are the chief differences. His observations on the

second and third stages and on the pairing itself are very similar

to mine.

The differences may be ascribed either to real differences of

behaviour of Zotanus calidris in different parts of its range, or

to errors or omissions of observation. Further watching alone

will reveal the truth.

Further watching too must elucidate the following points, all

of which ought to be known before a full discussion of the facts

and of their bearing on the theory of sexual selection can take

place :—

1. How often does each bird go through the act of pairing ?

2. Is pairing promiscuous, or do birds pair for the season, or

for life ¢

3. What is the relation between pairing and nest-building,

and between pairing and each act of oviposition ?

4. When does pairing begin in the spring, and for how long

is it continued 2

5. What is the relation of the love-flight, the combats, and

the calling from a perch, to the courtship proper ?

6. Does the female, who possesses all the structures used by

the male in his display, ever use them herself for ‘showing

off,” or for any other purpose (recognition signals, etc.) ?

With regard to 1, it is interesting to note that when the air

was calm the Nightjar-like note described above (p. 649) could

be heard at every hour of the night and day; on some nights

there were one or more birds giving utterance to‘it practically

all the time that I was listening. Now I am prepared to assert

that this call is only used during the second stage of the court-

ship, so that the number of courtships which advanced as far

as the second stage must have been very great (yet less than the

number of those which never reached the second stage at all;

cf. p. 651.)

654 MR. J. S. HUXLEY ON THE

The number of Redshanks near the head of the estuary (where

alone they really congregated) was never very large. I should

put fifty as an outside limit, and from observations on this and

other species (e. g. Oyster-catchers), I believe that they are the

same individual birds day after day. Therefore, even allowing

that only a fraction of the courtships which reached the second

stage were consummated, yet the number of acts of pairing must

have been many times greater than the number of birds; pro-

kably, therefore, each bird pairs several times a day.

Tt is thus likely, as Selous says, that the performance of the

courtship will vary very much in different circumstances ; there

may be satiety or eagerness in either sex, as well as timidity and

shyness in the female, at different times. Selous adduces the

Pheasant to prove that the display of the male may be gradually

much abbreviated and scamped after the first few pairings.

As regards 4, 1 have very meagre evidence. I can only say

that pairing had begun before the 8th of April, and was still

going on very vigorously when I left Wales on the 18th. All

the other questions I must leave for the present unanswered.

Selous’ and my observations, however, are in themselves enough

to establish one important point, namely, that the actions of the

birds which lead up to each single act of pairing are explicable

only on the Darwinian theory of Sexual Selection, or on some

modification of that theory.

On the one hand, there is a very marked display by the male: the

fanning of the tail in the first two stages, the lifting and flutter-

ing of the wings, and the high-stepping with the legs in the

second stage, are all obviously calculated to show off to the best

advantage certain conspicuous markings which are usually con-

cealed, while the rattling note of the second stage is to my mind

equally to be considered as an excitant.

On the other hand, there is an equally marked power of choice

shown by the female; it is perfectly clear that if a female Red-

shank does not want to pair with any given male, he has no

possible means of forcing her to do so. He can only persuade

her, or rather attempt to persuade her (by means of his display),

and if she is very unfavourable to his suit, she can even prevent

him from doing this, by flying off directly he begins.

Thus, though the male in this particular species has the

initiative, the final decision must rest with the female.

The chief postulate of Darwinian Sexual Selection therefore

holds good in the case of the Redshank:—The females have a

power of choice, and the cocks have to go through a display

before pairing. Moreover, the one certainly seems to stand ina

causal relation to the other.

It is interesting to note, however, that although Sexual

Selection is at work, yet it has not produced any appreciable

difference between the sexes. This can be explained in one of

three ways.

COURTSHIP OF THE REDSHANK. 655

(1) The markings used in display (red legs, white tail, and

whitish under surface of wings) were acquired previously by the

species for some distinct purpose (e.g. as recognition or oblitera-

tion markings) or possibly accidentally, and then were turned to

account by the cock as the ‘physical basis” of his display. The

hen would of course possess the markings too; she differs from

the male in not having secondarily acquired the instinct to display

them.

(2) The markings were acquired by the cock first or primarily

as secondary sexual characters, to form the basis of his display,

but were either at once or later found to be of use in other ways:

they would then tend to be transferred to the hen as well, either

from their first beginnings, or subsequently to their definitive

development in the cock, by the operation of Natural Selection.

(3) The markings were acquired for purposes of display, while

in other respects they are neither harmful nor the reverse; the

instinct to use them for display, however, depends on a physio-

logical stimulus only present in the male sex. Then they would

tend to be transferred to the female sex, for we generally find

that the two sexes resemble each other unless there is some

definite reason for their differing. It appears to be both more

primitive and easier for hereditary characters to be transmitted

equally to both sexes. ;

It is at present very hard to decide between these possibilities.

Of the three, the last appears the least probable. What does

emerge clearly, however, is that in considering the facts of Sexual

Selection, as so often elsewhere, we must be careful not to isolate

structure from function. When we speak of secondary sexual

characters, we usually think of structures only. In reality the real

character is the structure plus the instinct to use the structure,

for it is the use of the structure which alone has any significance

for the species: it is that which constitutes a unity, it is that

which has been really acquired by the species. For purposes of

convenience we separate it into two components—structure and

function; but in any question of its origin and history we must

always be careful to think of it asa whole. To take a concrete

example: if it were proved (as is probable) that the female Red-

shank never used her white tail, etc. for purposes of display, we

should be justified in saying that the Redshank showed secondary

sexual characters—these characters being the various actions of

display found in the male, and in the male alone,

This point of view will perhaps help to make more intelligible

the various cases which have been described where the sexes are

alike in plumage, but the male alone goes through a display.

656 MRS. E. W. SEXTON ON

35. Some Brackish-water Amphipeda from the mouths of the

Weser and the Elbe, and from the Baltic*. By E. W.

Sexton, Marine Biological Laboratory, Plymouthf.

[Received March 1, 1912: Read April 23, 1912.]

(Plates LX XIII. & LXXIV.+)

INDEX.

Page

ViaiatlOnlpee teehee reer peeemene can asus asreepeeanaatsu aut: LODO

Systematic: New Species under Gammarus ......... 657

While engaged in determining the Copepoda of the harbour at:

Bremerhaven, Herr W. Klie met with certain Gammaridea, some

of which were not easily referable to known forms. Through the

intervention of a friend these were submitted to me for examina-

tion, and the present paper contains the result, with notes on

some other collections of brackish-water Gammarus. There

proved to be three species in Herr Klie’s collection, one of

them (Leptocheirus pilosus Zaddach) already known, the other

two apparently new, which I described. But before the descrip-

tion was published I received a paper from Dr. Vanhéffen, which

contained the description and figures of one of these species

under the name of Corophiwm lacustre. I have therefore in-

cluded here only a few notes on this species. Dr. Vanhoffen

kindly allowed me to examine his specimens of Amphipoda from

the Frische Haff, and it is interesting to note that the three

species, Leptocheirus pilosus Zadd., Corophiwm lacustre Vanhoften,

and Gammarus zaddachi, sp. n., are present in both his and Herr

Klie’s collections. All three appear to flourish equally well in

absolutely fresh water as in the brackish water of river estuaries

and harbour basins.

The classification of the Amphipoda is rendered exceedingly

difficult by the changes or modifications resulting from each

successive moult. We know practically nothing yet of the

factors influencing the development of any given species, except

those of growth and sex. The modifications caused by these two

alone are responsible for an excessive multiplication of synonyms;

in some cases, Jassa for instance, almost every moult has been

given a different specific and sometimes a different generic

name by different observers. But in the species of Gammarus

* (Since this paper was communicated to the Society, under the title “Some

Amphipoda from Bremerhaven,” the author received some large collections of the

Gammarus described, which showed remarkable differences between those living in

salt and in fresh water. These facts have been included in the text, and the title

has been slightly modified. ]|—Ep. P. Z. §

+ Communicated by the Rev. T. R. R. Sreserne, M.A., F.R.S., F.Z.S.

{ For explanation of the Plates see p. 665.

Pi. ole, Saeed,

a. /\ he

E,W.Sexton del, Huth se. et imp.

AMPHIPODA FROM BREMERHAVEN.

Ei es OIG

Huth se. et imp.

AMPHIPODA FROM BREMERHAVEN.

E.W.Sexton del,

SOME BRACKISH-WATER AMPHIPODA. 657

described below, certain marked modifications occur which cannot

be referred to either of these influences. They would appear

to be caused by the animal’s environment, according as it lives

in fresh water or in salt. The difference in appearance between

a typical adult freshwater specimen and a typical marine or

brackish-water one is so extreme as to suggest their belonging

to distinct species, but structurally they are identical, and

intermediate forms are very common (see below, “ Moorflether

Concave”), apparently varying with the degree of salinity, though

on this point I cannot as yet speak definitely. Experiments

have been instituted at the Laboratory here with an allied species

to try and determine the question of the effect of salinity on

the animal and the length of time for such effect to become

evident.

GAMMARUS ZADDACHI, sp. n. (Pls. LXXIII. & LXXIV.

figs. 1-12.)

= 1844, Gammarus locusta Fabr. ?, Zaddach, Syn. Crust. Pruss.

: =1878. Gammarus locusta Fabr., Zaddach, Die Meeres Fauna

’ an der preussischen Kiiste, pp. 26-32.

= 1886. Gammarus pulex Kraepelin, Die Fauna der Hamburger

Wasserleitung. Abhand. Geb. d. Naturw. Verein in Hamburg,

130 rere sae

=1907. Gammarus pulex Volk, Mitteil. biol. Elbeuntersuch-

ung. Naturh. Museums in Hamburg. Verhandl. Naturw. Ver.

Hamburg.

=1911. Gammarus locusta L., Vanhoffen, Beitrige z. Kennt.

d. Brackwasserfauna im Frischen Haff. Sitzung. d. Gevxellsch.

naturf. Freunde, Berlin, 1911, no. 9.

It is with reluctance that I have felt myself obliged to institute

a new species in the already overcrowded and confused genus

Gammarus. The species of Gammarus are difficult to separate

except by the secondary sexual characters of the adult males;

the immature of all the species are practically indistinguishable

from each other, and even the females are not easy to differen-

tiate. The species now under discussion has been frequently

confounded with others, the freshwater form with G. pulex, and

the marine and brackish-water form with G. locusta and G. due-

benii; but it can be distinguished from them by a glance at the

antenne, both of which are characterised by clusters of long out-

standing hairs, and by the form of the 4th sideplate and the

3rd uropod.

The brackish-water specimens are characterised by their slender-

ness and transparency, and the tenuity of their epidermis, while

the freshwater ones are broad, very robustly built, the epidermis

thick, strong and opaque, the basal joints of the hinder perzo-

pods narrower especially in the old males, and with a much

denser supply of the long fine hairs developed on the antenne,

the perxopods, particularly the hinder ones, the pleon, the 3rd

658 MRS. E. W. SEXTON ON

uropods, and the telson ; in some cases these long hairs appear to

replace the spines of the brackish-water form, e.g. those inset on

the inferior margins of the 2nd and 3rd pleon epimera and the

posterior margins of the 4th and 5th perzopods.

The finest brackish-water specimens I have seen come from

the Konigsberg Collection, taken at Rauschen and Zoppot, the

largest freshwater ones from the Hamburg “ Wasserleitung.”

These are larger than any of the others examined by me, probably

owing to the more protected life they lead in the underground

pipes, and the rich and constant supply of food.

The collections I have examined are as follows, with the names

of those to whom I am greatly indebted for the opportunity of so

doing :—

1. Konigshberg Museum: Dr. Lithe. - Four tubes containing

Zaddach’s specimens and others from Danzig Bucht, Zoppot,

Koliebk, Redlan, and Rauschen, in all about 125. Most of the

specimens are of the typical brackish-water form, but in one

sample from Danzig in which a specimen of the freshwater

G. pulex was also found, many of the animals were “intermediate”

forms, more solidly built and with the long fine hairs developed

in great numbers.

2. Hrisches Haff: Dr. Vanhoffen. Two tubes labelled ‘‘ Gam-

marus locusta L. Frisches Haff bei Pillau Ostpreussen. 20.1x.1911.

Vanhoffen. 5.” One tube contained 16 specimens, 2-5-5 mm. in

length. It is interesting to note a few young Leptocheirus

pilosus with the young G. zaddachi; these two species, with

Corophium volutator and C. lacustre, seem to flourish equally well

in fresh or brackish water and are frequently recorded together.

There were 8 specimens of 4. zaddachi in the second tube, 3°5-

10 mm. long, all of the brackish-water form ; the largest, however,

approaching the freshwater form, strongly built and opaque.

3. Mouth of the Oder: Dr. Vanhéffen. One tube, ‘Gammarus

locusta L. Dievenow Odermundung Pommern 1889. Hilgen-

dorf. 5.” 26 specimens, 1 young male G. locusta and 25 G. zad-

dachi exactly like those described from Pillau.

4. Mouth of the Elbe, from Schulau to Gauert, together with a

collection from the Hamburg Wasserleitung; Dr. Steinhaus. This

is a most interesting series taken at regular stations in the Elbe

(see Volk, with map of the district), containing in all about 500

specimens. Commencing with the stations nearest the mouth of

the river, where the water has nearly the full salinity of the

North Sea, the details are as follows :—

‘“‘Siider-Elbe siidl. Blankanese. Grund. 11.vi.00.” 11. speci-

mens 4—10 mm.

‘““Hafen siidl. Nienstedten. Grund. 2.vi.00.” 35 specimens

15-10 mm.

‘“* Hafen sudl. Nienstedten. 11.vi.00.” 5 specimens, largest a

female 11 mm.

“ Tinkes Elb-Ufer sid]. Nierstedten. 2.1x.00.” 3 specimens,

largest a female 8 mm.

SOME BRACKISH-WATER AMPHTPODA. 659

* Altona. Ponton. 9.vii.00.” 107 specimens, all rather small,

averaging 8-9 mm. in length; the eyes in these have reddish-

brown pigment.

“« Altona. Due d’Alben. 1.x.00.” 1 specimen, a female.

All these belong indubitably to the brackish-water type. The

very young animals differ slightly from the adult, the eyes are

round, the peduncles of the antenne are about equal in length,

the pleon clusters are represented each by a single spine, and the

rami of the third uropod are much more unequal than in the

adult. Some of the specimens, especially those from Blankanese,

are more setose than the others.

“Hamburg.” The “ Fleet” referred to is one of the canals

which ran through the inner town. These canals are mostly

dependent on the tide, with mud-banks exposed at low tide.

‘Grasbrokhafen ” and ‘“ Indiahafen ” are blind alleys as it were

opening out of the Harbour basin ; they are deep and broad, the

bottom consisting of clay, sand and mud, with organic detritus

deposits providing food for a rich animal life.

“Fleet 1. Grund. 25.vi.00.” 1 specimen, a male 11 mm.

‘Fleet 1. Krotzprobe. 23.vii.00.” 1 specimen, a male, with

exceptionally large eyes.

““Grasbrookhafen. Grund. 19.xi.00.” 2 specimens 19-20 mm.

“Grasbrookhafen. Duc d’Alben. 20.vi.00.” 18 specimens

mostly small, the largest 9-10 mm.

“Tndiahafen. Ponton. Oberfliche. 22.v.00.” 77 specimens, all

very young.

The specimens from the “ Fleet” samples show intermediate

characters, forming the link between the two extremes of the

species. They are more solid in appearance than the brackish-

water form, but the chitin is not as thick as in the freshwater

animal. The spines and long hairs vary greatly in number, and

the narrowing of the basal joints of the hinder perzopods also

appears variable. Of the “ Grasbrookhafen” specimens the two

from the bottom are typical freshwater ones; the eighteen spe-

cimens from the “ Duc d’Alben” (piles driven in and used for

making ships fast) are all delicate in appearance, the largest

showing intermediate characters. Some of the small ones, 3-4 mm.

long, are extremely slender and compressed, almost thread-like,

and with very few hairs. The “ Indiahafen” specimens are all

very small, the brackish-water form.

“Alte Dove Elbe and Moorflether Concave.” Volk says of

these that, taken together, they form a quiet bay with the bio-

logical characters of a backwater (“ Altwasser”), but influenced

to some degree by the tide. These samples are the most interest-

ing of the series ; in some both the fresh- and the brackish-water

forms occur together, with the “intermediate” form.

“ Moorflether Concave. Grund I. 23.vii.00.” 1 specimen, a

female 9 mm., with the delicate appearance of the brackish-water

form, but with dense clusters of long hairs on antenne, pereeo-

pods, and telson,

660 MRS. E. W. SEXTON ON

‘¢Moorflether Concave. Oberfliche II. 23.vii.00.” 11 speci-

mens; the largest, males, measured 10-11 mm.; all the brackish-

water form.

“‘ Moorflether Concave. 23.viii.00.” 129 specimens; largest

15 mm. Freshwater, brackish, and “intermediate” forms, and

one specimen of the exceedingly narrow, thread-like form described

from the Grasbrookhafen sample.

“© Moorflether Concave. Ponton. 23.viil.00.” 5 specimens, the

largest, a male of 14 mm., shows the freshwater characters ; the

others are of the brackish-water type.

“‘Moorflether Concave. 24.ix.00.” 37 specimens, brackish-

water form, largest 12 mm.

The three following samples are all unmistakable brackish-

water form. The “ Prielen” are little channels cut in the banks

of the river; the current is less strong, and the growth of vege-

tation and animal life richer than in the Elbe.

“Dove Elbe. Oberfl. 2.vii.00.” 6 specimens, largest about

8 mm.

“T, Pr. O. 7.vi.00” (=Rechtsseitiger Elbpriel am Spaden-

land, Oberfliche). 6 specimens, largest 10 mm.

“7. P.r. O. 2.vii.00.” 14 specimens, largest 8 mm.

The remaining samples are all fine examples of the typical

freshwater form.

“ Kaltenhofe. ‘Wasserprobe aus Filter No. 16 entnommen

am 6. Dec. 1894.’” 3 specimens, the largest 22 mm.

‘¢ Hamburger Wasserleitung.” 5 specimens, 18-20 mm. long.

“Hamburger Wasserleitung. Samuelson.” 71 specimens.

Dr. Kraepelin has given a very interesting account of the condi-

tions of animal life in the underground “ Wasserleitung ” of Ham-

burg, conditions much more favourable than in the Elbe, owing

to the greater abundance of food, the protection from many

enemies, and the lesser temperature variations. He mentions

this species under the name of G'. pulea as second only in numbers

to Asellus aquaticus among the Edriophthalma met with in the.

series of samples taken.

5. Bremerhaven: Herr Klie. One tube “ Alter Hafen. x.11.”

10 specimens, the largest a female 9 mm., from which several of

the figures are drawn. ‘This sample contains one of the thread-

like form described above. These Bremerhaven specimens, which

come from salter water than the preceding, have scarcely any of

the long fine hairs developed, which so strikingly characterise the

freshwater animals.

6. Irish Lakes: British Museum (coll. by Major Trevelyan).

“Tough Nadarragh.” 4 large specimens, male and female,

about 18 mm. in length. Freshwater form, exactly like those

from the Hamburg Wasserleitung.

‘Lough Keenaghan.” 2 specimens, male and female G. pulex.

“ Lough Keenaghan.” 1 specimen, a female, freshwater form,

G. zaddach.

SOME BRACKISH-WATER AMPHIPODA. 661

“Tough Awaddy and Tullynabour.” 4 specimens, three males

and one female, largest 14-15 mm. Freshwater form.

“ Lough Erne.” 4 specimens; two G. zaddachi, male and female,

’ freshwater form; and two G. pulex.

The first to observe the species now under discussion was

Zaddach, after whom I have accordingly named it.

In 1844 he described it under the name of “ Gammarus locusta

Fabr. ?,” but pointed out the characters in which it differed from

Milne-Edwards’s description, adding that if the latter were correct,

his (Zaddach’s) species must be regarded as new. In his later

work he gives a more detailed description and figures of the species,

again, however, emphasizing the differences in the antenne and

the 3rd uropods. Zaddach’s specimens are still preserved at

Konigsberg Museum. The G. locusta, described by Chevreux

(Bull. Soc. Zool. v. xvii. p. 141) as inhabiting the Loire and

the mouths of rivers of Corsica and Provence, is very probably

the species here described.

The principal points of difference between this newly-established

species and G. locusta lie in the antenne, in the proportions of the

peduncle-joints of antenna 1, and in the sete, in the gnathopoda,

in the 4th side-plate, in the armature of the pleon, and in the

3rd uropod.

It differs from G. duebenii, specimens of which were kindly

sent to me by Professor Sars for comparison, in the following

points:—The upper antenna of G. duebenii is only sparsely fur-

nished with hairs, and has none of the regular graduated clusters

so characteristic of G. zaddachi; the hands of gnaths. 1 and 2 are

smaller, and practically subequal in size, in G. zaddachi the hand

of gnath. 2 is decidedly larger than that of gnath. 1; in the adult

G. duebenw the basal joints of pereeopods 3, 4, and 5 are all

expanded, with the hind corners free, while in 4. zaddachi these

joints are hardly expanded at all in the male, and only the 3rd

has the hind corner free ; in the largest G. dwebenii the 1st and

2nd uropods and the telson reach to the level of the peduncle of >

‘uropod 3, and the outer ramus of urop. 3 is twice the length of

the outer ramus of urop. 1 (in the smaller specimens the propor-

tions are as figured by Sars, Crust. Norway, vol. 1. pl. 177); in

G. zaddachi urop. | is considerably longer than urop. 2, and the

outer ramus of urop. 3 only half as long again as that of urop. 1;

the telson in G. dwebenii is shorter, broader, and more spinose.

From G. pulex it is distinguished at once by the shape of the

eye, small and rounded in G. pulex, large and reniform m

G, zaddachi; the antenne, gnathopods, 4th sideplate, pleon

armature, and the 3rd uropods also differ.

The Bremerhaven and Frisches Haff specimens are all young,

the largest, a female figured on Pl. LX XIII., measuring 9 mm.

The largest male in the Konigsberg collection measures 18°5 min. ;

the females are much smaller and broader, 12-13 mm., but none

662 MRS. E. W. SEXTON ON

of them are ovigerous. The largest Hamburg specimen measures

22 mm.

The body is slender and compressed, the brackish-water speci-

mens much more delicate in appearance than the freshwater

ones.

Head (fig. 1) about equal in length to the first two perzeon-seg-

ments, in G. locusta it is distinctly shorter ; lateral lobes obliquely

truncate, upper angle usually subacute in large specimens, and

obtusely rounded in smaller ones, rounded below, sinus small.

Side-plates smaller than in G'. locusta, about as deep as the corre-

sponding segments, 1—4 serrated at the anterior angle. The 4th,

which forms one of the distinguishing characters of the species,

is as broad as deep, with the posterior expansion short and round-

ing into the inferior margin. In G. locusta the posterior expansion

is very deep, with the hind margin straight and much more

serrated.

Pleon-segments 1-3 with the postero-lateral corners acutely

produced, with only two or three setules inset on the hind

margin. Segments 4-6 each with three groups of spines. ‘The

number of spines is very variable: generally speaking, the median

group has two spines, and the lateral groups three each set fan-

wise, and accompanied asa rule by long fine hairs, more numerous

in the freshwater animals; but many “ag the larger animals have

three spines in the median group of the 4th segment, and two

in the 5th and 6th, and four spines in the lateral groups of

the 4th and 5th segments, and three in those of the 6th. The

number varies, however, even in animals of the same size.

The spines are longer than in G’. ocusta, and inset at a rather

different level; the dorsal groups are only slightly raised, whereas

in G@. locusta they are elevated and prominent.

Eyes large, reniform, very dark; outer row of ommatidia

colourless.

Antenna 1 not quite half as long as the body, much more

setose in the male than in the female. The peduncle is longer

than in any other known species of Gammarus, nearly equalling

the peduncle of ant. 2 in length in the adult, and quite equal to

it in the young animal. It is furnished inferiorly with the out-

standing clusters of stiff setee characteristic of the species, some

of the sete in each cluster extending far beyond the rest, and

graduating in length to the distal end of each joint. The

1st joint is longer than the 2nd, but not as long as the 2nd

and 3rd taken together. The primary flagellum is 18-jointed

in the largest Bremerhaven specimen, with 5 joimts in the

accessory flagellum; the largest male from Rauschen had 33

joints in the primary, and 8 in the accessory, the smaller

animals averaged 23-27 in the one, and 4-6 in the other.

Zaddach gives the range as from 25 -35 in the primary, and 5-9

in the accessory. Each joint in the primary flagellum from about

the 5th carries a small stalked sensory filament in addition to the

small sete, and a long seta on alternate joints. In G. locusta the

SOME BRACKISH-WATER AMPHIPODA. 663

peduncle is noticeably shorter than that of ant. 2, extending only

to the distal end of the 4th joint of the latter, instead of to half

the length of the 5th, as in our species; both the peduncle and

flagella are very sparsely provided with sete, and the accessory

flagellum is much longer, 13-14-jointed in the male, with about

47 joints in the primary.

Antenna 2 shorter; 4th joint of the peduncle slightly shorter

than the 5th, both furnished inferiorly with the characteristic

clusters of sete. The flagellum is about as long as the 4th and

5th joints of the peduncle combined, 11-jointed in the largest

Bremerhaven specimen, 15-19-jointed in the large males from

Rauschen ; in both sexes calceole occur on the proximal joints

and clusters of sete similar to those of the peduncle.

In the male of G. locusta, and to a less degree in the female

also, both the peduncle and flagellum are clothed with dense tufts

of long fine delicate sete. The almost glabrous first antenna

and the exceedingly hairy second antenna of G. locusta are sutti-

cient to distinguish the species from G. zaddachi at a glance.

Gnathopoda.—The ditterence between the gnathopods of the

males of the two species will be better seen by referring to

Pl. LXXITI. figs. 2-5. In @. zaddachi (fig. 2) they are much

broader in proportion to their length, with the palm less oblique

and the palmar margin crenulate. In G. locusta (fig. 3) the

palmar margin is sinuous, the clusters of setze on the hind margin

are much denser, the sete, as in ant. 2, are very fine and long,

and the palmar spines are of different structure.

In the female (figs. 4&5) the second gnathopod only carries a

spine midway on the palm; the finger is much more pointed than

in the male.

Perceopoda.—The proportions differ from G. locusta, the 4th

joint being much shorter in proportion and broader, and the 6th

joint, except in pereopod 1, always exceeds it in length. The

2nd joint in pereopod 3 has the lower hind corner produced, sub-

acute; this joint in pereopods 4 and 5 is narrowed distally, the

hind corners not free, but with one or two strong spines inset at

the angle (see fig. 10). In most of the large males the basal joints

are long, very narrow, and scarcely at all expanded posteriorly; in

the female and immature specimens they are always shorter and

more expanded than in the adult male. Generally speaking the

posterior margins of the basal joints and the inferior margins of

the 2nd and 3rd pleon segments are beset with small spines in

the brackish-water form, and with long setiform spines or set

in the freshwater form, but this again is not an invariable rule,

some specimens having spines on the hinder pereopods and sete

on the pleon epimera or vice versé. ‘The hinder perzeopods in the

freshwater animals are covered with long fine hairs set in clusters

with the spines ; these hairs are not much developed in the animals

from salt water, but, on the other hand, they are provided with

more spines.

The wropoda are more slender than in G, locusta. In uropod 3

664 MRS. E. W. SEXTON ON

(fig. 11) the inner ramus is about three-quarters the length of

the Ist joint of the outer ramus; in G. locusta it reaches i the

distal end of the Ist joint, and both ramiare more richly furnished

with spines and plumose setae.

CoroPHIUM LACUSTRE Vanhoffen, 1911. (Pl. LX XIV. figs. 13-

fea)

1911. Beitr. z. Kennt. d. Brackwasserfauna im Frischen Haff.

The largest male measured 3°5 mm.; the largest female

5°) mm.

Body broad, hardly at all compressed. Side-plate 1 with three

very long plumose sete at the anterior angle.

Pleon-segments 4-6 coalesced, strongly resembling C. acutum

Chev.

Head. The rostrum in the male is much produced, apex sub-

acute; not so prominent in the female, apex obtuse. Lateral

lobes rounded, produced, but not to the level of the rostrum.

Eyes large for the genus, irregularly round, pigment black.

Antenna 1.—The peduncle in the male is clothed with long

hairs; lst joint nearly as long as the 2nd and 3rd combined ; in

the female (Pl. LX XIV. fig. 13) the first joint is longer than the

2nd and 38rd taken together ; in both, it is produced below apically

and tipped with one short stout spine; 3rd joint much shorter .

than the 2nd. The flagellum is shorter than the peduncle, about

9-jointed in the male and 7-jointed in the female; terminal

joint rudimentary, the last three jomts each carry a cluster of

sensory filaments.

Antenna 2 very stout; in the male the large 4th joint of the

peduncle carries two apical teeth, the outer one much the larger,

recurved. The 5th joint and the flagellum are furnished in-

feriorly with clusters of very long fine hairs. The flagellum

consists of two joints, the small terminal one with two strong

spines. In the female (fig. 14) the 3rd joint of the peduncle is

produced apically into a small process tipped with a spine; the

Ath joint has a similar process midway on the inferior margin,

and a long apical tooth with a spine inset at the tip: 5th joimt

almost as long as the 4th. The flagellum as in the male, but

showing one joint-division more; one spine on the terminal

joint.

Gnathopod 1 (figs. 15 & 16).—5th joint longer than the 6th,

tapering distally ; 6th as wide at the palm as at the base, but

with hind margin lightly curved; front margin with transver se

rows of setz; the palm convex, serrulate, fringed with six small

spines ; the finger curved, acute, inner margin serrulate, with

three or four setules inset subapically.

Gnathopod 2 (fig. 17) much as in C. triaenonyx Stebbing. The

2nd joint is expanded, more so in the male than in the female;

6th joint long, tapering distally. The finger is tridentate ; teeth

upturned, the 3rd the largest, with small stiff setee inset between.

SOME BRACKISH-WATER AMPHIPODA. 665

Pereopods 1 and 2 stoutly built; the posterior margin of the

2nd joint and both margins of the 4th are fringed with long fine

sete in the male, but carry only a few sete at intervals in the

female. The finger has the glandular aperture opening at

the tip.

The hinder perceopods are alike in both sexes. ‘The 5th is

long and slender, about twice the length of the 3rd. The fingers

of pereopods 3 and 4 are strongly recurved, that of perzeopod 5

falciform. The posterior margin of the 3rd is furnished with a

few simple sete, that of the 4th with plumose sets, while both

margins of the 5th are densely fringed with long and _ short

plumose setz ; the 6th joint of this pereeopod carries two or three

clusters of remarkably long sets posteriorly.

The incubatory lamelle are long, narrow, and fringed with

very long hairs, that of perzeopod 3 almost equals the pereopod

in length.

_ Telsen the same in both sexes, about twice as broad as long,

the apex obtusely truncate; a pair of mobile setz inset on each

side near the base.

EXPLANATION OF THE PLATES.

Prate LXXILI.

Fig. 1. Head of Gammarus zaddachi, sp. n., young female, 9 mm. Bremerhaven.

The clusters of hairs are much denser in the males and in the older

females. « 42.

2. Hands of gnathopods 1 and 2, G. zaddachi, large male,17 mm. Rauschen.

x 20.

8. Hands of ie ee G. locusta, large male, 17 mm. Plymouth

Sound. X 20.

4. Gnathopod1. Young female. Bremerhaven. G'. zaddachi. X 42.

5. ” 2. 0 ) 2 ” x 42,

6. Perzzopod 1. os a ss = x 42.

7. y) 2. re) ” 2 oy) x 42.

8. oy) 3. oy) 20 ” ce) x 42.

Prate LXXIV.

Fig. 9. Pereopod4. Young female,9mm. Bremerhaven. G.zaddachi, sp.n. X 42.

39 5 ” 2 oy) 29 oy 2” x 42.

11. Uropod 3. - FP be 5 a a ADE

12. Diagrammatic dorsal view of pleon segments 4-6 and telson, showing the

arrangement of the spine clusters; in the males and older females, par-

ticularly in the freshwater form, these clusters are always accompanied by

long fine hairs. Young female, 9 mm. X 42.

13. Antennal. Female. Corophiwm lacustre Vanhoffen. Bremerhaven. X 42.

14. oD 2. oy) oy) cy) » oP) x 42.

15. Gnathopod 1. Male. He As Pe 3 X 58.

16. Finger and palm, gnathopod 1. Male. Corophium lacustre Vanhoffen.

Bremerhaven. x 145.

17. Gnathopod 2. Male. Corophiwm lacustre Vanhoffen. Bremerhaven. X 58.

Proc. Zoou. Soc.—1912, No. XLIV. 44

666 MR. C. TATE REGAN ON

36. Descriptions of new Fishes of the Family Loricariidee

in the British Museum Collection. By C. Tare Reean,

M.A: F.Z:s. oe

[Received March 5, 1912; Read April 23, 1912. ]

(Plates LXXV.-—LXXVII.*)

INDEX.

Systematic: Page

Plecostomus hond@, SP. Weis. ..ecccccecen cence venseccecens 666

Chetostomus palmeri, SP. Ve...csecceeecsececeeerses eveee 667

Co Lepturmus; SPoMs \..crse.<o- teenies sector neenene ee eer se OON,

CO. paucispinis, SP. Me .2....200.e.ecpceseonees seeccetenccsees 667)

Xenocara heterorhynchus, Sp. N..........00.eeeeee.-e ss 668

NE MULLIS PUNAS SP o Was wa vatsereeciace)- seca ease eee OOS

Otocinclus maculipinnis, SP. UN. .........eeeseeneeeeeeee 668

Oxyloricaria taman®, SP. De wee. seeeececeeeeeeeeeeeee 669

OE UCEGIASON, Db TWh “sogncsconegdoscdaoseoncosmoseOpsagaccececcn OBE)

AAUEGAS COMAUUUI SOs Mossnooaconcan aco seo oodnoatprbeareaossece (Vi)

During the last three years several Loricariid fishes have been:

added to the British Museum Collection, including examples of

the following ten species new to science.

1. PLEcostomuS HOND#, sp.n. (PI. LXXVI. fig. 3.)

Depth of body 42 in the length, length of head 3. Depth of

head 13 in its length, breadth 17, snout 13, eye 7, interorbital

width 24. Length of mandibular ramus 3 in interorbital width ;

jaws with about 12 teeth on each side. Snout ovate; supra-

orbital margins slightly raised ; supraoccipital convex, without

distinct ridge; temporal plates not or but feebly keeled. Scutes

spinulose, weakly carinate, 27 or 28 in a longitudinal series,

7 between dorsal and adipose fin, 13 between anal and caudal;

supraoccipital entirely bordered posteriorly by a single scute ;

lower surface of head and abdomen with a few scattered granules

(probably covered with granuiar scales in the adult). Dorsal 17;

first ray nearly as long as head, last 2 as long; base more than

distance from tip of spine of adipose fin. AnalI 4. Pectoral

spine extending to anterior 7 of ventrals. Caudal emarginate,

the middle rays 4 as long as the outer ones, which are longer

than the head. Caudal peduncle 3 times as long as deep.

Reddish brown, with traces of darker spots on head and anterior

part of body; dorsal red, with three series of dark spots between

the rays; caudal with alternate red and dusky bars; lower fins

reddish, with dark spots.

Two specimens, 80 mm. in total length, from Honda, Colombia

(300-400 ft.), presented in 1909 by Sir Bryan Leighton.

* For explanation of the Plates see p. 670.

BP Z.S.1912.P1. LAXV.

A.H, Searle, del et lith.

C, PAUCIEEPINIS: 6. ©, PARMERTL.

CHAETOSTOMUS LEPTURUS.

IP VANS) WON Ae LOCAL

tone, gol

[eRe

a oat

deax,

ACH. Searle del et lth.

3. PLECOSTOMUS HONDAS.

2. xX. HETERORHYNCHUS.

1. XENOCARA MULTISPINIS.

(@)"SINNIGITNOVIW SNTONIOOLO & ‘INO LHBSIAT'O 'S &) AVNVWYL VIHVDIHOTAXO 'T

‘THT I Pp eees Hy *

Doe Ta eb SZ ck

NEW SOUTH-AMERICAN FISHES. 667

9. CH#TOSTOMUS PALMERI, sp.n. (Pl. LXXV. fig. 3.)

Nearest to C\. fischeri Steind. Depth of body 53.in the length,

length of head 33. Head a little longer than broad and nearly

twice as long as deep. Diameter of eye 64 to 8 in length of head,

interorbital width 3 to 34, length of snout 1} to 1#. Length of

mandibular ramus equal to interorbital width. Interoperculum

with 2 or 3 spines. 24 scutes in a longitudinal series, 6 or 7

between dorsal and adipose fin, 11 between anal and caudal.

Dorsal I 8; first ray 2 to ? the length of head, last 3 as long;

length of base equal to distance from posterior part of spine of

adipose fin. Anal 15. Pectoral spine just reaching base of

ventral. Caudal peduncle 22 times as long as deep. Coloration

uniform; dorsal and caudal with or without series of spots on

the rays.

Two specimens, 75 and 95 mm. in total length, from the Rio

Tamana, Rio San Juan, Choco, S.W. Colombia, collected by

Mr. G. Palmer.

3. CHETOSTOMUS LEPTURUS, sp. n. (PI. LXXV. fig. 1.)

Depth of body 54 to 6 in the length, length of head 34 to 32.

Head nearly as broad as long and almost twice as long as deep.

Diameter of eye 8 to 94 in length of head, interorbital width

3+ to 34, length of snout 13 to 14. Length of mandibular ramus

equal to interorbital width. Interoperculum with 5 to 8 spines.

25 scutes in a longitudinal series, 6 or 7 between dorsal and

adipose fin, 11 or 12 between anal and caudal. Dorsal I 8; first

ray nearly as long as head, last 2 to 3 as long; length of base

equal to or slightly more than its distance from adipose fin.

AnalI 5. Caudal obliquely emarginate. Caudal peduncle 33 to

4 times as long as deep. Coloration as in C. marginatus Regan,

from which this species differs especially in the form of the

dorsal fin and the more slender tail.

Three specimens, 135 to 215 mm. in total length, from the Rio

Tamana, Rio San Juan, Choco, 8.W. Colombia, collected by

Mr. G. Palmer.

4. CHHTOSTOMUS PAUCISPINIS, sp. n. (Pl. LXXYV. fig. 2.)

Depth of body 5 in the length, length of head 34. Head as

broad as long and 14 as long as deep. Diameter of eye 8 in

length of head, interorbital width 3, length of snout 13. Inter-

operculum with 2 spines. 24 scutes in a longitudinal series,

6 between dorsal and adipose fin, 10 between anal and caudal.

Dorsal I 9; first ray a little less than ? length of head, last 3 as

long; length of base equal to distance from tip of spine of adipose

fin. Anal I 5. Pectoral spine extending to base of ventral.

Caudal very slightly emarginate. Caudal peduncle 27 times as

long as deep. Back with dark cross-bars; dorsal fin with series

of spots on the rays; tips of all the fins reddish.

A single specimen, 90 mm. in total length, from Tado, Rio

San Juan, Choco, Colombia, collected by Mr. G. Palmer.

. 44%

668 MR. C. TATE REGAN ON

5, XENOCARA HETERORHYNCHUS, sp. n. (Pl. LX XVI. fig. 2.)

Depth of body 6 to 63 in the length, length of head 22 to 232.

Head longer than broad and more than twice as long as deep.

Diameter of eye 7 or 8 in length of head, length of snout 1? to

2, interorbital width 3. Length of mandibular ramus 11 to 13

in interorbital width. Snout with tentacles. Interoperculum

with about 10 spines, the longest about } the length of head.

23 or 24 scutes in a longitudinal series, 7 or 8 hepueen dorsal

and adipose fin, 11 or 12 Heiwebn anal and caudal. Dorsal I 7;

first ray 4 to = the length of head, last, when laid back, separated

by 3 keeled scutes from adipose fin ; base of dorsal equal to or

less than its distance from adipose fin. Anal13. Pectoral spine

extending to base of ventral. Caudal obliquely truncate. Caudal

peduncle 3 times as long as deep. Fins with series of dark spots.

Two specimens (¢), 58 and 80 mm. in total length, from

Uruhuasi, Peru (4000 ft.), collected by Messrs. H. & C.

Watkins.

This species is related to Y. bufoniwm, differing somewhat in

proportions and also in the much greater width of the naked

margin of the snout, into which the bony plates extend as a narrow

median prominence nearly reaching the apex of the snout.

6. XENOCARA MULTISPINIS, sp. n. (Pl. LX XVI. fig. 1.)

Depth of body about 5 in the length, length of head 23. Head

longer than broad and twice as long as deep. Diameter of eye

8 10 94 in length of head, interorhital width 22 to 24, length of

snout 14 +02. Length of mandibular ramus 2 in the interorbital

width. Snout with many long tentacles(¢). Interoperculum

with 20 to 25 spines, the longest } the length of head. 24 scutes

in a longitudinal series, 6 or 7 between dorsal and adipose fin,

10 or 11 between anal and caudal. Dorsal I7; first ray about

3 length of head, last about = as long; base equal to distance

from posterior part of spine of adipose fin. Anal I 4. Pectoral

spine extending to anterior 3 or middle of venta Caudal

obliquely tr uneate. Caudal peduncle 24 to 22 times as long

as deep. Traces of rather large pale spots on ade dorsal and

lower fins with series of dark spots; caudal dusky.

Three specimens (¢), 90 to 120 mm. in total length, from the:

Humboldt and Novo Rivers, Sta. Catherina, 8.E. Brazil, collected

by Mr. W. Ehrhardt.

This species is near to X. stigmaticwm Eigenm., and except for

the larger size of the spots on the head might as justly be referred

to that species as those that I described as X. séigmaticwum from

the Mogy-guassu River. Dr. R. von Ihering has written to me

that this locality is not 250, but only 25 miles inland of Santos.

7. OvocINCLUS MACULIPINNIS, sp. n.. (Pl. LX XVII. fig. 3.)

Closely related to O. nigricauda and O. perforatus. Depth of

body about 54 in the length, length of head 23. Diameter of eye

NEW SOUTH-AMERICAN FISHES. 669

about 7 in length of head, interorbital width 2%, length of snout

2t0 21, Occipital region evenly convex, without crests. Scutes

spinulose, not carinate, 22 to 24 in a longitudinal series. Lower

surface of head with a naked area in front of clavicles; abdomen

with 4 or 5 irregular, series of plates between the lateral series.

Dorsal I 7; origin a little behind base of pelvics, but not much

nearer to end of snout than to base of caudal. Anal 15. Pectoral

spine extending beyond middle of ventral. Caudal emarginate.

Caudal peduncle 3 times as long as deep. Fins with series of

dark spots.

Four specimens, 30 to 35 mm. in total length, from the La

Plata, presented in 1909 by Herr J. Paul Arnold.

8. OXYLORICARIA TAMANS, sp.n. (Pl. LX XVII. figs. 1, 1a, 1.)

Head 53 in the length. Breadth of head 1} (d) or 14 (@) in

its length, diameter of eye 7 to 8, interorbital width 33, length of

snout 12, Snout without distinct rostrum; supraorbital edges

not or scarcely raised; sides of the snout, in the male, with

bristles. 32 or 33 scutes in a longitudinal series (14-15+ 18) ;

lateral keels weak, obsolete anteriorly, united posteriorly. A

naked area surrounding the lips ; abdomen posteriorly with 3 series

of plates between the lateral series; an anal plate bordered

anteriorly by 3, these 3 again by 5. Dorsal elevated, its height

3 the length of the fish; pectoral spine extending to anterior 4

of ventral; caudal emarginate, the outer rays produced. Breadth

of body at level of last anal ray 44 to 5 in the distance from

the caudal. Anterior half of dorsal blackish ; caudal blackish at

the base; pectoral blackish, pelvics more or less dusky, anal

spotted anteriorly.

Two examples (3, @), 225 and 200 mm. in total length, from

the Rio Tamana, Rio San Juan, Choco, 8.W. Colombia (200 ft.),

collected by Mr. G. Palmer.

9. OXYLORICARIA LEIGHTONI, sp.n. (Pl. LX XVII. fig. 2.)

Head 42 in the length. Breadth of head 12 to 1} in its length,

diameter of eye 8, interorbital width 31 to 33, length of snout 14

to 2. Snout without distinct rostrum ; supraorbital edges not or

scarcely raised. 30 scutes in a longitudinal series (15+ 15); lateral

keels weak, united posteriorly. A naked area surrounding the’

lips ; abdominal plates posteriorly forming 3 or 4, anteriorly 6 or 7,

irregular series between the lateral series ; an anal plate bordered

anteriorly by 3. Height of dorsal equal to length of head ; pectoral

spine extending to anterior 7 of ventral; caudal emarginate, the

outer rays produced. Breadth of body at level of last anal ray

1 the distance from caudal. Fins with blackish spots; anterior

part of dorsal blackish at base and near tip.

Two specimens, 55 and 65 mm. in total length, from

Honda, Colombia (300-400 ft.), presented in 1909 by Sir Bryan

Leighton.

670 ON NEW SOUTH-AMERICAN FISHES.

10. ARGES CIRRATUS, sp. 1.

Length of head 33 in the length of the fish. Interocular width

rather more than distance from eye to posterior nostril, rather

less than 7 the length of head. Nasal flap produced into a barbel

which is as long as the diameter of the eve; maxillary barbel not

quite reaching gill-opening; teeth of outer series of premaxil-

laries unicuspid, acute; mandibulary teeth bicuspid. First dorsal

ray 3 the length of head; pectoral extending to base of ventral ;

ventrals inserted below anterior part of dorsal, nearly reaching

anal. Spine of adipose fin well developed, movable, connected

by a membrane with the caudal peduncle, inserted at a distance

from caudal equal to about 4 the length of the middle rays of

that fin. Distance from tip of snout to origin of dorsal fin 22

in the length of the fish, from base of last anal ray to caudal 532.

A few large dark spots on body; a blackish bar across middle of

caudal fin.

A single specimen, 24 mm. in total length, from the Rio

Condoto, Rio San Juan, 8.W. Colombia, collected by Mr. G.

Palmer.

EXPLANATION OF THE PLATES.

Prats LXXV.

Fig. 1. Chetostomus lepturus.

Fig. 2. 3 paucispinis.

Fig. 3 ce palmeri.

Pirate LXXVI.

Fig. 1. Xenocara multispinis.

Fig. 2 st heterorhynchus.

Fig. 3. Plecostomus honde.

Prate LXXVII.

Fig. 1. Oxyloricaria tamane (X24).

Figs. la, 1b. Heads of ¢ & 9.

Fig. 2. O. leightoni.

Fig. 3. Otocinclus maculipinnis (X2).

ON EPIDERMAL SHEATHS FROM BILL OF THE KING PENGUIN. 671

EXHIBITIONS AND NOTICES.

April 23, 1912.

Dr. 8. F. Harmer, F.R.S., Vice-President,

in the Chair.

The Secrerary exhibited a living specimen of a young female

Dorsal Hyrax (Dendrohyrax dorsalis) from Nigeria, recently

presented to the Society by Mr. J. L. McKellar.

The Secrerary exhibited a number of photographs of an

Elephant Kraal in Siam which had been presented to the

Society by the Rt. Hon. Sir Cecil Clementi Smith, P.C.,

G.C.M.G.

May 7, 1912.

Prof. EK. A. Mincuin, F.R.S., Vice-President,

in the Chair.

Mr. R. I. Pocock, F.R.S., F.LS., F.Z.S8., Superintendent of

the Gardens, exhibited a skin and a living specimen of a fawn

variety of the Brown Rat (Zpimys norvegicus), which had been

caught on an island in the middle of Loch Corrib, Co. Galway,

and presented to the Society by Lord Headley. Mr. Pocock

remarked that although similarly coloured varieties of this rat

had been caught now and again in different parts of England, it

was especially interesting to put on record Lord Headley’s state-

ment that it was quite common on the island, no fewer than

eleven having been trapped while others had been seen; and that

it did not occur, so far as was known, on the mainland. Typically

coloured brown rats lived on the island as well.

Mr. D. Sera -Smirx, Curator of Birds, exhibited two horn-like

sheaths which had been shed from the orange-coloured patch at

the base of the lower mandible of the King Penguin (Aptenodytes

pennantr) living in the Society’s Gardens. Mr. W. E. de Winton

had observed the shedding of this epidermal sheath in a bird living

in the Gardens in 1898 (P. Z.S. 1898, p. 900); but although the

present specimen had been carefully watched during two suc-

cessive moults in March and October 1911 (P. Z.S. 1912, p. 60),

no sign of this process was observed. The bird, however, went

through another complete moult in March to April of the present

year (1912), and shortly after this was completed the epidermal

covering of the orange-coloured patches became loose and finally

672 MR, G. A. BOULENGER ON

fell off; the pieces somewhat resembled the wing-cases of a large

beetle, being semi-transparent and of a clear orange-colour.

Dr. Francis Warp, F.Z.8., showed a number of photographs

and diagrams illustrating a method of observation of fishes, birds,

and NG under the water, the principle being that thé

subjects under consideration were Uluminated by natural hight,

and the observer, being in a dark chamber in the water, was ‘not

seen. The appearance of black-feathered birds was shown ;

these, by carrying down air-bubbles among the feathers, were

converted into reflectors : and a Water-Hen was shown br ight red

and then green as it reflected the different surroundings in

which it had been placed. Otters and Seals were also shown as

seen under the water, ‘The demonstration was illustrated by

numerous slides and by the cinematograph.

PAPERS.

37. On a Collection of Fishes made by Mr. A. Blayney

Percival in British East Africa to the East of Lake

Baringo. By G. A. Bounrnanr, F.R.S., F.Z.8.*

[Received April 2, 1912; Read May 7, 1912.)

(Plates LXXVITI.-LXXX.+)

INDEX.

Systematic : Page

QBGO Herverwwrds, SPoDs aver ssec cena cascnevinaes (ovuctawkces 673

Barbus argyrot@nia, SPoDe cc ccccscccccsececceesneae B74

BHAWAUS SPD KS REL NE RES, Pe, GGA

Amphilius owyrhinus, SPM. cee 675

TUAPIG MEVOWALG SMa dis, oss ssncesncsncancernennceneeey MORO

The collection here reported upon, presented to the British

Museum by Mr, Blayney Percival, is of special importance as

coming from a district the fishes of which had not been collected

before, from a watershed of its own without any communication

with sea. The river called Eusso Nyiro, or Guaso Nyiro, quite

distinct from the Southern Guaso Nyiro in German East Africa,

takes its source to the east of Lake Baringo, separated from it

and from Lake Rudolf by hill-ranges 1000 to 1500 feet higher

(4000 to 5000 feet above sea- -level) and is lost in the Lorian

* Published by permission of the Trustees of the British Museum.

+ For explanation of the Plates see p. 676.

P25. 1912) tay.

J.Green del.et lith.

LABEO PERCIVALI.

Nee a ee Mz) ev leprae

J.Green del.et lith.

Ze yWE TIVES.

1L.BARBUS ARGYROTANIA,

See er Ne WAS rs ENG INA A

“SNONIHYAXO SNITIHAINV

“UAT, Ye"Tep useag 'p

4 Or diel 2161 ies wa GL

FISHES FROM BRITISH EAST AFRICA. 673

Swamp. The Eusso Mara is a tributary of the Eusso Nyiro.

The Saya is a separate river, between the Anagata Borita and the

Eusso Nyiro, flowing north and lost in a small swamp some

50 miles north of Eusso Nyiro. Unless otherwise stated, the

fishes here listed were obtained in the Eusso Nyiro, below the

falls.

The fishes of this basin have much in common with those of the

Webi Shebeli and Juba, and nothing with those of Lake Rudolf.

Curiously, the small Barbus, B. nairobiensis and B. percivali,

previously discovered by Mr. Percival in the Nairobi (Athi River

system), on the east side of Kilimanjaro, were found again in

abundance in the Eusso Nyiro.

1. Mormyroes DELICIosuS Leach.

2. MormMyRUS KANNUME Forsk.

. 3, LABEO PERCIVALI, sp.n. (Pl. LX XVIII.)

Body strongly compressed, its depth 33 to 3? times in total

length. Head 4 to 42 times in total length, 14 to 13 times as

long as broad; snout rounded, 3 length of head; eye perfectly

lateral, 23 (young) to 3% times in length of head; interorbital

width 2, or a little over 4, length of head; lips with small papillee

forming transverse plice ; lower lip with a fringe of large papille;

rostral flap moderately large, with entire or indistinctly denticulate

edge; a small barbel in the corner of the mouth. Dorsal III

10-11, equally distant from end of snout and from root of caudal,

or slightly nearer the former; upper edge concave ; last simple ray

in adult males much produced, twice, or nearly twice, as long as

head, not or but little longer than head in females and young.

Anal ITI 5, reaching or nearly reaching root of caudal. Pectoral

considerably longer than head in males, reaching base of ventral,

the first ray of which falls below fourth or fifth branched ray of

dorsal, shorter in females. Caudal deeply emarginate, with long,

pointed lobes. Caudal peduncle 13 to 13 times as long as deep.

Scales 38-42 a 5 between lateral line and root of ventral,

18 round caudal peduncle. Silvery, back brownish grey ; vertical

fins greyish.

Numerous specimens, measuring 55 to 190 mm.

Allied to Z. newmanni Blgr. Well distinguished by the larger

eye and the shape of the dorsal in the males,

4, LABEO CYLINDRICUS Peters.

Also from the Eusso Mara and the Saya, and the Lorian

Swamp.

5. DiscoGNATHUS DEMBEENSIS Riipp.

‘Also from the Eusso Mara and the Saya.

674 MR. G. A. BOULENGER ON

6. BARBUS ERLANGERI Bler,

Also from the hot springs at Chandler Falls, upper Eusso Nyiro,

and from the Saya.

7. BARBUS NAIROBIENSIS Bler.

From the Eusso Nyiro above the falls.

8. BARBUS PERCIVALI Bler,

From the above locality, and also from the Saya.

The largest specimen measures 105 unm.

9. BARBUS ARGYROTANIA, sp.n. (Pl. LX XIX. fig. 1.)

Depth of body 2? to 3 times in total length, length of head 31

to 37 times. Snout rounded, shorter than the eye, which is 22 to

3 times in length of head and equals or nearly equals interorbital

width; mouth small, subinferior; lips feebly developed ; two

barbels on each side, anterior about 3 diameter of eye, posterior

as long as or slightly longer than eye. Dorsal III 8, equally

distant from posterior border or centre of eye and from base of

caudal, border feebly concave; last simple ray enlarged, bony,

strongly serrated behind. Anal III 5, not reaching caudal.

Pectoral a little shorter than head, not reaching ventral ; base of

latter below origin of dorsal. Caudal peduncle a little longer

than deep. Scales radiately striated, 30-31 a, 3 between lateral

line and ventral, 14-16 round caudal peduncle. Pale sand colour,

with a silvery lateral band which is sometimes edged above with

a streak of black pigment; fins white, often tinged with pale

orange at the base.

Total length 53 mm.

Numerous specimens from the Eusso Nyiro below the falls.

B. argyrotenia stands nearest to B. zanzibaricus Peters.

10. Barsus Mimus, sp.n. (Pl. LXXIX. fig. 2.)

Depth of body 23 to 22 times in total length, length of head

33 to 4 times. Snout rounded, much shorter than the eye, which

is 23 to 3 times in length of head and equals interorbital width ;

mouth small, subinferior; lips feebly developed; two barbels on

each side, anterior about 3 diameter of eye, posterior a little

longer than eye Dorsal IIT 8, equally distant from posterior

border or centre of eye and from base of caudal, border feebly

concave; last simple ray not enlarged, not serrated, as long as or

a little longer than head. Anal III 5, not reaching caudal.

Pectoral as long as head, not quite reaching ventral; base of

latter below origin of dorsal. Caudal peduncle as long as deep or

a little longer than deep. Scales radiately striated, 25-26 = 2-24

between lateral line and ventral, 12 round caudal peduncle.

Pale sand-colour, with a silvery lateral band, which is sometimes

FISHES FROM BRITISH EAST AFRICA. 675

edged above with a streak of black pigment; fins white and

transparent, or pale orange at the base.

Total length 55 mm.

Numerous specimens from the Eusso Nyiro below the falls.

Closely allied to B. neglectus Blgr. from the Nile, and strikingly

resembling the preceding species in general appearance, although

differing widely in the feeble, non-serrated last simple dorsal ray

and in the much lower number of scales.

11. Cuartas LAzerA Lacep.

Kusso Nyiro, near the Lorian Swamp.

12. EvurTrRopPius DEPRESSIROSTRIS Peters.

13. Bagrus urostiema Vincig.

EKusso Nyiro, above the Chandler Falls.

14. CLAROTES LATICEPS Rupp.

15, AMPHILIUS GRANDIS Bler.

16. AMPHILIUS OXYRHINUS, sp.n. (PI. LXXX.)

Depth of body 6 times in total length, length of head 32 times.

Head much depressed, slightly longer than broad ; snout pointed,

4 length of head; eye very small, 11 times in length of head,

25 times in interorbital width; posterior nostril midway between

eye and end of snout; maxillary barbel $ length of head, reaching

root of pectoral; outer mandibular barbel 2 length of head, inner

nearly 2. Gill-rakers rather long, 7 on lower part of anterior

arch. Dorsal I 6, much nearer end of snout than root of caudal.

Adipose dorsal low, 33 times as long as deep, twice as long as

rayed dorsal, its length 3 its distance from latter. Anal IIT 5.

Pectoral a little more than % length of head. Veutral a little

shorter than pectoral, far behind base of dorsal. Caudal feebly

emarginate. Caudal peduncle 13 times as long as deep. Reddish

brown above, spotted with blackish, dirty white beneath.

Total length 195 mm.

Distinguished from 4. grandis by the pointed snout, the longer

caudal peduncle, and fewer anal rays.

A single specimen from the Eusso Mara, a swift mountain-

stream.

17. SyNopDonTIS SCHALL BI. Schn.

18. SYNODONTIS GELEDENSIS Gthr.

From the Eusso Nyiro, close to the Lorian Swamp.

19, Tinapra niLorica L.

From the Eusso Nyiro, above and below the falls.

676 ON FISHES FROM BRITISH EAST AFRICA.

20. TILAPIA PERCIVALI, sp.n. (PI. LX XIX. fig. 3.)

Depth of body equal to length of head, 24 times in total length.

Head about twice as long as broad; snout rounded, with straight

or slightly convex upper profile, a little broader than long, ? post-

ocular part of head ; eye 33 (young) to 4 times in length of head,

greater than preorbital depth; mouth large, ? width of head,

extending to below anterior border of eye; lips very strongly

developed ; teeth moderately slender, in 3 or 4 series, 40 to 50

in outer series of upper jaw; 2 series of scales on the cheek, width

of scaly part rather less than diameter of eye. Gill-rakers short,

13 or 14 on lower part of anterior arch. Dorsal XJJJ-XV 11;

spines increasing in length to the last, which measures # length of

head ; longest soft rays $ length of head. Anal III (exceptionally

IV) 9-10; third spine 3 length of head. Pectoral about ? length of

head, not reaching origin of anal. Ventral reaching vent or anal.

Caudal rounded, subtruncate. Caudal peduncle deeper than long.

Seales cycloid, 30-31 =; lateral lines ag Dark brown, with

black vertical bars on the body, or entirely black.

Total length 75 mm.

Hot springs at Chandler Falls, Northern Eusso Nyiro, 2500 ft.

Caught in water about 100° Fahr.

Near allies of this new Zilapia are 7’. alcalica Hilgend. from

alkaline water in volcanic holes near the Nguruman Salt Lake,

into which the Southern Guaso Nyiro flows, on the boundary of

British and German East Africa, and 7. grahami Blgr.from Lake

Magadeé, a perfectly isolated hot soda lake in the Rift Valley,

British East Africa, not far from the Southern Guaso Nyiro.

The former has, like 7. percivali, D. XIII 11, but the scales on

the cheek are in two series and the caudal is truncate ; the latter

has D. XI 11-12, 3 series of scales on the cheek, and the caudal

peduncle is deeper than long.

EXPLANATION OF THE PLATES.

Pratt LXXVIII.

Labeo percivali, p. 678. 3 nat. size.

PuatE LXXIX.

Kig. 1. Barbus argyrotenia, p. 674.

2. As minus, p. 674.

3. Tilapia percivali, p. 676.

PuatEe LXXX.

Amphilius oxyrhinus, p. 675.

ON NEW CESTODES FROM THE TASMANIAN DEVIL. 677

38. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea. By Frank EH. Bepparp, M.A.,

D.Sce., F.R.S., F.Z.8., Prosector to the Society.

[Received April 2, 1912: Read May 7, 1912.]

(Text-figures 92-101.)

V. On A NEw Genus (Dasyurotenia) FROM THE TASMANIAN

Devi (Dasyurus ursinus), THE TYPE OF A NEW Famity.

INDEX.

Systematic : Page

Dasyurotaniay BEN. We c..cecseyersscnecessssseeceteetseses ODA

EDV POUUSEAN SP Mla ee Pos ooo ca ace oem ie eA

Résumé of anatomical points............0:ceeeeeeeeee. 695

The Cestoidea hitherto recorded from Marsupials appear to

belong to the following genera only, viz. Triplotenia, Moniexia,

Bertiella, Linstowia, Anoploteenia, Oochoristica, and Cittotenia.

To these must now be added an eighth genus, of which I have

recently examined examples from the Tasmanian Devil, in whose

small intestine they occurred.

Text-fig. 92.

Dasyurotenia robusta, nat. size.

Two views of the same two specimens, to show their attachment to the gut.

The left-hand figure represents the interior of the gut, the right-hand

figure the outside.

The accompanying drawings (text-fig. 92) show two of the

three specimens which I obtained 7 situ. Hach worm is firmly

imbedded at the head end in the wall of the gut. The two

678 DR. F. E. BEDDARD ON

drawings show the inner surface and the outer surface of the

intestine. The body of the worm is rather flat though at the

same time fairly thick, and is curved in a sickle-like fashion.

The length of the larger of the two individuals is about 17 inches,

and the greatest breadth does not exceed 5mm. There is nota

very marked difference in diameter at the neck end of the body,

and the worm thus presents a very solid and strong appearance,

which is correlated with the very firm way in which it is im-

bedded in the gut. The narrowness of the segments and the

lack of overlap of successive segments contributes further to the

stout appearance of the worm.

Text-fig. 93.

Head of scolex of Dasyuwrotenia robusta, shown by opening up the cyst on the gut.

The last two or three segments of the body are rather narrower

from side to side and are curved, thus tending to encircle each

other successively. This state of affairs is shown in the drawings

referred to. Anteriorly the strobila is seen to perforate the wall

of the intestine, which forms a fold round it, like a collar not

fitting very closely. The strobila at this point disappears from

view when the intestine is examined from its internal surface.

NEW CESTODES FROM THE TASMANIAN DEVIL. 679

When the piece of gut to which the worms are attached is turned

over they reappear on the outer surface (as is shown in text-

fig. 92) in the form of a cyst. This cyst forms a simple hemi-

spherical bulging upon the wall of the gut consisting apparently

of the peritoneum. When this peritoneal wall is cut through a

Text-fig. 94.

Transverse section through scolex of D. robusta at about the middle.

The central medulla shows the problematical vesicles referred to in the text in

the middle of its extent and the coils of the water-tubes at the ends,

vr. Irregular furrows upon scolex. ¢. Tube formed by these.

A second tube of the same nature lies above and to the right of that lettered.

cavity is opened up in which lies the large scolex. This is

illustrated in text-fig. 93. The scolex, it will be observed, does

not completely fill the cavity in which it lies.

680 DR. F. E. BEDDARD ON

The large size of the scolex of Dasyurotenia contrasts with the

relatively small or even minute scolex of other Tetracotylea, and

thus recalls that of the Tetraphyllidea and others of the ‘“‘ lower”

groups of Tapeworms. So also do certain other characters of

the scolex in this worm. The greatest diameter of the scolex is

3°5 mm., and it is of much the same length, so that its dimen-

sions are but little less than the diameter and thickness of the

body, and are greater than those of the neck-region which imme-

diately ensues. Broadly speaking, the scolex is of a globular form

and has a soft, easily indentable, irregular surface marked by faint

ridges and depressions, and not at all like the vertex of a typical

member of the Tetracotylea. Two flat plate-like areas are to be

seen when the scolex is examined in this way with a pocket-lens.

These are, as [£ believe, the expanded suckers which I have seen

in a contracted condition in another specimen whose scolex was

studied by means of transverse sections.

Transverse sections of the scolex (text-fig. 94, p. 679) are

approximately circular in outline, in the middle part at any rate.

This particular scolex, which I studied by the section method,

was apparently more retracted than the one represented in text-

fig. 93. For at first the sections (text-fig. 95) showed two semi-

circular masses closely applied along the straight margin which

is obviously the expression of an apical groove in the scolex.

It was lower in the series than this that the contour of the

sections became circular. At the body end of the scolex the

latter shghtly overlaps the neck for the whole of its circum-

ference. But although the sections through the scolex are on the

whole circular in form, the peripheral layer is not at all uniform

and shows numerous processes and grooves occurring everywhere,

which is an expression of the irregular grooving and ridging of

the scolex apparent when that region is examined with a hand-

lens. I could detect no symmetrical arrangement in these pro-

jections and grooves, which, however, may collectively represent

“bothria” meandering over the surface of the scolex.

It is furthermore the fact that these grooves are converted in

places into tubes which run along the interior of the peripheral

layer of the scolex in a longitudinal direction, though not for a

long distance. These short tubes end blindly. Their existence,

together with the grooves and ridges of which they are a further

development, must facilitate the adhesion of the scolex to the

surrounding walls of the cyst. While the more or less irregular

grooves and furrows upon the surface of the scolex might be put

down to irregular contraction of its outer layer, this can hardly

be the case—one would suppose—with these invaginated tubes.

In any case the arrangement of the suckers does not conform to

the arrangement usually met with among the Tetracotylea (to

which group this worm would be expected to belong). The scolex,

however, possesses four suckers which are of very small size when

compared with the diameter and circumference of the scolex.

NEW CESTODES FROM THE TASMANIAN DEVIL. 681

The greatest diameter of one of the suckers is not more than

one-tenth the diameter of the scolex at its widest part.

Text-fig. 95,

Section through apex of scolex to illustrate relative size of one of two

smaller suckers (s.).

g. Median groove formed by retraction of sucker.

On either side of the apical groove already mentioned was a

single cup-shaped sucker bearing hooks. I assume, therefore,

that these two suckers are really anterior in the fully expanded

condition of the scolex, and that they are the two disk-like bodies

Proc. Zoou. Soc.—1912, No. XLV. . 45

682 DR. F. E. BEDDARD ON

described above in the scolex, which was examined entire and by

the aid of a lens only. These two suckers were not exactly

opposite on the opposite sides of the groove. They represent

all that I can find in the scolex comparable to a rostellum. I

believe, however, that they do not represent the rostellum of the

Tetracotylea.

Text-fig. 96.

Two consecutive sections through one of two larger suckers bearing hooks.

A. Muscular layer of sucker. B. Cellular layer in which lie hooks (H.).

In these sections (text-fig. 96) the sucker appears cup-shaped,

the orifice naturally opening into the median transverse groove

NEW CESTODES FROM THE TASMANIAN DEVIL. 683

upon the scolex. The structure of the sucker is not like that of

the typical Tetracotylean sucker. In the latter there is one thick

layer of typically arranged muscle-fibres (into the nature of which

arrangement it is not my purpose to enter—it is well known)

which is quite distinct from the surrounding tissues of the scolex,

but never covered by them on its free surface, which faces towards

the aperture. In the present species the structure of this sucker

is illustrated in the accompanying text-figure (text-fig. 96), which

is drawn from one sucker only which appears to me to be rather

larger than its fellow, the two being thus mutually asymmetrical.

Examining the sections from below upwards, the sucker is seen

at first to be apparently like that of the Tetracotylea generally.

It consists of a strongly marked layer of muscle-fibres etc. lying

in a cavity of the body parenchyma. ‘This is shown in the

figure. As we approach more nearly to the rostellar region,

another layer in which the hooks are implanted pushes itself

in front of the sucker proper, as is also shown in the text-

figure referred to. This layer is continuous with and similar

in structure to the general parenchyma of the head. The

true (?) sucker thus gets as it were buried beneath a layer

of tissue bearing hooks. So far the structure of this sucker

will be plain from the sections figured. In the corresponding

sucker of the opposite side of the body the hooks are obvious

and quite similar, and imbedded in a perfectly corresponding

tissue. But I could find no trace whatever of the muscular

structure independent of these hooks.

In addition to these two suckers the scolex of Dasyurotenia

possesses two others, thus making the normal four. The latter

are roughly opposite to each other, but do not—obviously,

at least —alternate with the others. These two suckers

are opposite to each other and lie on the outside of the

scolex, and are about on a level with the hooked suckers

already described. It is quite possible that when the scolex

is fully expanded and not warped through unequal contrac-

tion with alcohol, the four suckers might alternate with more

regularity than is apparent in my preparations. In any case,

these suckers are quite different from those already described.

They are smaller than, at any rate, the larger of the more apically

placed suckers; and there is no trace whatever of any hooks

associated with them. Each appears (see text-fig. 95) in section

to have the form of a flat disk. Their structure seems to be

quite like that of suckers generally among Tapeworms. What

is very apparent about these suckers is their very small size com-

pared with the wide periphery of the scolex. I could not detect

them at all on an examination of the scolex with a lens (see

text-fig. 93, p. 678).

We have next to consider the histological structure of the

scolex, which presents certain peculiarities. ‘The anterior region

of the scolex has no distinction into medullary and cortical layers.

The commencement of such a differentiation is marked by the

A5*

684 DR. F. E. BEDDARD ON

depth of the invagination of the anterior end of the scolex referred

to above. This part of the scolex presents a very lax appearance

in sections, which is doubtless to be correlated with the irregular

outline of the scolex. Through this lax tissue stray in every

direction muscle-fibres which no doubt effect the retraction of this

anterior part of the scolex. The main mass of tissue in which the

muscular fibres run has the usual amorphous ground-substance of

the Cestoid body with many nuclei; but I have not made a

particular histological study of these tissues.

Text-fig. 97.

Part of transverse section through neck-region of Dasyurotenia robusta.

C. Cortical layer outside of longitudinal muscles (Z.i.).

me. The thin medullary layer.

Further back in the scolex the medullary region appears as

quite distinct from the cortical. In the latter the muscular

fibres gradually collect into a massive bundle of longitudinally

running fibres which immediately surround the medulla and

occupy a great deal of the cortex. Within the scolex there is no

subdivision of this muscular mass into bundles such as I shall

NEW CESTODES FROM THE TASMANIAN DEVIL. 685

describe in the strobila and even in the neck-region of the same.

In the medullary region at either end is to be seen the water

vascular tube, which is here rather coiled, a tube appearing five

or six times in a single cross section. In the middle of the

medullary area are certain remarkable structures, which are

represented in text-fig. 94 (p. 679). These are apparently

hollow spheres excavated in the parenchyma but with very

definite walls. These vesicles contain lumps of an amorphous

matter’ into the nature of which I have not enquired. There

is no connection that I could find between these vesicles and

the water vascular system. I am quite uncertain as to their

nature. :

The neck-region of the worm after it has issued from the cyst

which contains the scolex is represented in transverse section in

text-fig. 97. It will be observed at once that the medullary»

parenchyma is very greatly reduced. It forms a thin layer very

much narrower than the extremely thick cortical layer. Nor

eould I discern in it the typical retiform appearance of the

Cestoid medullary parenchyma with spherical masses of granular

matter lying between the meshes of the network. The whole of

the available space, save between the individual testes at the two

sides of each proglottid, is occupied by the rudimentary female

organs, which together with the testes seem to occur in the very

first segments. ‘The whole space thus occupied by the generative

masses and by what remains of the medullary parenchyma is not

one half of the diameter of the cortex. It has been already

pointed out that at the posterior end of the scolex, a little before

it merges with the neck, the medulla is surrounded by a very

thick layer of muscles composed of large fibres. In this layer no

marked arrangement of the fibres into bundles could be made

out. But in the next few sections, which we here speak of as

the neck, these fibres are very definitely disposed in a series of

bundles (text-fig. 98).

These bundles are disposed in four to six layers, and at the

two lateral extremities of each segment they are rather more

numerous than medianly. The bundles consist of a large number

of individual fibres, often as many as sixty or so. They are sepa-

vated from each other by a dense nucleated tissue. Later in the

body we have the same four to six rows of bundles of fibres ;

but in these latter segments each row is separated from those

which lie above and below it by a delicate layer of transverse

fibres. These fibres are not apparent in the neck-region. I

imagine that this enormously powerful muscular system is corre-.

lated with the distance to which the scolex is enabled to force

itself into the wall of the intestine of its host. These rows of

bundles of longitudinal fibres are reminiscent of what is charac-

teristic of the family Acoleide, as is to be seen, for example, in

the genus Proterogynotenia,* where they have beer figured by

* Abh. Senck. Ges. 1911, p. 260, fig. 14.

686 DR. F. E. BEDDARD ON

Fubrmann. Outside of these layers of fibres the cortex presents

no features which differentiate it in any marked way from other

allied forms.

Text- fig, 98.

1 . 3

i < op kowy

t./71..

Section through the muscular layer of the cortex, taken at a point further —

back than that represented in text-fig. 97.

4.m. Bundles of longitudinal fibres. #.m. Transverse fibres.

The water vascular system of this tapeworm is in more than

one respect remarkable. In transverse sections through the

ripe proglottids only two vessels are as a rule visible, one on

each side of the body. The most careful examination often

failed to reveal the presence of another, even minute, tube. Nor

did longitudinal sections show any trace of this second vessel.

It is not, however, really absent in this worm, as I found it -

to be in Thysanotenia lemuris, for it is present as a very minute

tube in some segments. The vessels which are obviously present

’ NEW CESTODES FROM THE TASMANIAN DEVIL. 687

are of very large size, and it appeared to me that that on the pore

side of the segment was a little the larger of the two. This, then,

is the first important point about these vessels, 7. ¢. that there is

a very lavge ventral tube on each side of the body and a very

minute dorsal tube. In following out a series of transverse

sections, it is seen that the lumen of the large ventral water-

vessels is occasionally occluded by a delicate diaphragm-like

sheet of membrane which is abundantly nucleated. There is no

question of a narrowing of the calibre of the tube, but of an

actual membrane which extends partly across it here and there.

In sagittal sections the existence of these membranes stretch-

ing partly across the lumen of the water vascular tube is quite

obvious. They occur, moreover, on both sides of the body, that is

to say in the case of both ventral tubes. The reason for emphasizing

this fact will be apparent later. In the longitudinal sagittal

sections referred to it will be seen that there are several of these

membranes which stretch a good way across the water-vessel,

and though two membranes arising from different sides of the

vessel do not actually meet, the edge of each stretches beyond the

edge of the other, so that the tube would appear, when viewed in

the direction of its length, to be entirely occluded. It is note-

worthy that these diaphragms, so to speak, arise indifferently

from both sides. The exact arrangement will be plain from the

annexed drawing (text-fig. 99, p. 688). Ihave noticed that in some

of the posterior proglottids the lumen is actually occluded once

in each proglottid. The two ends of two oppositely projecting

membranes are connected by a continuous though very thin

membrane which connects the thicker extremities of the lateral

projections. This median part appears to be nowhere deficient,

and the water vascular system is thus divided up in these regions

of the body into a series of chambers.

T presume that these numerous membranes stretched across the

large vessels correspond to what has been figured in other tape-

worms as valves. I have emphasized the fact that they occur on

both sides of the body, because they carry on the pore side the

genital ducts which actually perforate their substance and lie in

their thickness. This perforation is limited to the larger of these

valve-like structures which arise from the outer side, and it has

been produced I imagine by the extension round the ducts of the

water-vessel, which is of much greater diameter in the posterior

than in the most anterior segments. In the case of these latter,

as already mentioned, the generative ducts pass to one side of the

water-tube.

Another important feature in the water vascular system of this

tapeworm is the total absence of transverse vessels uniting the

longitudinal trunks in such segments. This state of affairs is

not unknown among other Cestoidea—it occurs, for example, in

Hymenolepis acuta *—but it is not common. Nor is this lack of

* y, Janicki, Zool. Anz. Bd. xxvii. 1904, p. 776.

688 DR, F. E. BEDDARD ON

transverse vessels compensated by any network of excretory tubes

pervading the medulla, such as is met with in the genera Jnermi-

capsifer and Zschokkeella *.

Text-fig. 99.

Sagittal section to illustrate ventral water vascular tube.

V. Projecting valves. v.d. Vas deferens. v.a. Vagina.

T. Testes.

* Beddard, P. Z.S. 1912, p. 596, and literature quoted there.

NEW CESTODES FROM THE TASMANIAN DEVIL. 689

The Generative organs of this tapeworm begin to be recog-

nisable very early in the body, only a segment or two behind the

head. But it is a long way back before the ovaries are ripe. As

in the vast majority of Cestoidea, the testes ripen earlier than the

ovaries. This being the case, the testes are recognisable earlier

as distinet bodies, and only cease to be so clear in the more pos-

terior segments, where the uterus is gorged with eggs. A remark-

able point of interest in the generative organs is the fact that the

duct leading to the exterior, or to be more exact the formative

mass of cells which will be both vagina and cirrus sac, is seen to

alternate in position in relation to the single water vascular tube

and the nerve-cord. In all segments the actual opening is on the

same side of the body, but the generative duct passes towards it

either between the water-tube and the nerve-cord or outside of

both ; in the latter case only to one side—there is no alternation

between a dorsal and a ventral position. We shall see, when the

cirrus sac and vagina come to be described later, that there is

also variation in the exact relationship in position between these

two.

The ovaries are large and consist of two wings, which are

symmetrical or very nearly so, the middle point between them

being the middle line of the body, where are situated the shell-

gland etc. The ovaries are posterior in the segment, and behind

them lie the vitelline glands. These latter are of much the same

shape as the ovaries, and in rather immature segments differ only

from them by their rather darker staining with hematoxylin.

They also form two wings symmetrical with their middle point,

and are in contact with the ovaries in front. The two glands are

composed of many lobes, which reach as far as the testes at the

sides; altogether an ovary occupies fully half of the segment, and

rather more when it is fully mature. Immediately in front of it

are the sperm-duct and vagina. At the sides are to be found the

testes, which are also dorsal to it.

The vagina offers no very remarkable character. It has at first

a contracted lumen, which widens out for a considerable space,

and then contracts again before it suddenly opens into the rather

pear-shaped receptaculum seminis. The course of the vagina is

quite straight between its two ends and oblique in direction.

The narrow part of the vagina which opens into the receptaculum

is of some length, but shorter than the wider part. The vagina

shows quite the same characters in the most mature proglottids.

The receptaculum seminis is full of spermatozoa, and very

frequently contained ova at its wider end, close to where the

oviduct opens into it. The receptaculum and the vagina lie

anteriorly in the segment in front of the ovary, but behind the

uterus.

The uterus of this worm is persistent and found as a large

cavity extending right across the segment in the most mature

proglottids that I have examined. It begins as a small rounded

cavity lying in the front part of each segment.

690 DR. F. E. BEDDARD ON

The testes of this worm are chiefly massed at the two sides of

the proglottid, but these two masses are connected by a string

of testes which pass dorsally along the proglottid. They therefore

nearly surround the ovary and the female organs generally, so far

partly resembling Cyclorchis. In transverse sections the testes

appear circular in section; but sagittal sections such as that

represented in text-fig. 99 (p. 688) show that the form of the testes

is that of a flat plate, for in those sections they appear more linear

in shape. These sections also show quite plainly that each testis

Text-fig. 100.

hee AD

A portion of the coil of the sperm-duct (sp.) gorged with sperm.

ce. Interstitial prostatic cells.

lies in a space, unless, indeed, the appearances produced are due

to shrinkage through reagents. In any case, however, the testes

of these worms frequently present the appearance of being sur-

rounded by the spermatozoa which they produce, thus showing

that a chink exists or can be formed for their reception when

pressed out of the testis. The testes are very numerous and

quite crowded together,—so much so that the delimitations

between successive segments so far as concerns these organs

NEW CESTODES FROM THE TASMANIAN DEVIL. 691

are not at all visible in sagittal sections; they appear as a

continuous mass. ;

The sperm-duct runs straight for a short way after it has left

the cirrus sac. It forms a copious coil (text-fig. 100) occupying the

middle of the body and lying partly dorsally to the receptaculum

seminis and rather nearer to the pore side of each proglottid. The

rest of the coil—that part which is nearest to the cirrus sac—is

Text-fig. 101.

Sagittal section showing in consecutive segments the varying relations of

cirrus sac (the larger tube) and vagina.

to the pore side of the receptaculum, and therefore ventral in

position. In fact, regarded as one coil, this region appears oblique

in direction in transverse sections of the proglottids. Between the

individual loops of the sperm-duct are cells which quite fill up

the interstices, and are thus numerous in proportion to the width

692 DR. F. E, BEDDARD ON

of those interstices. The cells are rather clear, with well-stained

nuclei. They correspond exactly, as it seems to me, with the

prostatic cells of Jnermicapsifer and Zschokkeella, dealt with in

these genera by v. Janicki* and myself t, and which appear to

occur also elsewhere. In immature segments the cells bulk more

largely than the coils of the sperm-duct. But the reverse is the

case in the mature proglottid.

The genital ducts open into a common cloaca genitalis, which in

its turn opens on to the exterior. The cloaca genitalis is of some

depth, but it is not borne upon any process of the body. Into it

open, close together, the vagina and the cirrus sac, whose mutual

relations I have investigated by means of sagittal sections (text-

fig. 101) through portions of the strobila. There is a con-

siderable variation in these relations. Although the vaginal pore

is apparently never in front of the male pore, it is not always

directly behind the male pore. The commencing vagina lies

obliquely behind the commencing cirrus sac—the direction of the

obliquity being now dorsal now ventral in this segment and in that.

There is, in fact, an irregular alternation from segment to seg-

ment; sometimes the two tubes are not merely oblique, but actually

dorsal or ventral to each other as the case may be, lying there-

fore side by side in sagittal sections. This recalls to mind the

alternation that occurs in certain (but not all) species of the

genus Monezia, where the vagina may be dorsal or ventral to the

crus sac. But in this latter genus the alternation is of the

right and left set of generative organs of a single segment.

The cirrus sac of this tapeworm is large and has the very common

flask-shape. The neck-1egion has very thick circular muscular

walls, forming a sheath which thins out over the more distended

region of the sac. In less mature segments (in which, however,

the testes are fully developed, though with no mature sperma-

tozoa) the cirrus sac is elongated, gradually diminishing in breadth

towards the external pore; there is no marked division into neck

and flask. It is very long and extends towards the middle line

of the body, a little beyond the water vascular tube of its side of

the body, or at least reaches the internal side of that tube. The

cirrus runs straight from end to end of the cirrus sac and

anteriorly presents a moniliform appearance, which is due to suc-

cessive dilatations of the lumen of the cirrus. This region of the

cirus is both preceded and succeeded by a perfectly straight

section of that tube with very narrow lumen and thick walls.

In later segments the cirrus sac acquires the flask-shape already

referred to. Coincidently with this is an actual shortening of the

length of the entire sac and a coiling of the cirrus within it.

The cirrus sac in these and in subsequent segments hardly reaches

beyond the outer edge of the water vascular tube. It seems clear

therefore that the shortening is due to an actual contraction of

* Denkschr. Ges. Jena, xvi. 1910.

+ P. ZS. 1912, p. 602.

NEW CESTODES FROM THE TASMANIAN DEVIL. 693

the length caused by a bulging of the walls of the cirrus sac due

in its turn to the rapid growth and consequent coiling of the

cirrus. When the cirrus sac is in this fully formed condition, the

cirrus itself is differentiated more thoroughly into those regions

less markedly indicated in earlier stages. ‘The sperm-duct enters

the cirrus sac at the apex and its lumen contracts to a fine line

for a short space near to its entry. This is particularly obvious

in the last few segments of the body, where the sperm-duct has

become much dilated before its entry into the cirrus sac, and thus

offers a greater contrast to this exceedingly narrow region.

In these more mature cirrus sacs the flask-shape has been

acquired, as already mentioned. But the neck of the flask is

much longer than the body part. The latter is so fully occupied

by the coils of the cirrus itself that there is but little of the inter-

stitial packing tissue to be seen. At its entry into the cirrus sac

and for a considerable time thereafter the duct is thick-walled,

with a very narrow lumen, and much coiled. This region of the

cirrus is succeeded by a not very long but coiled tract, which is

much wider and has thinner walls. ‘The lining membrane bears

numerous spinelets. Finally, the distal region of the cirrus is

again thick-walled and with a narrow lumen: it opens into the

genital cloaca without any alteration of character. In the most

posterior segments of the body the greater part of the cirrus sac

is filled with sperm, the posterior region alone showing a group of

coils of the cirrus. Whether the anterior part of the cirrus has

become ruptured, as it appeared, or has been simply enormously

expanded and its walls reduced to extreme tenuity by the enclosed

sperm, I am unable to say.

The systematic position of this tapeworm is difficult to fix with

any confidence. ‘The generative system, and, indeed, the internal

anatomy generally, presents no differences of importance from

many Tetracotylea; and there are, indeed, no reasons so far why

the worm should not be placed in the Anoplocephalide, which

family, as has been pointed out, contains nearly all the Marsupial

tapeworms. On the other hand, the very much developed layers

of longitudinal muscles in the body-wall suggest the family

Acoleide. The difficulty, however, of accurately placing the worm

lies in the peculiarities of the scolex. There is no doubt that it

contrasts very considerably with the general form of the scolex

among the ‘Vetracotylea in a number of points, of which the

principal ones are :—(1) its large size, both relatively to the body

and actually; (2) the presence of numerous grooves which cannot

be, at any rate, entirely artefact, as they are converted here and

there into tubes running within the thickness of the head;

(3) the relatively minute size of the four suckers and the fact that

two of them and two only are furnished with hooks*,

* Furthermore, these hooks are distinctly hollow at their broader end, “like a

Ruminant’s horn,” as Shipley (Willey’s Zool. Res., Entozoa, 1900) notes of Callio-

bothriwm, one ot the Tetraphyllidea.

694 DR. F. E. BEDDARD ON

These characteristics are collectively different from anything

met with among the Tetracotylea that is known to me. They

are not, however, inconsistent with the conditions known to occur

among the Tetraphyllidea, if we may admit the grooves upon the

scolex to represent the bothria of such tapeworms. ‘The suckers

would then correspond with the small accessory suckers so

frequently possessed by these latter worms, and their small size

relatively to the scolex would be thus intelligible. The apparently

numerous bothria not reducible (by me) to symmetry is suggestive

of a type like Phyllobothrium* slightly modified, or perhaps

Peltidocotyle t. We cannot, however, place Dasyurotenia among

these Tetraphyllidea on account of the Tetracotylean character of

its yolk-gland. But with reference to this gland it may be borne

in mind that it is in structure much more diffuse than is usual

with the generally solid vitelline gland of the Tetracotylea.

The genus may be thus defined :—-

Dasyurotenia, gen. nov.

Stoutly built worms with large scolex bearing four small suckers,

of which the inner two bear hooks. No rostellum, but anterior end

of scolex, including hooked suckers, retractile. Segments very

short. Inner layer of (longitudinal) muscles very thick, consisting

of four to six rows of bundles of fibres. Ventral excretory tubes

large, with numerous valves not communicating with each other in

the strobila. Dorsal vessels minute, not always visible. Genital

pores unilateral. Testes numerous, chiefly lateral, anterior, and

dorsal. Vas deferens with a large coil in middle of segment sur-

rounded by prostatic cells ; cirrus sac large, cirrus with spinelets.

Ovaries with two wings, median and posterior and. ventral in

position, in front of vitelline gland, which is also symmetrical.

Shell-gland median, dorsal. Receptaculum seminis present, nearly

median, ventral. Uterus sac-like, persistent, fills nearly whole of

ripe proglottid. Lggs thin-shelled.

Hab. Marsupials.

The species I term “robusta” on account of its very stout build.

It is, however, quite impossible for me to venture upon an

enumeration of the peculiar specific characteristics for the present.

This genus and species cannot, as I think, be identified with

any other form that has been described from an Australian

Marsupial. From the present genus we only know a species

described by myself § a little time since as Anoplotenia dasyuri.

Nor can I identify it with any of the genera enumerated at the

beginning of this paper in other Marsupials. In fact, Bertiella is

* Bronn’s ‘ Thierreich,’ Bd. iv. Cestoiden, pl. xli. fig. 10.

+ Ibid. pl. xlui. fig. 1.

{ Middle line of female apparatus only slightly displaced towards pore side.

§ P.Z.S. 1911, p. 1003.

NEW CESTODES FROM THE TASMANIAN DEVIL. 695

the only one of these genera to which it bears any likeness in the

reproductive system, and from this genus the characters of the

scolex at once distinguish Dasyurotenia. Indeed, its inclusion

among the Tetracotylea (=Teenioidea) is not, to my mind, an

obvious certainty. In any case the hooked suckers exclude it

from the family Anoplocephalide, to which nearly all the

Marsupial tapeworms belong up to the present.

The most salient points of anatomical interest in this worm

appear to me to be the following :

(1) The immense size, relatively speaking, of the scolex and the

small size in comparison with it of the suckers. The fact that

two suckers are armed with hooks while the other two suckers

are not so armed.

(2) The great thickness of the longitudinal muscles, which

consist of at least four layers of bundles each containing very

many individual fibres of considerable stoutness.

(3) The existence for the most part of only a single water

vascular tube on each side of the body, which is, moreover, in the

posterior segments completely divided up into a series of com-

partments, one to each segment, and whose lumen is also here and

more anteriorly divided by delicate septa jutting into its cavity.

Furthermore, by the fact that these tubes are not connected

in successive segments by transverse vessels, as is so nearly

universally the case.

(4) As a remarkable structural feature, which is at present

mysterious in nature, may be mentioned the isolated cavities in

the medullary region of the head which have no connection with

the water vascular tubes.

(5) An anatomical feature of some importance is the very

variable relation to each other in position of the extremities of

the male and female ducts, which is correlated with an orifice

upon one side of the body only. An alternation in the position

of the external pore may, we know, be accompanied with difference

in the relative position of the ducts as, for example, in the double

series of genital tubes of Moniezia.

(6) In view of the very considerable peculiarities of structure

briefly indicated in the foregoing résumé, it may be worth men-

tioning, as a remarkable fact, that the generative organs do not

show any marked features of interest as compared with those of

other tapeworms.

696 MR. R. E. TURNER ON

59. Studies in the Fossorial Wasps of te Family Scoliide,

Subfamilies Elidinee and Anthoboscine. By RowLanp

EK. Turner, F.Z.S., F.E.S.

[Received March 29,1912: Read May 7, 1912.]

(Plates LX XXI.-LXXXITI.*)

InDEX.

Geographical Zoology : Page

Ant hobOScaMmMOIsttlDItLONNOLGE eee e epee teeee ea see eee 726

Systematic :

PAMENODOSCE) OCCUPULAIES SPuMe .aeeeneeee treet eeeeeerene ee Oe

Braunsomeria, gen. n. au Te Meee aa cee ee MOOI

Pe Giatrathenpets SRT Aes, eA Se ee GOS

A AIA ESS 0 0G | Galcroeaseee badcdaocgeeaSsdone: Oar)

TE iso OF |S PsMek ess eRe. teehee hoe ene ee cas eeccheas eeRY see

5 EIS CLLOU IA SDs Ml. Sause fs texcatdbeers sites ste ou oiies rete sage ae eM

Bg ui CULES) NaLLACiCe: Sp OMe cee ngcmecue nae aise Genanee eel 704,

i 1 CHOICHUCHIDIS, SIDS WS dob bonhocaa daboeecoonsocnneeo ON

m Fe QU PAU UCAS Dene cer aehenteebiee cestnccpesee atten ao

Ps Bil) GUTUCEN UC SD. Meme meee ata cre ces choco eats cee ALO

» Sy i CORT LBGEUS, SU DG Sohcnorsulecororionbong secon. »//117

a yf DI RSpoliataispamls cess hel oksce seca stet ee TALL

Mazine braunsi, Sp.) gages) ee eas sscheesssnae-esece serene OO

oy CROMMADIAED Novis WO) TOES antec sddeoocee paceenorncaazanens 7/2444

» constrictiventris, sp. N. ........ Re araee Ole

» sauakinensis Grib. to Uniiooonen oe ease LO)

Sith pSCLQUICCL SDE Me SRE coi esc. Bah ena eS,

PETIA RTT ACIS) SILI. ist Seanbaned Eee ieaaaciaeee a deeeecane 702

Odontothynnus Cam. 6 to Anthobosea.. ‘i 724.

Plesia continua Cam. $ of Myzina pada prnaliag caer 703

» erythronota Cam. to Anthobosea .................. 738

» leucospila Cam. ,, 2, I DR eee 739

» melanaria Cam. ,, 5 pCR ee ROG,

The following notes on the Elidine and Anthoboscine will, it is

hoped, facilitate the study of these neglected groups, the latter of

which especially has been very little understood by many authors,

who have touched on it merely as describers of new species,

Saussure alone having seriously studied the group. The best -

work on the Elidinz has been done by the same author, but the

material at his disposal was very limited.

I am indebted to Dr. Brauns, of Willowmore, 8. Africa, for

valuable assistance with many carefully collected specimens. ‘The

material available is still insufficient for a revision of the species

of Myzine, as to which much confusion still exists.

The species which I have not seen are marked with an asterisk.

* For explanation of the Plates see pp, 753-754,

TP Gasea ase, xe,

West, Newman chr.

Horace Knight del.et lith.

FOSSORIAL WASPS.

SN aie Seles dL zyrel al es UD O:SIN

Catherine A.M.Pearce del. West,Newman lith.

WING NEURATION OF FOSSORIAL WASPS.

her

ee ae

Ses ‘s

IE) Las lh S hile Jel ey OOS)

Catherine A.M.Pearce del. West, Newman lith.

EXOSKELETAL STRUCTURES OF FOSSORIAL WASPS.

STUDIES IN THE FOSSORIAL WASPS. 697

Family ScoLiipDa,

Subfamily Exp.

BRAUNSOMERIA, gen. noy.

2. Apterous; mandibles acute at the apex, with a rather in-

distinct tooth on the inner margin near the apex; antennze twelve-

jointed, the first joint of the flagellum very small and almost con-

cealed by the apex of the scape. Head almost rectangular, the

posterior angles slightly rounded ; eyes oval, touching the base of

the mandibles, rather small, separated by a distance at least as

great as their own length from the posterior angles of the head;

ocelli absent, their position indicated by large punctures. Thorax

much narrower than the head; pronotum rather longer than its

greatest breadth; mesonotum very short, almost covered by the

pronotum, the tegulz more or less developed ; scutellum narrower

than the pronotum, broader than long ; median segment nearly as

long as the pronotum, flattened on the dorsal surface, broadened

from the base to the apex. Sides of the head and thorax and base

of the abdomen thinly covered with long hairs. Abdomen longer

than the head and thorax combined, shining, the apical segment

long and more or less acute at the apex, stricture between the first

and second ventral segments well developed. Intermediate cox

rather widely separated, posterior coxze contiguous, intermediate

and posterior tibiz spinose, tarsal ungues simple.

6. Winged; stigma rather large, situated at about three-fifths

from the base of the wing; radial cell shorter than the stigma;

three cubital cells, the second and third small, not reaching the

apex of the radial cell, each receiving a recurrent nervure; cubital

and discoidal nervures not continued beyond the cells. Medial

cell of the hind wing not emitting veins from theapex. Antenne

in the typical species long and slender, thirteen-jointed, the first

joint of the flagellum almost concealed in the apex of the scape,

the antennz much longer than the abdomen ; antennal tubercles

well developed. Head strongly convex; ocelli present. First

abdominal segment with a short petiole, the segment, including

the petiole, a little longer than the second segment, suddenly

widened at the apex of the petiole. Apical segment with a re-

curved spine, the apical emargination of the dorsal segment

shallow. Eyes entire, not emarginate.

The characters given here for the male will doubtless be found

not to apply to all species of the genus; but the important

characters in the neuration separating the males from Myzine

are the larger stigma, the blunter apex of the radial cell, the fact

that the cybital and discoidal nervures are not continued beyond

the cells as in Myzine, and that no veins are emitted from the

median cell of the hind wing, there being two veins in Myzine.

Type of the genus, Braunsomeria quadraticeps.

Proc. Zoo. Soc.—1912, No. XLVI. AG

698 MR. R. E. TURNER ON

BRAUNSOMERIA QUADRATICEPS, sp. n. (Pl. LXXXTI. figs. 9, 10;

PIPEX XM. Ge. 7)

2. Rufo-ferruginea ; mandibulis apice, vertice, capite lateribus ;

segmentis abdominalibus tribus basalibus nigris ; femoribus tibtisque

Juseis, calcarvis albidis.

Long. 8 mm.

@. Head broader than long, fully half as broad again as the

pronotum ; mandibles with a blunt tooth on the inner margin

near the apex, another near the middle of the inner margin,

acute, and another smaller nearer the base. The whole insect

shining, with a few scattered punctures. Pronotum longer than

broad, a little longer than the median segment. Dorsal abdo-

minal segments broadly depressed at the apex, the basal portion

of the segments produced into a slightly raised rounded mark

on each side; apical segment very narrowly rounded at the

extremity.

3. Niger; mandibulis, clypeo macula mediana nigra, scapo

subtus apice, tuberculis antennalibus, macula parva frontali, mar-

gine interiore oculorum, linea’ undulata verticali, pronoto macula

utringue antice et fuscia lata postice, mesonoto macula quadrata,

scutello fascia lata, postscutello, mesopleuris fascia, segmento dor-

sali primo fascia apicali, ceteris fascia apicali macula nigra

utrinque, segmentisque ventralibus 2-6 fascia bisinuata apicali

pallide flavis ; alis hyalinis, venis testaceis.

Long. 6 mm.

3g. Antenne slender, longer than the abdomen, the inter-

antennal tubercles prominent. Clypeus very short and broad,

very shallowly emarginate at the apex. Head very strongly

convex, cheeks as broad as the eyes. Head and thorax coarsely

but not very closely punctured, median segment finely and closely

punctured-rugulose; abdomen shining, very sparsely and shallowly

punctured. Pronotum shorter than the mesonotum, narrowed

anteriorly, the anterior margin straight, posterior margin very

feebly arched. Median segment steeply sloped posteriorly, not

truncate. Petiole of the basal abdominal segment occupying less

than half the length of the segment, the remainder of the segment

slightly inflated; the first segment, including the petiole, only a

little longer than the second. The segments not constricted ;

hypopygium forming a long recurved spine; apical dorsal segment

convex, shallowly emarginate at the apex. Stigma large, twice

as long on the costa as broad, nearly twice as long as the radial

cell, which is broadly rounded at the apex. Three cubital cells

on the right side, two on the left; on the right the second

abscissa of the radius is very short, the third about equal to the

first and second combined, but shorter than the second transverse

cubital nervure, second recurrent nervure received close to the

apex of the third cubital cell.

Hab. Willowmore, Cape Colony ; January (Dr. Brauns).

The female is the type.

STUDIES IN THE FOSSORIAL WASPS. 699

T have little doubt that links will be discovered connecting both

sexes of this genus with J/yzine through the short-winged Pseudo-

meria section in the females, and through species with somewhat

more extended neuration in the males. But the apterous con-

dition of the female and the differences of neuration pointed out

in the description of the male seem to me to be suflicient reason

for founding a new genus. The female shows a strong resem-

blance to female Thynnide of the genus 4irone, also to the

Bethylid genus Apenesia. I have not been able to examine the

mouth-parts, but it is likely that they would show atrophy of

some parts. The entire eyes of the male are also noticeable as

contrasted with the shallowly emarginate eyes of MWyzine.

BRAUNSOMERIA ATRICEPS, Sp, 0.

2. Nigra; mandibulis basi, clypeo, antennis, therace, segmento

mediano, pedibus pygidioque apice ferrugineis,

Long. 5 mm.

@. Mandibles acute, with a very small ill-defined tooth on the

inner margin near the apex, Head rectangular, a little broader

than long, very slightly convex, shining, with a few scattered

punctures. Thorax shining, with a few scattered punctures on

the pronotum, the median segment more closely punctured.

Pronotum longer than broad, slightly narrowed anteriorly,

narrower than the head by about one-third; mesonotum very

short, the tegule rather better defined than in quadraticeps ;

scutellum rounded posteriorly, broader than long, Median seg-

ment a little shorter than the pronotum, slightly broadened from

the base, a little broader than long and obliquely sloped poster-

iorly ; sides of the thorax and median segment sparsely clothed

with long yellowish hairs, Abdomen shining, finely aciculate,

the basal segment truncate anteriorly, with a ‘short petiole not

more than half as long as the posterior cox ; the third segment

the broadest; a semicircular small raised mark on each side of

dorsal segments 2-5; sixth segment smooth, pointed at the apex.

The constriction between the two basal ventral segments is well

marked. The eyes are smaller than in quadraniceps and are

separated from the posterior angle of. the head by a distance

equal to about three times their own length.

Hab. Algoa Bay, Cape Colony ; November (Dr, Brauns).

Myzine(?) sttema, sp,n, (Pl. LXXXI. fig, 11; Pl, LXXXITI.

fig. 13.)

3. Niger; mandibulis basi, pronoto anguste postice, tegulis,

tibiis subtus tarsisque basi pallide flavis; alis hyalinis, venis

perlucidis, stigmate maximo, pallide flavescenti; oculis haud

emarginatis, antennis abdomine br evioribus ; cellula radiali oblite-

rata, cellula cubitalt secundo poene obliterata.

Long. 7 mm.

g. Clypeus very short, transverse. Antenne about as long as

16*

700 MR. R. E. TURNER ON

the thorax and median segment combined, not very slender, of

about even thickness throughout, inserted a little nearer to the

eyes than to each other. Eyes not emarginate, their inner

margins parallel; posterior ocelli a little nearer to each other

than to the eyes. Head convex, closely punctured, with a

frontal suleus reaching to the ocellus; antennal tubercles not

developed. Thorax rather sparsely punctured. Pronotum rather

short, as broad as the head, the anterior margin straight, the

posterior margin widely and feebly arched. Scutellum large, a

little shorter than the mesonotum. Median segment short, almost

smooth, with a median sulcus, truncate posteriorly. Abdomen

subsessile, the basal segment broad, not constricted at the apex

on the dorsal surface, deeply divided from the second on the

ventral surface, all the segments sparsely punctured and shining ;

the apical segment rather deeply triangularly incised for the

reception of the Jong aculeus of the hypopy gium; the whole

abdomen about equal in length to the head, thorax, and median

segment combined. Stigma. very large, about twice as long as

the greatest breadth ; only one cubital cell and one recurrent

nervure, which is received on the cubitus just beyond the angle

of the cubital cell, the cubitus continued just beyond the point of

reception of the recurrent nervure, the radial cell and all neura-

tion beyond the stigma obliterated, Median and submedian cells

of the hind wing present, but no neuration beyond them, the

median cell not extending very far beyond the submedian.

Hab. Willowmore, Cape Colony (Dr. Brauns).

This very distinct species will probably prove to be generically

distinct from J/yzine. It approaches most nearly to WM. swalei

Turn. and J/. braunsi Turn., but differs in the reduced neuration,

the entire eyes, the more robust and subsessile abdomen, and the

much broader stigma. But until the female is known I prefer to

leave it provisionally in Myzine. From Braunsomeria, to which

the neuration approaches more nearly than to M/yzine, it may be

distinguished by the much more robust build, the much shorter

and stouter antenne, and the deeper emargination of the apical

segment.

MyzINE BRAUNSI, sp.n. (Pl. LXXXI. fig. 14; Pl. LXXXTI.

fig. 14.)

3. Niger; mandibulis basi, pronoto fascia angusta postice,

segmentis dorsalibus 2-6 macula transversa mediali apice, mcacr-

laque curvata laterali utrinque, tegulis, tarsis articulo apicali

excepto, tibiis anterioribus omnino, intermedtis posterioribusque

basi pallide flavis ; alis hyalinis, venis perlucidis, stigmate pallide

testaceo.

Variat segmentis dorsalibus macula mediali obliterata.

Long. 5-8 mm.

3. Clypeus much broader than long, closely punctured and

shallowly emarginate at the apex. Antenne gradually thickened

STUDIES IN THE FOSSORIAL WASPS. 701

towards the apex, the terminal joint twice as thick as the fourth ;

the antenne about as long as the head, thorax,and median seg rene

combined. Eyes slightly emarginate on the inner margin ;

posterior ocelli nearly as far from each other as from the eyes.

Head and thorax closely and not very finely punctured ; pronotum

narrower than the head, shorter than the mesonotum, narrowed

anteriorly, the anterior margin straight, the posterior margin

widely but not strongly arched. Median segment transversely

rugose, with a longitudinal depression in the middle, truncate

posteriorly, the surface of the truncation coarsely transversely

striated. Abdomen narrower than the thorax and nearly half as

long again as the head, thorax, and median segment combined,

finely and sparsely punctured, the segments scarcely constricted

at the base; petiole of the basal s sesment very short, the segment

abruptly broadened and slightly swollen, nearly as Jong as the

second segment without including the petiole. Apical segment

convex, the incision at the apex subtrianeular, not quite as deep

as its ‘apical breadth. Tarsal ungues simple. Cubital and

discoidal nervures not continued beyond the cells, stigma not

rounded on the inner margin, three times as long as the greatest

breadth ; radial cell acute at the apex, produced far beyond the

third cubital cell, second and third abscisse of the radius nearly

equal in length, second recurrent nervure received just before the

middle of the third cubital cell. Median cell of the hind wing not

emitting any veins from the apex.

Hab. ‘Willowm ore, Cape Colony; January to March (Dr. Brauns).

This is allied to JZ. swalei Turn., but differs in the shorter and

stouter basal abdominal segment, in the sculpture of the median

segment, and in the tr anslucent nervures of the wings. Both

differ from typical Myzine by not having the cubital and discoidal

nervures continued beyond the cells and in the absence of the two

veins emitted from the apex of the median cell of the hind wings,

MYZINE CONSTRICTIVENTRIS, sp. n. (Pl. LXXXII. fig. 15;

Pl. LX XXIII. fig. 12.)

3. Niger, dense albo-pilosus ; mandibulis basi, pronoto margine

posteriore, tegulis basi, segmentis dorsalibus 2-6 macula mediana

apicali et macula transverse uth ‘inque, tibiis basi tursisque pallide

flavis ; alis hyalinis, venis brunneis.

Long. 10-12 mm.

3. Clypeus broad and short, shallowly emarginate at the apex.

Eyes distinctly convergent towards the clypeus, the inner margin

not emarginate, almost straight. Antenne inserted nearer to

each other than to the eyes, almost as long as the abdomen,

moderately stout and of almost even Pigienee throughout.

Posterior ocelli a little nearer to the eyes than to each ‘other.

Head and thorax closely and strongly punctured ; pronotum

short, narrower than the head, the anterior margin straight, the

posterior margin very feebly arched. Median seoment coarsely

702 MR. R. E. TURNER ON

rugose, almost vertically truncate posteriorly. Abdomen much

jonger than the head and thorax combined, not very slender,

sparsely punctured, the segments rather strongly constricted at

the base, subsessile, the first segment no longer than the second.

Sixth dorsal segment more coarsely punctured, not convex, the

lateral margins raised, the apical emargination shallow, much

broader at the apex than deep. Madial cell broad, nearly twice

as long on the costa as the greatest breadth ; second abscissa of the

radius distinctly longer than the third, second recurrent nervure

received close to the middle of the third cubital cell.

- Hab. Willowmore, Cape Colony; October to January (Dr.

Brauns).

This species is easily distinguished by the absence of any

emargination of the eyes, the shallow emargination of the apical

dorsal segment, the strongly constricted abdominal segments, and

the broad radial cell. The cubital and discoidal nervures are

continued very little beyond the cells, and the two nervures

emitted from the apex of the median cell of the hind wing are

very short.

MyYZINE UMBRATICA, sp. 0.

2. Nigra, mandibulis pygidioque fusco-ferrugineis ; segmentis

dorsalibus 2-3 macula laterali utrinque alba; alis fusco-violaceis.

Long. 10 mm.

@. Shining and almost smooth, coarsely but not very closely

punctured round the base of the antenne; a few scattered

punctures on the vertex, pronotum, mesonotum, and scutellum,

pro- and mesopleure strongly but not very closely punctured ;

median segment closely and rather finely punctured at the base,

almost smooth in the middle and at the apex, a few obscure striz

at the posterior angles; abdomen almost smooth, with a few

small punctures on the apical portion of. the segments. Long

black pubescence on the sides of the thorax; calcaria whitish.

Hyes rather narrowly ovate, cheeks as broad as the eyes; ocelli

small, the posterior pair about as far from each other as from the

eyes: Antenne smooth and shining, thescape beneath punctured

and clothed with long hairs. Head subrectangular, half as broad

again as long, much broader than the thorax. Pronotum about

twice as broad as long, the posterior margin almost straight.

Mesonotum only half as long as the pronotum and a little shorter

than the scutellum, the parapsidal furrows very distinct. Apical

segment of the abdomen convex, long, and pointed. Wings of

moderate length, reaching to the fifth abdominal segment, the

stigma situated about halfway between the base and apex;

heuration similar to that of rufifrons Fabr.

Hab. Fourteen Streams, Cape Colony; January (Dr. Brauns).

There is only a very obscure median sulcus on the median

segment. The slit in the fore wing extends from the termination

of “the cubitus just beyond the third cubital cell to the mar gin of

the wing.

STUDIES IN THE FOSSORIAL WASPS.

LXXXITI.

ABDOMINALIS Guer.

MyzZIne (Pl.

Pl, LX XXIII. fig. 5.)

10.

11.

13.

. Sixth dorsal segment ‘punctured ;

. Abdomen wholly bright ferruginous

. Lateral margins of the Wiedianl ‘segment acute ;

. Apical or two

Meria abdominalis Guér. Rev. de Zool. iii.

703

Lie ype)

365 (1839), 9

Plesia continua Cam. Rec. Albany Mus. i. p. 299 (1905), d

Hab. Willowmore; Burghersdorp, Cape Colony.

Taken in copula by- Dr. Brauns.

Key to the Ethiopian Species of Elis (Mesa).

Females.

. Basal joint of posterior tarsi with a row of spines

beneath

Basal joint of posterior tarsi unarmed or with a scopa

only beneath .....

Radial cell distinctly ‘separated ‘from the costa for

more than half the Jength. Black, two apical

abdominal segments red .

Radial cell separated from ‘the costa at the apex

only

. Abdomen ferruginous, the basal segment only black.

Abdomen black

. Thorax and abdomer entirely black

Thorax more or less red, the prenot shorter than

the scutellum

‘calearia of the

hind tibie black

Sixth dorsal segment striate; calcaria whitish

. Vertex red, much more sparsely punctured than the

front, mesonotum and scutellum black. Spur of

the Cae tibie not strongly bent near the

base .

Vertex black, “as ‘closely. ‘punctured _ as the “front,

mesonotum and scutellum red. Spur of the

posterior tibie strongly bent near the base .........

Abdomen black, the two apical segments sometimes

ferruginous red

head,

thorax, less, and abdomen black 45

Lateral margins ot the median segment not acute ...

. Median segment twice as broad as the length in the

middle, the lateral carine sharply defined. An-

tenne fusco-ferruginous at the base, black at

Apex tee

Median seoment ‘about three times as broad as the

length in the middle, the lateral carme not

sharply defined. Antenne wholly orange

Head, thorax, and abdomen black

Head, thorax, and abdomen more or less red _

Legs black. Punctures of front and pronotum coarse

and confluent longitudinally

Legs ferruginous. Punctures of front and pronotum

not coarse and well separated ..

apical segments of abdomen ferru-

sinous red, head and thorax black ..

Abdomen w holly black, head and sometimes thorax

more or less red...

Two apical segments of the abdomen red ; “fore wings

fuscous except at the apex; sixth dorsal segment

punctured ......

Apical segment of abdomen only red ; ; wings s hy aliue ;

sixth dorsal: segment striate

alicie Turn.

(=P)

*#H. abdominalis Guér.

oO.

*#. peringueyi Sauss.

*E. hottentota Sauss.

EH.

. adelogamia 'Turn.

auriflua Turn.

. torrida Sm.

10.

* ryy

. saussuret Turn.

. canthocera Gerst.

11.

12.

. nnotata Turn.

- erythropoda Turn.

13.

14.

apicipennis Turn.

HH. pyxidata Turn.

The colour in the female is

variable, the head being sometimes ferruginous.

704 MR. R. E. TURNER ON

14. Wings subhyaline; head, pronotum, mesonotum, and

scutellum red; vertex punctured ..................... 2. heterogamia Sauss.

Wings fusco-violaceous REAGENT Ee Sea Rea ee eeeee ne 15.

15. Vertex punctured ; pronotum closely punctured; legs

black ytd. eeepites ae ks, th oe ee eb ae be MM oom TMS

Vertex smooth; pronotum sparsely punctured; legs

TO: ioctl sae CR ReE Sea eC Re oth LL RCE RDO 16.

16. Head red, thorax black oo... eects 2B ruficeps ruficeps Sym.

Thorax more ordess red) es. aise 48 s-c oss cess fecee.caeee) Es ruficeps atopogamia _

{Sauss.

Males.

1. Apical dorsal segment not incised at the apex ........ 2.

Apicil dorsal segment more or less incised at th

ADOT Seactiebiocae Nicoen toca so sieacee nae at wee eames eee 4.

2. Third dorsal segment of the abdomen measured

from the transverse basal furrow distinctly shorter

than its basal width. Abdomen wholly without

Sell Owsmia an: lun ey un ee ene ye eee arene aL cee ee EE. incerta Turn.

_ Third dorsal segment measured from the transverse

basal furrow distinctly longer than its basal

width. Abdomen with very small yellow

TY QUT OS! 2). vain eee eta oe ae Ee ren 3.

3. Apical dorsal segment flattened, with raised margins

and a median carina; pronotum entirely black ... EH. asmarensis Turn.

Apical dorsal segment convex, the margins not raised,

without a distinct median carina; pronotum with

a yellow band on the posterior margin ..,............. E. ametalla Turn.

4, Abdomen entirely black, sometimes with blue sheen . 5.

Abdomen banded with yellow ............. 6.

5. Wings more or less shaded with fuscous or viola-

ceous ; incision of the apical segment half as deep

as its apical width. Length 15-18 mm. ............. EB. ruficeps Sm.

Wings clear hyaline; incision of apical segment

much less than half as deep as its apical width.

AMGemves ih We mane aes to eset aa ade eelgneea eee modosal Gucrs

6. Apical dorsal segment flattened with raised margins ;

head large, the cheeks more than half as broad as

GN OMOV OS Ieate CARE en chasse cArcdeee nek ee er ee

Apical dorsal segment convex, with a median carina ;

head small, the cheeks much less than half as

EH. capitata Sm.

bidadkasstheseyestcarruyasusacae ins cana ene Us

7. Not very slender ; the apical dorsal seement with an

incision nearly as deep as broad at the apex ...... E. spoliata Turn.

Very slender; incision of the apical dorsal seement

uot more than half as deep as its apical breadth... H. longiventris Turn.

Burs (Masa) aticra, sp.n. (Pl. LXXXI. fig. 12; Pl. LX XXII.

fig. 8; Pl. LXX XIII tis! 9%)

2. Nigra; segmentis abdominalibus quinto sextoque rufo~

Jerrugineis ; pedibus albo-pilosus, cateartis albidis; alis nigro-

eeruleis.

Long. 18 mm.

9. Head and thorax coarsely punctured-rugose, the punctures

on the front finer than on the vertex; clypeus very broadly

rounded at the apex, with two or three indistinct teeth on the

margin; scape shining and sparsely punctured, clothed beneath

with long whitish hairs; the nine apical joints of the flagellum

opaque. Front thinly clothed with white hairs, the interantennal

prominence well developed and feebly bilobed. Inner margin of

the eyes slightly sinuate ; posterior ocelli about twice as far from

ye eee

STUDIES IN THE FOSSORIAL WASPS. 705

the eyes as from each other. Pronotum nearly as broad as the

head, widely emarginate anteriorly ; scutellum with a large

triangular rugose area from the base to the narrowly truncated

apex, the sides smooth and opaque. Postscutellum and median

segment smooth and opaque, a few large punctures on the middle

of the postscutellum ; median segment raised towards the median

line, with a narrow margined median groove, the posterior trunca-

tion of the segment coarsely but shallowly punctured, the sides of

the segment closely obliquely striated. Abdomen shining, finely

and very sparsely punctured, the punctures larger on the ventral

than on the dorsal surface, and rather closer at the apex of the

segments than at the base; the apical dorsal segment broadly

rounded, and longitudinally punctured-striate. Second abscissa

of the radius a little shorter than the third; first recurrent

nervure received beyond the middle of the second cubital cell,

second at two-thirds from the base of the third cubital cell.

Radial cell detached from the costa for about half its length.

Hab. British East Africa, Makindu, 3300 ft. (S. A. Weave);

April 5-7, 1911 (4. & BR. C.).

This fine species does not seemn to be very nearly allied to any

other, being well distinguished from the rujficeps group by the

very coarse sculpture of the head and thorax. The basal joint of

the posterior tarsi is furnished with a closely-set comb of small

spines beneath.

The description of Cosiia donaldsoni Fox corresponds rather

closely to this species, but the clypeus is tridentate on the apical

margin, not rounded, with indistinct teeth as in the present

species, and I think Fox was too careful a worker to have con=

fused the genera.

Huis (Mesa) AURIFLUA, sp. n.

Q. Nigra ; mandibulis basi, pronoto, mesonoto scutelloque rufo-

Serrugineis ; alis infuscatis.

Long. 12 mm.

@. Head and pronotum closely and rather strongly punctured ;

mesonotum and scutellum much more sparsely punctured, smooth

in the middle; pleurz coarsely punctured ; median segment finely

and closely punctured, the median groove narrow and shallow,

margined by low carine, the posterior truncation more shallowly

punctured. Abdomen rather closely and not very finely punctured,

the apical segment closely longitudinally striated. Sides of the

median segment obliquely striated.. Clypeus transverse at the

apex. Hyes very teebly and widely emarginate on the inner

margin; posterior ocelli further from the eyes than from each

other. Head more than half as broad again as long, broader than

the thorax. Pronotum short, not as long in the middle as the

scutellum, the anterior margin straight, the posterior margin

widely arched. Pubescence sparse and whitish, calcaria whitish.

Radial cell only separated from the costa at the apex, which is

subtruneate; first abscissa of the radius as long as the second,

706 MR. R. E, TURNER ON

but shorter than the third; recurrent nervures received close to

the middle of the second and third cubital cells.

Hab. Johannesburg, Transvaal (Kobrow). Received from

Dr. Brauns.

Nearly allied to adelogamia Turn. and diapherogamia Sauss.

From the former it may be distinguished by the entirely black

head, and by the closer and more even puncturation of the head

and pronotum, the vertex being almost smooth in adelogamia;

from diapherogamia it may be distinguished by the much shorter

pronotum, the stronger and closer puncturation, and the colour of

the head. #. hova has the pronotum longer than in the present

species or adelogamia.

The comb on the underside of the basal joint of the posterior

tarsi is present, but the teeth are few. The upper spur of the

hind tibiz is strongly bent near the base.

Eis (Mesa) ADELoGAMIA Turn.

Plesia (Mesa) adelogamia Turn. Ann. & Mag. Nat. Hist. (8) i.

p. 503 (1908), °.

Hab. Maseru, Basutoland; Lichtenburg, Transvaal.

This is nearly allied to Z. auriflua, but differs as noticed in the

key.

Eis (Mesa) ruricers Sm. (PI. LX XXII. figs. 9, 10;

Pl. LXXXITI. figs. 2, 4, 10, 15, 16.)

Myzine ruficeps Sm. Cat. Hym. B. M. iii. p. 75 (1855), 9 ;

Turner, Ann. & Mag. Nat. Hist. (8) i. p. 508, ¢; Turner, Ann.

& Mag. Nat. (8) vii. p. 304 (1911), 35 @.

Eis (M&sA) RUFICEPS, subsp. ATOPOGAMIA Sauss.

Plesia( Mesa) atopogamia Sauss.in Grandidier, Hist. Madagascar,

xx. p. 244 (1892), ©.

Plesia (Mesa) diapherogamia Sauss.; Distant, Naturalist in the

Transvaal, p. 225 (1892), °.

Plesia (Mesa) disjuncta Turn. Ann. & Mag. Nat. Hist. (8) i.

p- 902 (1908), ¢.

Llis (Mesa) ruficeps, subsp. atopogamia and diapherogamia Turn.

Ann. & Mag. Nat. Hist. (8) vii. p. 304 (CISIEDS oi QS

Hab. Zanzibar ; Nyasaland; Transvaal.

The colour-differences between atopogamia and diapherogamia

are not constant, though in the former the mesonotum is usually

red and in the latter black. The wings of the male are darker in

specimens from Nyasaland than in those from the Transvaal.

A single male received from Harar, in 8. Abyssinia, has the wings

entirely hyaline, slightly iridescent.

Eis (MzsA) HETEROGAMIA Sauss.

Plesia (Mesa) heterogamia Sauss. in Grandidier, Hist. Mada-

gascar, xx. p. 244 (1892), 2.

Hab. Delagoa Bay ; Manica; South Nyasaland.

STUDIES IN THE FOSSORIAL WASPS. 707

Exits (Mss) nova Turn.

‘Plesia (Mesa) hova Turn. Ann. & Mag. Nat. Hist. (8) i. p. 504

(1208), 2. |

_ Hab. Madagascar.

Probably the female of nodosa Guér.

*His (MEsA) ABDOMINALIS Gueér.

Plesia abdominalis Guér. Rev. Zool. i. p. 57 (1838), 2

Plesia (Mesa) abdominalis Sauss. in Grandidier, Hist. Madagas-

car, xx. p. 244 (1892), ©.

Hab. South Africa.

Eis (MEsA) APICIPENNIS, sp. n.

©. Nigra; segmentis abdominalibus quarto apice, quinto sextoque

rufo-ferrugineis ; mandibulis basi antennisque ferrugineis, cal-

carus albidis ; alis fuscis, apice anguste hyalinis.

Long. 10 mm.

@. Clypeus finely punctured, broadly rounded at the apex.

Head sparsely and rather finely punctured, shining, almost

smooth round the anterior ocellus; interantennal prominence

bilobed, divided by a longitudinal sulcus which does not reach the

anterior ocellus. Scape shining, very sparsely punctured ; flagel-

lum opaque, the two basal joints shining. Posterior ocelli nearly

as far from each other as from the eyes. Pronotum and pleure

closely but not coarsely punctured; mesonotum and scutellum,

more sparsely punctured; median segment opaque, finely

punctured, the punctures more or less confluent longitudinally,

the median groove shallow and not distinctly margined. Abdo-

men shining, finely and rather closely punctured, with a very

short petiole, the first segment broadly truncate at the base ; the

sixth dorsal segment punctured, rounded at the apex. The sides

of the median segment are closely obliquely striated. Third

abscissa of the radius at least half as long again as the second ;

first recurrent nervure received just beyond the middle of the

second cubital cell, second at two-thirds from the base of the

third cubital cell. Radial cell not distinctly separated from the

costa, narrowly rounded at the apex.

Hab. British East Africa, Makindu, 3300 ft. (S. 4. Weave);

Auprilg fy hoi CA ee he. C.):

Nearly allied to #. pyaidata Turn. from N.E. Rhodesia, but

differs in the broader shape of the third cubital cell, the position

of the second recurrent nervure, the colour of the scape, of the

wings, and of the fourth and fifth abdominal segments, in the

finer and sparser puncturation and in the sculpture of the

pygidium. The wings are clear hyaline beyond the radial and

cubital cells. Basal joint of the posterior tarsi with a scopa of

white hairs beneath ; spur of the posterior tibia bent near the

base.

708 MR. R. E. TURNER ON

Eis (MzsA) rorripa Sm.

Myzihe torrida Sm. Desc. new spec. Hymen. p. 178 (1879), 9.

2. Nigra; mandibulis basi, scapo apice abdomineque toto

Ferrugeneis ; tegulis testaceis; tibiis tarsisque fusco-ferrugineis ;

alis hyalinis, venis nigris.

Long. 12 mm.

Q. Clypeus short, transverse at the apex, subcarinate in the

middle, finely punctured at the base, smooth at the apex, Front,

pronotum, and mesopleure closely and strongly punctured, vertex

a little more sparsely punctured; mesonotum and scutellum.

coarsely but sparsely punctured; pronotum as long as the

scutellum. Median segment twice as broad as the length in the

middle, not Hag almost smooth, but not shining ; the median

groove narrow but! well defined, with the margins of the groove

raised. Abdomen shining, il a few scattered punctures, the

apical dorsal segment finely longitudinally striated. Basal joint

of the posterior ‘tarsi unarmed beneath, with a scopa of very fine:

hairs. Radial cell detached from the costa at the apex, third

abscissa of the radius longer than the second, but & little shorter

than the first, second recurrent nervure received just beyond two-

thirds from the base of the third cubital cell. .

Hab. Gambia (ex coll. Shuckard).

* Eis (MEsA) PERINGUEYI Sauss.

Plesia (Mesa) peringueyi Sauss, in Grandidier, Hist. Madagas-

ear, xx. p. 245 (1892), ©.

“ Areola radialis apice minute truncata. Secunda v. recurr. in

ipso medio margine tertiz ar. cubitalis exserta. Metatarsus

posticus subtus scopa spinarum brevium instructus. Majuscula,

nigra, nitida, cinereo-hirta. Caput et thorax cribrosa. Caput

validum, quam pronoto paulo latius. Metathorax leeviusculus,

superne tenuiter punctatus, subtiliter carinatus (carina a latere

visa subbituberculata) utrinque pago polito; ejus facies postica

plana, subrugulata ; metapleura strigata. Epipygium elongato-

trigonale, punctatum, Spine tib. post. nigre, acute. Metatarsus

posticus subtus pectinatus, pilis vel spinis albidis intermixtis,

spinisque nonnullis dilatatis. Ale bruneo-nebulose, venis fuscis ;

2* ar. cubit. intus valde acuta; 2® vena recurrens tyansversalis,

cum Vv. discoidali angulum fere rectum etficiens.”

*Euis (Musa) norrentora Sauss.

Plesia (Mesa) hottentota Sauss. in Grandidier, Hist. Madagascar,

xx. p. 245 (1892), Q.

“ Areola radialis apice minute truncata. Secunda v. recurr. in

ipso medio margine tertie ar. cubitalis exserta. Metatarsus

posticus subtus scopa spinarum brevium instructus. Minor,

nigra, cinereo-hirta. Antenne imo apice flavo. Caput et thorax

sat tenuiter cribrosa. Caput quam thorax vix latius. Meta-

thorax lviusculus, subtilissime punctato-rugulatus, superne-

STUDIES IN THE FOSSORIAL WASPS. 709

obsolete roundato-carinatus; facie postica punctulata, supra

distincte angulata, obtusangula, fere rectangula. Epipygium

trigonale, striolatum, margine levi. Spin tib. post. albescentes,

squamose. Alze subhyalinz venis bruneis et fuscis ; parte apicali

nebuiosa, 2° ar. cubit. intus breviter acute producta; 2* vena

recurrens arcuata, obliqua, In alis posticis v. discoidalis longius

ultra venulam transverso-discoidalem furcata, Long 14 mill. ;

al, 10 mill, (Africa meridionalis).”

*EiIs (Mesa) capensis Lep.

Tiphia capensis Lep. in Hist, Nat, Insect., Hym. iii, p, 554

(1845), &.

ee Caput nigrum, supra nigro, subtus rufo subvillosum, An-

tenne nigre, articulo primo nigro hirto. Thorax niger rufa

subhirtus. Abdomen nigrum, ay eaaiinnes Pedes nigri, rufopallido

subvillosi, femoribus duobus posticis angulatis compressisque,

Ale rufo- fuscee, nervuris costiaque rufo-fuscis ; squama nigra,

Cellula radialis elucee Femina.” y

“« Long. 7 lignes.

“ Cap de Bonne Espérance. Musée de M. Serville.”

The figure shows that this species is an //¢s.

Exis (Mesa) rnnorara Turn.

Plesia (Mesa) innotata Turn. Ann. & Mag, Nat. Hist. (8) i,

p- 906 (1908), 2.

Hab. Loangwa River, N.E. Rhodesia ; 8. Nyasaland.

Exis (Musa) saussurer Turn.

Plesia (Mesa) saussurei Turn, Trans. Ent. Soc. London, 1910,

jon See WS

Hab. Madagascar.

This is near /. canthocera, but the median segment is longer in

proportion and much. more distinctly margined; the colour of

the antenne is also different.

Huts (Musa) XANTHOCERA Gerst,

Myzine xanthocera Gerst. Arch. f. Naturg, xxxvii. p, 353 (1870),

2; v.d. Decken, Reise in Ost-Afrika, Gliedethiere, p. 339, pl. 14,

fig. 5 (1873), 2.

Plesia (Mesa) xanthocera Sauss, in Grandidier, Hist, Madagas-

car, xx. p. 245 (1892), ©.

Hab. Howick, Natal; Zoutpansberg, Transvaal; Mozambique ;

Harar, Abyssinia. .

Exis (Mzsa) EryrtHROPODA Turn,

Plesia (Mesa) erythropoda Turn. Ann. & Mag, Nat. Hist. (8) i.

p- 505 (1908), @.

Hab. Lake Ngami.

The scopa beneath the basal joint of the hind tarsus is rather

710 MR. R. E. TURNER ON

coarse at the base, but I do not think that any spines are

present.

Eis (Mzsa) INCERTA, sp. n.

3. Niger, cano-pilosus ; clypeo macula mediana minuta, tibwis

anticis supra pallide flavis; tegulis pedibusque fuscis; alis

hyalinis, venis nigris ; pygidio haud inciso.

Long. 13 mm.

¢. Front coarsely reticulated, the vertex punctured; thorax

closely but not coarsely punctured, median segment punctured-

rugose, abdomen finely and closely punctured. Pronotum shorter

than the scutellum, the anterior margins straight, the angles

subacute. First abdominal segment petiolate, the narrow petiole

scarcely more than half as long as the dilated apical portion of

the segment, which is constricted at the apex; second segment

twice as broad at the apex as at the base, all the segments slightly

constricted at the base. Apical dorsal segment sparsely punctured,

without an incision, somewhat convex, subcarinate in the middle,

pointed at the apex, the lateral margins raised near the apex.

Third abscissa of the radius a little longer than the second, which

is fully twice as long as the fourth. First transverse cubital

nervure oblique, sharply bent close to the cubitus, second

recurrent nervure received just before one-third from the base of

the third cubital cell, curved outwards below the middle, the ends

slightly curved inwards.

Hab. Howick, Natal (J. P. Cregoe); Cape Colony.

This is very near the description of Plesta carbonaria Cam., but

in that species the seventh dorsal ‘segment is said to be shortly

incised at the apex, and the fourth abscissa of the radius is almost

as long as the second or third.

Exis (Mss) capirata Sm.

Myzine capitata Sm. Cat. Hym. B. M. i. p. 74 (1855), 3

6. Niger, albido-pilosus ; elypeo, mandibulis, linea obliqua

utringue supra antennas, pronoto fascia angusta postice, tegulis

basi, segmento dorsali primo fascia apicali emarginata, segmentis

dorsalibus et ventralibus 2-6 fascia bisinuata angusta flavis ;

‘pedibus flavis nigro-variegatis ; alis hyalinis, venis testacers.

Long. 14-17 mm.

3S. Head large, much prester than the thorax, the cheeks more

than half as broad as the eyes; antenne stout, the apical joint

truncate at the apex. Head rugose, thorax closely punctured,

median segment punctured-rugose; abdomen shining, glossed

with blue, very finely and closely punctured. Pronotum shorter

than the scutellum, slightly narrowed anteriorly, the anterior

margin straight, the angles not prominent, posterior margin

widely arched. First abdominal segment petiolate, the petiole

only about half as long as the rather strongly swollen apical

STUDIES IN THE FOSSORIAL WASPS. 711

portion, the apex slightly constricted. Apical dorsal segment

flattened, subcarinate in the middle, the lateral margins raised

and nearly parallel, the apical incision subtriangular, not quite as

deep as the apical width. Second and third abscissve of the radius

nearly equal in length ; second recurrent nervure strongly curved

outwards below the middle, joining the cubitus just before one-

fourth from the base of the third cubital cell.

Hab. Johannesburg, Transvaal; Kroonstad.

The type is much damaged and without wings; the details of

neuration are taken from a more recent specimen in which the

cubitus of the hind wing on the right side is almost interstitial

with the transverse median nervure, but is further removed

towards the base on the left side.

Exis (Musa) spouiata, sp. n.

3. Niger; mandibulis, clypeo macula parva nigra utrinque,

linea utrinque supra antennas, pronoto angulis anticis et fascia

postice, tegulis, segmento dorsali primo fascia angusta apicali,

segmentis dorsalibus et ventralibus 2-6 fascia bisinuata angusta,

coxis apice, femoribus posticis supra, tibiis anticis et intermediis

supra tarsisque subtus pallide flavis ; pedibus rufo-testaceis ; alis

hyalinis, venis nigris, stigmate rufo-testaceo.

Long. 13 mm.

3g. Clypeus short and broad, rather narrowly produced in the

middle and very feebly emarginate at the apex. Hyes widely

emarginate ; cheeks very nruch narrower than the eyes ; posterior

ocelli more than half as far again from the eyes as from each

other. Antennal tubercles well developed ; antennz longer than

the head, thorax, and median segment combined, of even thick-

ness throughout. Head punctured-rugose, much wider than the

thorax; the whole thorax finely and closely punctured, with

sparse white pubescence ; pronotum a little longer in the middle

than the scutellum, narrowed anteriorly, the anterior margin

straight, posterior margin strongly arched. Median segment

rounded posteriorly, not truncate, very closely but not coarsely

punctured. Abdomen slender, slightly shining, very finely and

sparsely punctured, petiolate; the petiole occupying the basal

third of the first segment, the apical two-thirds elongate pyriform,

the whole segment half as long again as the second, which is

gradually broadened from the base, a little longer than the apical

width ; third segment broader than long. Apical dorsal segment

slightly convex, subcarinate longitudinally in the middle, shining,

with a few large punctures, the apical emargination almost as

deep as its breadth at the apex. Radial cell pointed; second

abscissa of the radius shorter than the third, but much longer

than the first; second recurrent nervure received at one-third

from the base of the third cubital cell.

Hab. Algoa Bay, Cape Colony; March (Dr. Brauns).

Nearly allied to #. capitata Sm., but in that species the head

712 MR. R. E. TURNER ON

is larger, the cheeks much broader, and the first abdominal

segment a little shorter and more swollen towards the apex.

The apical dorsal segment in capitata is flat, not convex, and

has the sides distinctly raised into marginal carine.

Enis (Mesa) LONGIVENTRIS, sp. n. (Pl UXXXT. fig. 13;

Pl. LXXXITI, fig. 11.)

3. Niger; mandibulis, elypeo, linew obliqua utringue supra

antennas, pronoto fuscia anguste postice, segmento dorsali primo

Fascia angusta api ‘cali, se gmentis dorsalibus et ventralibus 2-6 fascia

angusta apicali bisinwata, coxis subtus. tibiis tarsisque anterioribus

supra, intermediisque subius albidajlavis; alis hyalinis, venis

negris.

Long. 10-13 mm.

é. Very slender. Head rugose; thorax strongly and closely

punctured, median segment punctured-rugose, Antenne rather

slender, as long as the head, thorax, and median segment com-

bined, of even thickness throughout. Pronoatum longer than the

scutellum, the sides almost p: avallel, the anterior margin straight,

with acute angles, the posterior margin widely arched. Sides of

the head and thorax rather thicky clothed with long white

pubescence. Median segment longer than broad, rounded pos-

teriorly. Abdomen slender, shining, very finely punctured ; the

basal segment as long as the second and third combined, petiolate,

the narrow petiole nearly as long as the slightly swollen apical

portion ; second segment br oadened from the base, about equal

in length to the third, the segments scarcely constricted at the

base; the apical dorsal segment convex, subearinate in the

middle, sparsely punctur ed, the punctures large, the apiece al

incision shallow, not as deep as its breadth at the apex. Wings

reaching to the apex of the fourth dorsal segment; second and

third abscisse of the radius usually about equal in length, the

fourth distinctly shorter; second recurrent nervure received just

before one-third from the base of the third cubital cell, curved

outwards in the middle.

Hab. Willowmore, Cape Colony (Di, Brawns).

This corresponds rather closely with the description of Plesia

incisa Cam,, but in that description the incision of the apical

dorsal segment is said to be twice longer than wide, and some of

the details of neuration are not quite the same. But the latter

character is of little importance in the genus, especially as to the

length of the seeond and third abscisste of the radius. Many

yariations oecur which are not of specific value.

Ets (Mesa) AsMArensis Turn,

Plesia asmarensis Turn, Ann. & Mag. Nat, Hist. (8) in, p: 481

(1909), ¢

Hab. Erythvea,

STUDIES IN THE FOSSORIAL WASPS. 713

Enis (Musa) ameraria Turn.

Lilis (Mesa) ametalla Turn. Ann. & Mag. Nat. Hist. (8) vii.

p. 305 (1911),

Hab. 5. Nyasaland.

Almost certainly the male of innotata,

Exis (Musa) Noposa Guér.

Myzine nodosa Guér, Dict, pitt. hist, nat. v. p. 584 (1837), 3;

Sauss. in Grandidier Hist. Madagascar, xx. p. 240 (1892), ¢.

Hab. Madagascar.

* Eis (Musa) cLAVATA Sauss,

Myzine clavata Sauss, in Grandidier, Hist, Madagascar, xx.

p. 242 (1892), ¢.

Hah. Transvaal,

- *Hinis (Musa) CARBONARIA Cam.

Plesia carbonaria Cam. Rec, Albany Mus. i, p, 317 (1905), 3.

Hab. Dunbrody, Oape Colony.

*Hiis (Mesa) RETICULATA Cam.

Plesia reticulata Cam. Rec, Albany Mus, i. p, 300 (1905), 3.

Hab. Brak Kloof, Cape Colony,

*Euis (Mesa) RUrO-FEMORATA Cam,

Plesia rufo-femorata Cam, Ree, Albany Mus, i.p. 298 (1905), 3

Hab. O’okiep, Cape Colony,

* Eis (Musa) rvc1sa Cam,

Plesia incisa Cam. Ree, Albany Mus. i, p, 320 (1905), 3.

Hab. Dunbrody, Cape Colony, .

Key to the Oriental Species of Elis (Mesa),

Kemales.

1. Sixth dorsal sezment ey Shilatedeeneseet cs 2.

Sixth dorsal segment punctured... ' 8.

2. Abdomen more or less ferruginows 3:

Abdomen wholly black 4 4,

3. Clypeus with a median carina ; “two ‘apical ‘abdominal

segments and the apex of the fourth black............ HL. dimidiata Guéy.

Clypeus without a distinct carina; apical abdominal

segment only black . ay vs... HH, mandalensis Magy.

4, Median seement distinctly margined poster iorly y. sa LE. fuscipennis 8m.

Median segment not Halietyy ened shag Brea 5.

Bee WInes LUSCOUB! vss. 272 see ta eoes Cece 6.

Wings subhyaline... ite

6. Pronotum longitudinally rugose ; “second abscissa of

the radius as long as the third. Length 16 mm.... EH. mandibularis Sm.

Pronotum coarsely punctured ; second abscissa of the

radius much shorter than the third. Lengthl0 mm. E. petiolata Sm.

Proc. Zoo. Soc.—1912, No. XLVII. A7

714 MR. R. E. TURNER ON

- Propleure sparsely punctured ......... 0.06000 ccc cee cee aee

Propleur finely striated.. seugepas Weve

8. Abdomen not marked with ‘yellow Rr ERR SA

Abdominal segments with yellow apical bands .........

9. Head black ; abdomen black or black and ferrugimous.

Head red; abdomen steel-blue ..................0.. cee vee ees

. Abdomen mostly ferruginous..

Abdomen entirely black, or with the ‘apical. segment

only ferruginous ..... SP

. Four basal seoments of the abdomen ‘ferruginous: Beaks

Four apical segments of the abdomen ferruginous eee

Apical segment of the abdomen ferruginous

Apical, segment of the abdomen black

Mesonotum closely punctured 0.0.0.0... ces ece eee eee ee

Mesonotum shining, almost impunctate ...............6..

Legs ferruginous; head coarsely and closely punctured.

Legs black ; head sparsely punctured

Myzine anthracina

recorded by Bingham

FE. claripennis Bingh.

LE, ustulata Turn.

*H. picticollis Mor.

10.

E.. tricolor Sm.

11.

12.

EF. bengalensis Cam.

E. apimacula Cam.

13.

14,

*#. fedtschenkot Sauss.

*H. dubia Mor.

FE. rothneyi Cam.

LE. opacifrons Turn.

as Indian

Australian and belongs to the genus Anthobosca; and Myzine

combusta also recorded as Indian is undoubtedly West Indian,

and a true Elis.

Males.

1. First dorsal segment elongate, peice more or less

nodose at the ADER! ccs.ncc Pa eer een ah ae

First dorsal segment subsessile, “not nodose at the

apex, broader “at the apex than LOG AG See eer A cee

. Seventh dorsal segment distinctly incised at the apex ;

abdomen wholly black.. :

Seventh dorsal segment not incised at the apex.

3. Legs black; seventh dorsal segment smooth and

shining at the BPE Kat dvr ssctn cack deans muck sa meas gee

Legs tusco - ferruginous; seventh dorsal segment

Coamselhympuncbured vance neces sare concer nee cee

4. Abdomen wholly black, without yellow une eS

sometimes glossed with blue .. : a

Abdomen more or less marked with yellow ..

5. Median segment without a distinct sulcus ; “pyg idial

area clearly defined, with marginal caringe reaching

nearly to the base of the seoment . Bis centss

Median segment with a distinct. median “sulcus:

pygidial area less clearly defined, the marginal

carine lower and only on the apical third of the

segment .

6. First abdominal segment very little, if ab ‘all, longer

than the hind femur and trochanter combined ......

First abdominal segment much longer than the hind

femur and trochanter combined ........................

7. Head punctured-rugose; pronotum without a yellow

LONG Waa eer ire hs Bossa Anh cate omen ens tights. cag con sctsn

Head punctured ; pronotum with a yellow band on the

posterior margin .

8. Wings hyaline; “pronotum ‘with a yellow band on the

posterior margin, closely punctured ..

Wings pale fusco-hyaline ; pronotum without a | yellow

band, rather sparsely punctur ed .

. Narrow. petiole of first. abdominal segment. “scarcely

more than half as long as the swollen apical portion

ot the segment .

Narrow petiole of first abdominal ‘segment “nearly a as

long as the swollen apical portion of the segment ..

Ws)

E. dimidiaticornis

[Bingh.

E. burmanica Bingh.

E. leta Bingh,

Ft. dimidiata Guér.

EH. mandibularis Sm.

HL. nurseit Turn.

EE. mandalensis Magy.

FL. extensa Turn.

EE. petiolata Sm.

Ei. claripennis Bingh.

Myzine pallida Sm. and Myzine orientalis Sm., placed by Bing-

ham and Smith in Wyzine, together with many species of lis,

STUDIES IN THE FOSSORIAL WASPS. Hala:

belong to the genus Zswara Westw., which also belongs to the

Elidine.

Exis (Mesa) pimip1ata Guér.

Myzine dimidiata Guér. Dict. pitt. hist. nat. v. p. 584 (1837),

Methoca orientalis Sm. Cat. Hym. B. M. iii, p. 66 (1855), 3

(nec Smith, 1875).

Myzine madraspatana Sm, Cat. Hym, B. M. iii. p, 72 (1855), @.

Myzine violaceipennis Cam, Mem. Manchester Lit. & Phil. Soe.

xlii. p. 21 (1898), ©,

Hab. The whole of India, except the North-West.

2. Head and thorax coarsely rugosely punctured, smooth

round the anterior ocellus and at the base of the mesonotum;

median segment opaque, with a narrow median gulcus in which is

a low carina, the sides of the sulcus raised and forming low

carine, the segment not margined at the base of the posterior

truncation. Radial cell distinctly separated from the costa for

more than half its length, third abscissa of the radius very

distinctly longer than the second. Sixth dorsal segment finely

longitudinally striated ; basal joint of hind tarsi with a scopa of

short hairs beneath.

Black ; second, third, and the base of the fourth abdominal

segments ferruginous. Oalcaria whitish ; wings fusco-violaceous.

3. Head and thorax closely punctured; median segment

rugose ; basal third of first abdominal segment forming a narrow

petiole, apical two-thirds swollen and slightly constricted at the

apex, the whole segment more than half as long again as the

second segment. Abdomen shining, very finely punctured ;

pygidial area coarsely punctured, fully twice as long as broad,

the sides raised and forming carine, a well-defined median carina,

the apex narrowly truncate, not emarginate.

Black ; the abdomen slightly glossed with blue; base of the

mandibles, fore tibie in front, fore tarsi in front, base of the

tegule and the hind margin of the pronotum narrowly and

obscurely pale yellow. Wings hyaline at the base, fusco-hyaline

beyond the basal nervure.

Length 17 mm,

The association of the sexes was suggested by me in 1908, and

positive proof was published by Mr. Lefroy a year later, a pair

having been taken in coitu by Mr. Dutt at Pusa.

Eis (Musa) ManDrBuLaris Sm,

Methoca mandibularis Sm. Trans, Ent, Soc, London, p. 301

(1869), 3.

Plesia (Mesa) mandibulavis Turn. Ann. & Mag, Nat. Hist, (8)

i. p: 009 (1908); 3.

Plesia (Mesa) purpureipennis Turn. Ann, & Mag, Nat. Hist.

(8) i. p. 508 (1908), @.

This will almost certainly prove to be the Chinese subspecies

AT*

716 MR. R. E. TURNER ON

of 1. dimidiata Guér., from which the female differs in the

entirely black abdomen and the greater length of the second

abscissa of the radius, which is nearly as long as the third. The

male differs from dimidiata in having the second abscissa of the

radius very distinctly longer than the third, in the paler colour

of the apical portion of the wings, in the presence of a distinct

longitudinal suleus on the medieth segment, and in the somewhat

shorter and broader form of the first abdominal segment. The

pygidial area of the male is also wider and less distinctly margined

than in dimidiata.

Ifab. Shanghai.

Eis (Musa) BENGALENSIS Cam.

Myzine Ee Cam. Mem. Manchester Ties & Phil. Soe.

xiii. p. 21,(1898), 2.

The four basal abdominal segments are ferruginous ; the wings

violaceous, the base of the hind wings hyaline. Pronotum

sparsely but coarsely punctured.

Length 15 mm.

This is quite distinct from dimidiata, and seems, as Cameron

suggests, to be more nearly related to mandalensis.

Eis (Musa) MANDALENSIS Magy.

Plesia mandadlensis Magy. Ann. Mus. Civ. Gen. (2) xii. p. 257

(1892), 2°

@. The five basal abdominal segments are ferruginous.

Calcaria and spines of the hind tibie whitish. Wings hyaline,

dark fusco-hyaline beyond the basal nervure of the fore wing

to the apex, the apex of the hind wing fusco-hyaline. Pronotum

closely and somewhat coarsely punctured; sixth dorsal segment

finely longitudinally striated.

On Head and thorax closely and rather finely punctured ;

median segment punctured-rugose ; abdomen shining, micro-

scopically punctured, First abdominal segment scarcely longer

than the hind femur and trochanter combined, the narrow petiole

a little more than half as long as the moderately swollen apical

portion of the segment; second segment longer than the third

by about one-quarter. Seventh dorsal segment pointed, not

incised ; pygidial area well defined, very narrow, coarsely

punctured and with a well-marked median carina. Third

abscissa of the radius distinctly longer than the second.

Black; mandibles at the base, clypeus, apex of the inter-

antennal prominence, posterior margin of the pronotum, tegule,

an apical band on dorsal segments 1—6, strongly bisinuate on

segments 2-6 and on ventral segments 2-6, tarsi, fore and

intermediate tibiz in front, and the base of the hind tibiz yellow.

Wings hyaline, nervures fuscous.

Length, 2 10 mm., ¢ 10-11 mm.

Hab. Mandalay, Burma.

Taken in copuld by Colonel Bingham.

STUDIES IN THE FOSSORIAL WASPS. Tit

Enis (Mesa) rormneyi Cam.

Myzine rothneyi Cam. Ann. & Mag. Nat. Hist. (7) x. p. 88

(1902); 2.

@. This fine species is easily distinguished, being black with

ferruginous legs; the wings fusco-hyaline, flushed with purple.

The sixth dorsal segment is coarsely punctured, the punctures

tending to become confluent longitudinally towards the apex.

Head and thorax very coarsely punctured, rugose on the

pronotum.

Hab. Khasi Hills, Assam.

Eis (Mzsa) FUSCIPENNIS Sm.

Myzine fuscipennis Sm. Cat. Hym. B. M. iii. p. 72 (1855), 2 ;

Bingh. Fauna Brit. India, Hymen. i. p. 67 (1897), @.

Q@. This species may be distinguished from the female of

wetiolata Sm., which it closely resembles, by the sharply margined

median segment, the posterior slope of which is abrupt and steep,

not gradual as in petiolata. The colour is black; the calearia

whitish, mandibles fusco-ferruginous, wings fuscous. The spines

on the outer margin of the hind tibiz are black in fuscipennis,

whitish in petiolata.

Length 12 mm.

Hab. India.

The type is unique in the British Museum collection and was

obtained from Shuckard in exchange, so that the locality is

uncertain. Bingham’s description is taken from the type, but

he evidently confused the species with peticlata, a specimen of

which is labelled fuscipennis by him in the British Museum

collection. I have not seen the specimens he records from

Burma, but consider it very doubtful if they belong to this

species.

Eis (Mzsa) PETIOLATA Sm.

Myzine petiolata Sm. Cat. Hym. B. M. iii. p. 72 (1855), 3.

Myzine ceylonica Cam. Ann. & Mag. Nat. Hist. (7) v. p. 18

(1900), 2.

Plesia (Mesa) petiolata Turn. Ann. & Mag. Nat. Hist. (8) 1.

p- 512 (1908), d 2.

Q. Differs from fuscipennis Sm. in the absence of a distinct

margin separating the dorsal surface of the median segment

from the surface of the posterior slope and in the white spines of

the hind tibize. From claripennis Bingh. it differs in the fuscous

colour of the wings, the somewhat coarser puncturation, and the

lesser length of the second abscissa of the radius, which is only

about half as long as the third in petiolata and almost or quite as

long as the third in claripennis. In fuscipennis the second

abscissa of the radius is shorter than in petielata, being distinctly

less than half as long as the third.

718 : MR. R. E. TURNER ON

3. This is very near the male of claripennis, but may be

distinguished by the less elongate petiole, the narrow basal

portion of which is scarcely more than half as long as the.

moderately swollen apical portion of the segment.

Length, ¢ 2, 10 mm.

fab. Bengal, Bombay, and Ceylon.

Eis (Mesa) CLARIPENNIS Bingh.

Myzine claripennis Bingh. Fauna Brit. India, Hymen. i. p. 68

(189i) 9"

Myzine hortata Nurse, Journ. Bombay Nat. Hist. Soc. xiv.

p. 81 (1902), 2

@. Differs from petiolata as noticed under that species.

Bingham’s description of the species is inaccurate as to the

median segment, and this has misled Nurse. The segment is not

smooth ane Sines and the longitudinal impression, though

not very long, cannot be said to be tr ‘angular.

¢. The male, which has not been previously. described, closely

resembles petiolata Sm. Head and thorax closely punctured,

most coarsely on the front, median segment rugose; abdomen

very slender, shining, minutely punctured. First abdominal

segment more than half as long again as the hind femur, the

narrow petiole very little shorter than the feebly swollen apical

portion ; second segment very narrow at the base, nearly half as

long again as the third segment. Seventh dor sal Segment not

incised at the apex, pointed, convex and without a distinct

pygidial area.

Black ; mandibles, clypeus, the apex of the interantennal

prominence, posterior margin of the pronotum, an apical narrow

band on dorsal segments de 6, strongly bisinuate on segments

9-6, base of the tegulze, fore tarsi, anterior and intermediate

tibiee in front, base of hind tibie and base of intermediate and

hind tarsi pale yellow. Wings hyaline, nervures fuscous.

Length, ¢ 8mm., 2? 10 mm.

Hab. Burma, Ceylon, Bengal, and Deesa.

The male has the two basal abdominal segments more slender

than in petiolata, the first with the apical portion less swollen,

the basal narrow. petiole longer in Propersony the second

narrower at the base.

Exits (Mesa) usrunara Turn.

Plesia (Mesa) uistulata Turn. Ann. & Mag. Nat. Hist. (8) i.

polo 1908), Q.

Q. This is nearest to claripennis Bingh., but is a larger and

more robust species, somewhat more closely punctured, with the

wings distinctly darker and the punctures on the abdomen

larger.

Hab. Yunzalin Valley, Tenasserim.

STUDIES IN THE FOSSORIAL WASPS. 719

Eis (Mzsa) opactrrons Turn.

. Plesia (Mesa) opacifrons Turn. Ann. & Mag. Nat. Hist. (8)

1, p. 509 (1908), °.

2. This black species may be distinguished by the very sparse

puncturation of the head, pronotum, and sixth dorsal segment.

The wings are pale fusco-hyaline. It is a larger and more robust

species than petiolata.

Hab. Salwen Valley, Tenasserim.

*Eis (Mzsa) pusra Mor.

Plesia dubia Mor. Hor. Soc. Ent. Ross. xxiv. p. 627 (1890), 2.

This species is black, with the apical abdominal segment,

tarsi, hind tibiz, mandibles, and antenne beneath ferruginous,

the mesonotum shining and almost impunctate, the sixth dorsal

segment finely and closely punetured.

Length 113 mm.

Hab. Turkestan.

Puesta (Mesa) APIMACULA Cam.

Myzine apimacula Cam. Journ. Bombay Nat. Hist. Soc. xiv.

p-. 272 (1902), 2.

2. This is allied to dubia Mor., but differs in having the four

apical abdominal segments ferruginous, the wings paler, and the

puncturation somewhat different.

Hab. Deesa, N.W. India.

The male described as belonging to this species by Colonel

Nurse is Pecilotiphia albomaculata Cam., but has three cubital

cells instead of two as in the type of the species, which is

evidently an aberration from the usual neuration. The male

has the form of a Myzine, not of a Mesa, but it is by no means

improbable that Nurse is correct in the association of the sexes,

though he informs me that he does not recollect his reasons for

placing them together. ‘The apical dorsal segment is deeply

incised as in Myzine. Cameron’s action in forming a new genus

on a specimen with abnormal neuration is quite unjustified and

due to ignorance of the variable character of neuration in the

Scoliide, but I am at present doubtful if Pecilotiphia should be

treated as a synonym of Myzine or of Elis (Mesa).

“Eis (Misa) FEDTSCHENKOI Sauss.

Plesia fedischenkot Saussure in Fedtschenko: Turkestan,

Scolude, p. 29 (1880), 2 (Plesia tartara on plate).

Plesia tartara Sauss. |. ce. pl. ii. fig. 12, 9.

This seems to differ from dubiw Mor. in the coarser and closer

puncturation of the mesonotum. The name tartara is used on

the plate through a fault in the editing, and has unfortunately

been recorded in Dalla Torre’s Catalogue as a distinct species.

720 MR. R. E. TURNER ON

*Eis (Mesa) prorrconiis Mor.

Plesia picticollis Mor. Hor. Soc. Ent. Ross. xxiv. p. 624 (1890), 2.

This appears to be a very distinct species, strongly marked

with yellow on the head, thorax, and abdomen. The colour would

suggest an Anthobosca allied to A. arabica Turn., but the structure

of the apical abdominal segment and the indistinct striation of

the metapleure render it unlikely that it belongs to that genus.

Exis (Mesa) TRicotorn Sm. (Pl. LX XXT. fig. 16; Pl. LX XXII.

fig. 11.)

Myzxine tricolor Sm. Journ. Linn. Soce., Zool. ii. p. 91 (1858), ©.

This fine species is easily distinguished by the large size

(19 mm.), the bright red head, and steel-blue abdomen. The

wings are fusco-hyaline, flushed with purple, almost hyaline at

the base. The head is large, quadrate, sparsely punctured,

smooth on the vertex; pronotum coarsely punctured, median

segment subconcave on the posterior slope; sixth dorsal segment

strongly punctured. Second abscissa of the radius nearly as

long as the first and third combined.

Hab. Borneo (typical) ; Dibrughar, Assam; W. India (7. &.

Bell).

The only specimen I have seen from Assam has the head

distinctly longer than broad, longer behind the eyes than in the

typical form, and the scape and three basal joints of the flagellum

are red; the median segment is not at all concave on the posterior

slope. The differences will probably prove to be subspecific.

Mr. Bell informed me that he bred this species from the larva of

a longicorn beetle.

Eis (Musa) ? pimipraticornis Bingh. (Pl. LXXXI. fig. 15.)

Myzine dimidiaticornis Bingh, Journ, Linn. Soc., Zool. xxv.

p. 423 (1896), ¢ ; Turn, Ann. & Mag. Nat. Hist. (8) i. p. 501

(1908), 3.

3. Antenne stout, a little longer than the thorax and median

segment combined. Front rugose, vertex sparsely punctured.

Pronotum transversely rugulose, much longer than the meso-

hotum, the sides almost parallel. Head slightly narrowed and

produced from behind the eyes. Thorax and median segment

rugose, the median segment distinctly margined posteriorly and

vertically truncate. First abdominal segment vertically truncate

anteriorly, with a distinct transverse carina above the base of

the truncation, attached to the abdomen By a very short petiole,

nearly as broad as the second segment, the sides parallel.

Apical dorsal segment not incised at theapex. Without a pygidial

area. Second abscissa of the radius as long as the first and third

combined.

Black; scape and four basal joints of the flagellum, clypeus,

and apex of the interantennal prominence dull ferruginous ;

abdomen glossed with steely blue. Wings hyaline; fusco-

STUDIES IN THE FOSSORIAL WASPS, 721

violaceous from the basal nervure of the fore wing to the apex

and at the apex of the hind wing.

Length 13 mm.

Hab. Kumaon, N.W. India.

Although this resembles J/yzine rather than Zlis in the form

of the first abdominal segment and the antenne, I consider that

it will probably be found to be the male of Z#. tricolor Sm. The

proportions of the cubital cells, the colour of the abdomen and

antenne, and the colour of the wings are all very similar in the

two forms and unlike any other species, either of Myzine or His.

I consider it probable that the group of Zlis containing dubia,

fedtschenkoi, and apimacula will also be found to have males

showing the facies of Myzine rather than of Hlis. These cases

render it very difficult to reach absolute certainty in distinguishing

the males of Elis from Myzine, though all males with a long

petiole may be assigned to Lis.

I think Methoca rugosa Cam. (Mem, Manchester Soc. 1896)

from Ceylon will prove to be a local form of this species, and also

an individual aberration as far as the loss of one of the transverse

cubital nervures is concerned. It is certainly not a Methoca.

Enis (MzsA) nurset Turn.

Plesia nursei Turn. Ann. & Mag. Nat. Hist. (8) ui. p. 480

(1909), ¢.

The first abdominal segment is no longer than the hind femur

and trochanter combined, the narrowed petiole being rather less

than half as long as the strongly dilated apical portion of the

segment. In general appearance there is a strong resemblance

to petiolata, but there is no yellow band on the pronotum and

sixth abdominal segment. It is also a larger species, and the

shorter petiole is a good distinguishing character, in which the

resemblance is nearer to mandalensis than to petiolata.

Hab. Simla.

Exts (Musa) Extensa Turn.

Plesia (Mesa) extensa Turn. Ann. & Mag. Nat. Hist. (8) i.

p. 511 (1908), 3.

The first abdominal segment is much longer than the hind

femur and trochanter combined ; the antenne are longer than the

head, thorax, and median segment combined. The wings are

strongly suffused with yellowish brown, and the yellow bands on

the abdomen are reduced to short transverse lines on each side of

dorsal segments 2—5.

Length 12 mm.

Hab. Upper Burma.

Eis (Musa) tzra Bingh.

Myzine leta Bingh. Faun. Brit. India, Hymen. i. p. 70

(1897), 3.

3. Black; the legs fusco-ferruginous; wings hyaline. First

(22 : MR. R. E. TURNER ON

abdominal segment a little longer than the hind femur and

trochanter combined, the narrow petiole about half as long as

the dilated apical portion ; seventh dorsal segment distinctly

incised at the apex, the incision nearly as deep as its apical

width ; second segment about half as long again as the third.

Second abscissa of the radius distinctly longer than the third.

Pygidial area clearly defined and strongly punctured. Hind

tibiz more distinctly serrate than in other species of the genus.

Length 9-11 mm.

fab. Moulmein, Tenasserim.

Enis (Mesa) BuRMANICA Bing.

Myzine burmanica Bing. Faun. Brit. India, Hymen. i. p. 70

(1897), 3s.

3. This differs from deta in the black colour of the legs; the

pygidial area is flatter, less distinctly punctured, and not distinctly

margined, and the third abscissa of the radius is distinctly longer

than the second. Whether these differences are of specific value

or not it is not easy to say from a single specimen of each, but I

am inclined to regard them as merely variations of one species.

fab. Amherst, Tenasserim.

ELIS SELLOWI, sp. 0.

@. Nigra ; postscutello linea transversa, segmento dorsali primo

macula magna utrinque, segmentisque dorsalibus 2-5 fascia basali

angusta utrinque flavis ; femoribus, tibiis tarsisque rufo-testaceis ;

mandibulis fusco-ferrugineis ; alis fusco-hyalinis, costa obscura,

venis fuscis.

Long. 11-13 mm.

2. Clypeus strongly punctured, broadly rounded at the apex,

with a longitudinal carina from the base almost reaching the

apex. Front strongly and closely punctured, vertex and ocellar

region sparsely and less deeply punctured, cheeks smooth, an

arched impressed mark above the posterior ocelli; scape strongly

punctured. Thorax deeply and closely punctured, especially on

the pronotum and mesopleure, the dise of the mesonotum and

the scutellum rather sparsely punctured; propleure obliquely

striated. Median segment closely punctured in the middle,

opaque and almost smooth near the anterior angles; the posterior

slope almost vertical, concave in the middle and shagreened, with

short transverse striz on the sides, the sides of the segment

strongly striated. Abdomen shining, with a few scattered

punctures; the apical segment closely longitudinally striated,

with a few large punctures between the strie, very broadly

rounded at the apex. ‘The three abscisse of the radius almost

equal in length; first recurrent nervure received just beyond the

middle of the second cubital cell, second just beyond one-quarter

from the base of the third cubital cell. The ventral abdominal

STUDIES IN THE FOSSORTAL WASPS. 723

segments are very sparsely punctured, with a row of piliferous

punctures near the apex ; the pubescence is whitish.

Hab. San Juan del Rey, Brazil (Sellow). Two specimens.

Type in the Berlin Museum.

ELIS MAJOR, sp. n.

9. Nigra; fascia angusta transversa supra antennas, margine

interiore oculorum anguste, linea pone oculos, postscutello, segmentis

dorsalibus primo secundoque macula magna obliqua utrinque,

segmentisque 3-5 fascia lata sub-basali flavis ; mandibulis, tibris

tarsisque fuscis ; alis flavo-hyalinis, venis testaceis.

Long. 24 mm.

9. Clypeus coarsely punctured, broadly rounded at the apex,

raised and longitudinally carinate in the middle. Front coarsely

punctured, the prominence between the antenne almost trans-

verse at the apex, vertex and ocellar region a little more sparsely

punctured, cheeks very sparsely and more finely punctured ;

posterior ocelli nearly twice as far from the eyes as from each

other. Thorax closely and coarsely punctured, especially on the

pronotum and mesopleure ; propleure finely obliquely striated.

Median segment smooth, opaque, rugosely punctured in the

middle, with a few irregular transverse striz at the apex and

also on the sides of the posterior truncation ; the sides of the

segment striated, with a small yellow band near the apical angle.

Abdomen shining, with a few scattered punctures, the apical

segment closely longitudinally striated and broadly rounded at

the apex. Ventral segments smooth, more or less punctured at

the apex. First and third abscisse of the radius about

equal in length, the second nearly half as long again. First

recurrent nervure received beyond the middle, sometimes at

two-thirds from the base of the second cubital cell, second

received between the middle and one-quarter from the base of

the third cubital cell. Calcaria whitish.

Hab. Central Brazil (Sellow).

Described from three specimens in the Berlin Museum.

The probable male of this species has the usual yellow markings :

the mandibles, clypeus, a small spot on each side above the base

of the antenne, the margins of the pronotum, interrupted in the

middle on the anterior margin, the tegule, a spot on the meso-

notum, one on the centre of the scutellum, a large spot on the

mesopleure below the anterior wings and another before the

intermediate cox, the postscutellum, a lateral longitudinal band

on each side of the posterior slope of the median segment, apical

bands on dorsal segments 1-6, straight and narrow on segments

46, broader and shallowly emarginate on segments 2-3 and very

deeply emarginate on the first segment, the tibie, tarsi, and the

greater part of the femora yellow. No yellow spots on the apical

segment. Wings hyaline, nervures black. Emargination of the-

apical segment broader at the apex than deep, the dorsal surface

124 MR. R. E. TURNER ON

of the segment flattened and distinctly margined lateraily,

shallowly longitudinally depressed in the middle and sparsely

punctured ; the basal segment distinctly less than twice as long

as the breadth at the apex. Clypeus broadly rounded or sub-

truncate at the apex, not emarginate. Second recurrent nervure

received at the base of the third cubital cell, almost interstitial

with the second transverse cubital nervure; third abscissa of the

radius a little longer than the second. Pronotum siightlv

emarginate anteriorly.

Length 19 mm.

The details of neuration in the Hlidine are not reliable for

specific differences, often showing slight differences on the

opposite sides of the same insect. ‘The really important

distinctions in the males of “lis are to be found in the structure

of the apical and basal segments of the abdomen.

Eis compusta Sm. (Pl. LX XXII. fig. 12.)

Myzine combusta Sm. Deser. new spec. Hymen. p. 179 (1879),

@; Bingh. Faun. Brit. Ind., Hymen. i. p. 67 (1897), @.

The type is from Shuckard’s collection. I do not know why

Smith gave the locality as India or Africa, nor why Colonel

Bingham included it in the Fauna of India without any note of

doubt. The structure is that of the true American section of

Elis, and I look on it as a mere colour-variety, or possibly a local

race confined to one island, of H. ephippiwm Fabr., a West Indian

species which I have recorded from St. Thomas, St. John, Antigua,

and Porto Rico.

Subfamily ANTHOBOSCIN &.

Genus AnNTHOBOSCA Guér.

Anthobosca Guér. Voy. ‘ Coquille,’ Zool. ii. p. 214 (1839).

Cosila Guér. Voy. ‘ Coquille,’ Zool. ii. p. 249 (1839).

Dimorphoptera Sm. Trans. Ent. Soe. London, p. 238 (1868).

Myzine Burm. Stett. ent. Zeit. xxxvii. p. 168 (1876) (nec

Latreille).

Odontothynnus Cam. Rec. Albany Mus. 1. p. 161 (1904).

_Austrotiphia Cockerell, Bull. Mus. Comp. Anat. Harvard, p. 49

(1906).

ee Cockerell, Bull. Mus. Comp. Anat, Harvard, p. 49

1906).

Hic Turns);Proe: Winn: sSoc. Nis, Wie) soe page.

(1907).

There has been so much confusion in connection with this

interesting genus that I think it may be useful to give a short

account of the species, which may serve as a basis for a future

monograph. Owing to the great differences between the sexes,

they were for a long time placed in different families, most of the

STUDIES IN THE FOSSORIAL WASPS. 725

males being placed in Anthobosca in the Thynnide, and the

females in Cosila among the Scoliide. Guérin used both

names in the same work, but although he placed Cosila

chilensis correctly, with the Scoliide, he failed to see the

relationship to Anthobosca australasie which he had described

in an earlier portion of the same work and classed with the

Thynnide.. Smith in 1868 placed a single male in his

genus Dimorphoptera, which is a synonym of Cosila, but both

before and after placed other males in Anthobosca among the

Thynnide. He also described both sexes of A. albomaculata Sm.,

which were taken coupled by Bates, as Myzine, but this work was

published after his death and without his revision. Burmeister

also in 1876 correctly associated the sexes, but placed his species

in Myzine. Although three or four males had been correctly

associated with the females, they do not appear to have been

connected in any way with the males described in the genus

Anthobosca till my revision of the Australian species of the

genus appeared in 1907. The best work on the genus was done

by Saussure in 1892; but he treated the males as unknown,

except in the case of chilensis, and did not connect Cosila with

Anthobosca. In Dalla Torre’s great Catalogue, published in

1897, there is much confusion in regard to this genus, species

appearing under the genera Zhyniwus, Myzine, and Cosila. There

is much confusion over the genus in the recent papers of Cameron

on South African Hymenoptera. Ashmead treats the group as a

family, Cosilide, not as a subfamily of the Scoliide, which I

consider the more natural course. But he places Anthobosca in

the Thynnide and Dimorphoptera in his family Myzinide. He

includes in his family Cosilide several genera of doubtful

affinities, of which, in my opinion, Vursea should be treated as

an aberrant genus of the Sphecoidea, whilst Maurillus belongs to

the Pompilidee. The position of Sierolomorpha and Dicrogeniwm

seems to me very doubtful, but I have not seen specimens. J am

compelled to look on Anthobosca as the only genus which can

be placed in the subfamily Anthoboscine with any certainty,

Ashmead states that the intermediate coxe in his Cosilide are

contiguous or nearly so; but this is quite incorrect as to the

females, and even in the males the separation is quite distinct,

In my key to the species I have included several species which I

have not seen, one or two of which may possibly not belong to

the genus.

The females are distinguished from other Scoliidee by the

absence of a deep groove between the two basal ventral segments

of the abdomen. The hind coxe are separated as in Tiphia,

not contiguous as in Hlis; but the intermediate cox are less

widely separated than in either of those groups, though very little

less so than in Hlis. The males are distinguished from all other

Scoliidee by the unarmed rounded hypopygium—in this character

approaching most nearly to the 'Thynnide of the genus Hirone.

736 MR. R. E. TURNER ON

Tt is quite possible that the genus will have to be subdivided

owing to differences of structure, especially in the tarsal ungues,

which in most of the Australian species have a blunt lobe at the

base, but in the African, South American, and a few Australian

species are bifid. The neuration is so variable, even in the same

species, that I do not consider that genera should be founded on

the shape of the radial cell alone, on which character the sub-

genera Colobosila Sichel and Cadlosila Saussure have been based.

In two Australian Species the cell is acute at the apex, in others

blunt, and African species may similarly be divided into two

groups. But until more species are known in both sexes any

subdivision would be rash and unnecessary.

The geographical distribution of the genus is interesting, being

almost entirely Southern, including Australia (where the species

are most numerous), South America as far north as the Amazon,

Madagascar and South Africa spreading up the East African

coast to Suakin and crossing to Aden. Iam aware that many

regard similar cases of distribution as a proof of southern origin,

and explain them by former northern extensions of the Southern

continent connecting at different times with southern extensions

of the land in the Southern Hemisphere. But taking into

consideration the enormous depth of the Southern oceans I

cannot look on this explanation as satisfactory, and think that it

is more reasonable to look on this and other similar cases of

distribution as instances of the survival in the south of genera

which in former times had a much more extensive range. It

has been pointed out by Darwin that the struggle for existence

is more severe on large land-areas than on smaller ones, owing to

the more complex conditions of life from the larger number of

existing species which are able to come into competition. Now

the land-areas in the south are very much smaller than in the

north, so that it is reasonable to suppose that many genera may

have been able to survive in the south with little or no modi-

fication, which have been exterminated by the more severe

struggle for existence in the north. In this case we should

expect to find fossil remains of such genera, or at all events of

nearly related forms, in the north; and in a great number

of cases such fossils have been found. In the present genus I

look on Cockerell’s Geotiphia found fossil in Colorado as absolutely

congeneric with existing South-American species ; and the plant

genus dAraucaria and many others, which are now southern, had

in early geological times a wide range in the Northern Hemi-

sphere. It may, of course, be argued ‘that such genera originally

had their home in the south and at one time extended their

range northward, but in that case it may be conceded that

they may have reached the different portions of the Southern

Hemisphere by way of the north and not from a southern

continent. I must own to an objection to calling up continents

from the extreme ocean depths “‘ nisi dignus vindice nodus.”

10.

11.

12.

13.

16.

STUDIES IN THE FOSSORIAL WASPS. 72

Key to the Species of Anthobosca.

Females.

. Species from the Old World......

South-A merican species

. Australian species .....:..

Atrican and Arabian species _

. Tarsal ungues with a lobe at the base ‘either ‘rounded

or pointed... Bi

Tarsal ungues without a basal lobe, deeply ‘bifid .....

. Radial cell pointed at the apex

Radial cell blunt at the apex .......... ins a eee tes

. Pubescence black, calcaria black, abdomen wholly

LAC Kec area ini ccacntoat teas eaten ae ueceas anette pees

Pubescence whitish, calcaria white, abdomen usually

with a yellow spot on each side of the third dorsal

segment .....

F Wines short, the length ‘of the costa of the fore w ving

not exceeding two and a half times the breadth of

the mesonotum. Pronotum, median segment, and

three basal abdominal segments rufo-testaceous ...

Wings longer, the length of the costa of the fore

wing three times as great as the breadth of the

mesonotum. Thorax and abdomen black .........

. Antenn# orange; stigma about three times as long

on the costa as broad Se aR iat ede Ae ae

Antenne black or fuscous; stigma scarcely more

than twice as long on the costa as broad ............

. Legs bright ferruginous

Legs black, sometimes fusco- ferruginous beneath —

. Second abscissa of the radius no longer than the

third ; first recurrent nervure received just before

the middle of the second cubital cell ..................

Second abscissa of the radius longer than the third;

first recurrent nervure received at one-quarter from

the base of the second cubital cell .

Wings fuscous; four basal abdominal ‘segments ‘with

cilize of white scale-like hairs .............2s.e0000- ++.

Wings hyaline or subhyaline; abdominal segments

without cili# of scale-like hairs ....................00.

Pronotum and scutellum strongly punctured .........

Pronotum and scutellum Spee and very nue,

punctured .....

Third cubital cell nearly a as ‘long on the radius as on

the cubitus or longer; fir st recurrent nervure

received slightly beyond the middle of the second

CUPILACEM Me eet oteneee Reber esc ma conic senrerseee

Third cubital cell much shorter on the radius than

on the cubitus; first recurrent nervure received

well before the middle of the second cubital cell ...

Front punctured; legs wholly black; length 11 mm.

Front smooth and shining; legs fusco-ferruginous

beneath ; length 7 mm..

. Abdomen black ; second and third cubital ‘cells each

receiving a recurrent nervure ............-.0060cee eee eee

Abdominal segments 2-4 very broadly rufo-testaceous

at the base; second cubital cell receiving both

LECHIECHUINE ULES a teem se ote vis arencivsecvalcvelaceiee

. Sixth dorsal segment strongly striated at the base;

radial cell detached from costa for half its length .

Sixth dorsal segment punctured; radial cell detached

from costa at extreme apex only ..

Radial cell subacute at apex; abdomen not marked

with yellowish white.. 5

Radial cell broadly rounded or 1 narrowly truncate at

apex; abdomen marked with yellowish white ......

—

Sue

A, australis Sichel.

A. signata Sm.

A, fastuosa Sm.

A, flavicornis Sauss.

10,

A, cognata Sm,

A, strandi Turn,

at ie

12.

A. albopilosa Turn,

A. argenteocincta Grib,

A. anthracina Sm,

13.

A. unicolor Sm,

A, levifrons Sm,

15,

A, clypeata Sm.

A, nubilipennis Turn.

A. occipitalis Turn,

20,

728 “MR. R. E. TURNER ON

17. Apical segment at least of abdomen red; wings

HUNEO APOC OWS! coc son sad tus aunose Kod uewebsgdncensnoenb coerce 18.

Abdomen wholly black; wings flavo- or fusco-hyaline. 19.

18. Apical abdominal segment only red; thorax not

coarsely punctured — .......--.eeeeeseeeseeeeeeeeeseeessenee *A, melanaria Cam.

Two apical abdominal segments red; thorax very

coarsely sculptured ............:cesscsesesseeeesesseaseees *A, donaldsoni Fox.

19. Wings flavo-hyaline; third transverse cubital nervure

received close to the apex of the radial cell ......... A. insularis Sm.

Wings fusco-hyaline; fourth abscissa of the radius

RIV NOTE? YS) TN IWUAE Foooaarpagcosoh908 con bo peo Aoh opp danscaone

20. Radial ceil narrowly truncate at apex. Length 5mm.

Radial cell broadly rounded at apex. Length 9mm.

. natalica Turn.

. minima Turn.

bs be

COUNTING Satansesoouuageicnvonedonmanteese Coasdopon vad ddooded 21.

21. Head and thorax black with pale yellow markings ... 22.

Head and thorax without yellow markings, thorax

black or more or less red ............s0.ceeeerereeeeeeeee A. erythronota Cam.

Oe, oN oslovaner 1a AA DEATHS A dooheacas soouaaden ceoenosdDdoscachoas *A. sauakinensis Magy.

Aibdomentdhlack a eererce eee wae barce enact ne cetera rer A. arabica Turn.

23. Abdomen black without markings ............-...0.s0 ee 24.

Abdomen marked with yellow or white.................. 25;

24, Antenne orange; head and thorax closely puuctured. A. chilensis Guér.

Antenne black; head and thorax very sparsely

FOVUNMVGUUTREGL cue odn. dou codeine hoawasincd ebhave dusesa change ocaad Sec A. carbonaria Burm.

25. Wings flavo-hyaline; abdomen with four or five

THA(ETOH HOLE yp GN? WEHOGIS oo Sco sesios Joktonn coh eansen osGno 26.

Wings fusco-hyaline; abdomen with one or two

yellowashelaterallispOts) ee eeessceseses sere area 27.

26. Legs ferruginOUS ....-..c--ecececeesnecenseecerceteesernseen> A. ENLENNALG SM.

Legs black ......... * A. erythropyga Burm.

27. Vertex smooth and shining; third dorsal segment

Only vg belaterallispOts |e eee teeeaacteit eto oaene ered

Vertex sparsely punctured; second and third dorsal

segments with whitish lateral spots .................. 4d. albomaculata Sm.

A. bipunctata Perty.

1, AnrHoBoscA AUSTRALIS Sichel. (PI. LX XXII. figs. 1, 2;

Pl, LX XXIII. figs. 1, 3, 6, 14.)

Cosila australis Sichel, Sauss. et Sichel, Cat. spec. gen. Scolia,

p, 261 (1864), 2.

Dimorphoptera scoliiformis Sm. Trans. Ent. Soc. London, p. 238

(1868), ¢@.

Cosila (Callosila) australis Sauss. in Grandidier, Hist. Mada-

gascar, xx, p. 232 (1892), 9.

Dimorphoptera nigripennis Sm. Trans. Ent. Soc. London, p. 239

(1868), 3.

@. The radial cell is pointed and detached from the costa at

the apex, receiving the second transverse cubital nervure close to

the middle, the second abscissa of the radius almost as long as the

third. First recurrent nervure received close to the middle of

the second cubital cell, second at one-third from the base of the

third cubital cell. Third cubital cell fully half as long again on

the cubital as on the radial nervure. Basal lobe of tarsal ungues

bluntly rounded. Basal joint of hind tarsi with a scopa beneath,

in which is a row of longer black spines. Sparsely punctured,

shining; pronotum and scutellum more closely punctured ;

pygidium densely clothed with long black pubescence.

Black, with black pubescence. Wings fusco-violaceous.

Length 23 mm.

STUDIES IN THE FOSSORIAL WASPS. 729

3. Second abscissa of the radius about half as long as the

third. Basal lobe of tarsal ungues bluntly rounded. | Apical

joint of the flagellum about half as long again as the penultimate

and slightly curved ; the eighth, ninth, tenth, and eleventh joints

subtuberculate im the middle beneath. Posterior tibie very

feebly spined on the outer margin. Pronotum short, scarcely

narrowed anteriorly. First abdominal segment very little longer

than the second, broader at the apex than long. Head and

thorax closely and not very minutely punctured.

Black, with cinereous pubescence, calcaria black. Wings

fusco-violaceous.

Length 17 mm,

Hab. HK. Coast of Australia between Sydney and Brisbane.

The association of the sexes is not certain. The structure of

the male antenne is very remarkable.

2. ANTHOBOSCA SIGNATA Sm.

Myzine signata Sm. Cat. Hym. B. M. iii. p. 75 (1855), @.

Cosila biguttata Sichel, Sauss. et Sichel Cat. spec. gen. Scolia,

p. 262 (1864), ¢.

Dimorphoptera signata Sm. Trans. Ent. Soc. London, p. 238

(1868), @.

Cosila (Callosila) signata Sauss. in Grandidier, Hist. Madagascar,

‘exp. 232 (1892), 2).

Var. Dimorphoptera morosa Sm, Trans. Ent. Soc, London, p. 239

(1868), ©.

®. Radial cell pointed and detached from the costa at the

apex, second recurrent nervure received at about one-quarter

from the base of the second cubital cell. Other details of

neuration as in australis Sich. Basal joint of hind tarsi with a

comb of thickly set spines beneath. Basal lobe of tarsal ungues

bluntly rounded. Shining and sparsely punctured, the front and

pronotum more closely punctured; sixth dorsal segment of

abdomen rugosely punctured.

Black, with whitish pubescence, the calcaria white. Wings

fusco-violaceous. Third dorsal segment with an orange spot on

each side.

Length 22 mm.

Hab. EK. Australia, from Sydney to Cairns.

I look on morosa as a small variety in which the orange spots

on the abdomen are wanting. The second abscissa of the radius

is distinctly shorter than the third in the type of morosa, owing to

the sharp curve toward the base in the second transverse cubital

nervure, but I do not think that this distinction is of specific

importance.

3. AnTHoposca FasTuosA Sm. (Pl. LXXXT. fig. 3.)

Dimorphoptera fastwosa Sm. Trans. Ent. Soc. London, p. 240

(1868), @.

Myzine fastuosa D. T, Cat. Hym. viii. p. 123 (1897).

Proc. Zoou. Soc.—1912, No. XLVIII. 48

730 MR. R. E. TURNER ON

@. Wings short, the length of the costa of the fore wing not

exceeding two and a half times the breadth of the mesonotum.

Radial cell broad, the apex detached from the costa and narrowly

rounded. Third abscissa of the radius nearly half as long again

as the second, and more than twice as long as the first; second

transverse cubital nervure received at about two-fifths from the

base of the radial cell. Second cubital cell not very strongly

produced towards the base on the cubitus, receiving the first

recurrent nervure near the middle, second recurrent nervure

received just beyond one-quarter from the base of the third

cubital cell. Basal joint of hind tarsi finely pectinate beneath ;

basal joint of fore tarsi with six spines above and about eight

shorter ones beneath; basal lobe of tarsal ungues bluntly rounded.

Sparsely punctured, more closely on the front, pronotum, and

abdomen ; the apical abdominal segments more strongly punctured

than the three basal segments.

Rufo-testaceous; head, mesothorax, and three apical abdominal

segments black; mandibles at the base and scape fusco-ferruginous ;

pubescence rufo-testaceous. Wings hyaline, tinged with yellow ;

nervures testaceous, the stigma black.

Length 17 mm.

Hab. Champion Bay, W. Australia.

The process beneath the hind femora near the apex is much less

developed in this species than in most Australian species, though

the femora are by no means convex beneath as in most of the

South-American species and in A. nubilipennis.

4. ANTHOBOSCA FLAVICORNIS Sauss.

Cosila (Callosila) flavicornis Sauss. in Grandidier, Hist. Mada-

gascar, xx. p. 233 (1892), ©.

@. Radial cell with a slight angle at the apex ; second abscissa

of the radius shorter than the first, the third longer than the

first and second combined, third cubital cell of about equal

length on the radial and cubital nervures. Second cubital cell

much produced towards the base on the cubitus; first recurrent

nervure received about the middle of the second cubital cell,

second just before the middle of the third cubital cell. Basal

joint of hind tarsi with a scopa beneath, in which are a few

spines longer than the hairs; basal joint of fore tarsi with six

stout spines above and a row of very fine spines beneath; basal

lobe of tarsal ungues rounded. ‘The lobe beneath the apical

portion of the hind femora has the lower margin nearly straight,

only very slightly rounded, and is strongly developed. Shining

and rather sparsely punctured, the anterior half of the pronotum

and the median segment closely and finely punctured and sub-

opaque.

Black, with white pubescence; the flagellum orange; calcaria

white, the spines of the tibie and tarsi reddish brown. Wings

fusco-hyaline, nervures fuscous.

STUDIES IN THE FOSSORIAL WASPS. 731

Length 11-14 mm.

Hab. Victoria.

There is also a specimen in the National Collection from

Tasmania and another from Cairns, Queensland. In the former

the radial cell is rounded at the apex and the tibie and tarsi are

reddish ; in the latter the radial cell is almost truncate at the

apex, though narrowly, and the second abscissa of the radius is

as long as the first. I do not think that small differences of

neuration in this genus will prove to be of specific value.

5. ANTHOBOSCA COGNATA Sm.

Dimorphoptera cognata Sm. Descr. new spec. Hymen. p. 188

(1879), @.

@. Radial cell rather broadly rounded at the apex; the four

abscissee of the radius of about equal length; the third cubital

cell much longer on the cubitus than on the radius. Recurrent

nervures received close to the middle of the second and third

cubital cells; the second cubital cell strongly produced toward

the base on the cubitus. Basal joint of hind tarsi with a scopa

beneath in which are five or six longer spines; basal joint of fore

tarsi with six long and rather slender spines above and a closely

set row of very small spines beneath ; basal lobe of tarsal ungues

less bluntly rounded than in flavicornis. The lobe beneath the

apical half of the hind femora is very broadly rounded. Closely

and finely punctured, more finely and closely on the abdomen

and median segment than on the head and thorax, some larger

punctures on the disc of the mesonotum.

Black, with sparse whitish pubescence; mandibles at the hase

and legs, except the cox, ferruginous.

Length 11 mm.

Hab. Swan River, Western Australia.

6. ANTHOBOSCA STRANDI Turn.

Anthobosca strandi Turn. Proc, Zool. Soc. London, p. 306

(1910), 2.

@. Radial cell broadly rounded at the apex; second abscissa

of the radius twice as long as the first and distinctly longer than

the third; first recurrent nervure received at one-quarter from

the base of the second cubital cell, second at one-fifth from the

base of the third cubital cell. Apical lobe beneath the hind

femora broadly rounded and strongly developed. Pronotum,

median segment, and abdomen very finely and closely punctured,

front coarsely and closely, mesonotum and scutellum less coarsely

and rather sparsely punctured.

Black ; the pubescence pale fulvous on the head and pronotum,

grey on the sides of the abdomen, fusco-ferruginous on the apical

dorsal segment ; apex of the scape and flagellum beneath fusco-

ferruginous, a dull ferruginous spot on each side close to the

summit of the eyes; mandibles at the base, tegule, tibie, tarsi,

48*

(32 MR. R. E. TURNER ON

femora at the apex, and the whole of the hind femora ferruginous.

Wings subhyaline, nervures fuscous, stigma ferruginous.

Length 13 mm.

Hab. B. Australia; probably from Victoria.

I have not a specimen of this species before me at the time of

writing.

7. ANTHOBOSCA ALBOPILOSA Turn.

Anthobosca albopilosa Turn. Proc. Linn. Soc. N.S.W. xxxii.

p- 520 (1907), 2.

Q. Radial cell slightly angular at the apex; third abscissa of

the radius the longest, but shorter than the first and second

combined, the first*scarcely as long as the second. First re-

current nervure received near the middle of the second cubital

cell, second just beyond one-quarter from the base of the third

cubital cell. Second cubital cell less strongly produced towards

the base than in flavicornis. Basal joint of hind tarsi with a

scopa beneath, in which are a few longer spines; basal joint of

fore tarsi with six or seven well-developed spines below and the

usual six above; basal lobe of tarsal ungues rounded; lobe

beneath the apical half of the hind femora broadly rounded and

less strongly developed than in flavicornis. Deeply and rather

closely punctured, the abdomen more shallowly punctured.

Black; the pubescence on the head and sides of the thorax

white, on the abdomen and dorsal surface of the thorax black ;

calearia black; four basal dorsal segments of abdomen and

ventral segments 2-4 with apical ciliz of scale-like white hairs.

Wings fuscous, with faint violet reflections.

Length 14 mm.

Hab. Queensland.

8. ANTHOBOSCA ARGENTEOCINCTA Grib.

Cosila argenteocincta Gribodo, Ann. Mus, Civ. Stor. Nat. Genova,

Xvili. p. 261 (1883), 2.

Q. Radial cell with a slight angle at the apex; second abscissa

of the radius longer than the first, the third equal to the first

and second combined, the third cubital cell a little longer on the

cubitus than on the radius, First recurrent nervure received

just beyond the middle of the second cubital cell, second at one-

third from the base of the third cubital cell; the second cubital

cell moderately produced towards the base on the cubitus. Basal

joint of hind tarsi with a close row of small spines beneath ; basal

joint of fore tarsi with seven spines above, and a close-set row of

slender spines beneath; basal lobe of tarsal ungues bluntly

rounded; lobe beneath the basal half of the posterior femora

broadly rounded. Shining, finely and very sparsely punctured ;

the median segment subopaque and very finely and_ closely

punctured on the sides; abdomen shallowly but much more

coarsely punctured.

STUDIES IN THE FOSSORIAL WASPS. ee

Black, with sparse grey pubescence ; flagellum beneath fuscous ;

tarsal ungues ferruginous ; calcaria white. Wings light fuscous,

shaded with bronze ; nervures black.

Length 13 mm.

Hab. Adelaide, 8. Australia.

I think I have identified this species correctly. Gribodo

mentions a tubercle at the base of the first ventral segment,

which does not seem to be any more developed in the specimen

described above than in other species.

9. ANTHOBOSCA ANTHRACINA Sm. (PI. LX XXII. fig. 3.)

Myzine anthracina Sm, Cat. Hym. B. M. iii. p. 71 (1855), 9.

Myzine sabulosa Sm. Cat. Hym. B. M. iii. p. 76 (1855), 9.

Dimorphoptera anthracina Sm. Trans. Ent. Soc. London, p. 238

1868).

pisos subulosa Sm. Trans. Ent. Soc. London, p. 238

1868).

Ausrotitphia kirbyt Cockerell, Bull. Mus. Comp. Anat. Harvard,

p. 49 (1906), @.

Q. Radial cell very bluntly rounded at the apex; second

abscissa of the radius shorter than the first, the third abscissa

half as long again as the first and second combined. First

recurrent nervure received just beyond the middle of the second

cubital cell, which is more sharply produced towards the base

than in wiicolor Sm. or levifrons Sm.; second recurrent nervure

received at two-fifths from the base of the third cubital cell,

which is longer on the radius than on the cubitus. Basal joint

of hind tarsi finely pectinate beneath; basal joint of fore tarsi

with six spines above and eight finer beneath; basal lobe of tarsal

ungues bluntly rounded. Finely and rather sparsely punctured,

pronotum more coarsely punctured, median segment very finely

and closely punctured with a smooth line in the middle.

Black with whitish pubescence; spines of the tibiz and tarsi

white, those of the anterior pair slightly reddish. Wings

hyaline, shaded with fuscous towards the apex, nervures pale

testaceous.

Length 14 mm.

Hab. S.K. Australia; Adelaide to Sydney.

The description is taken from the type of awuthracina. The

type of sabulosa has the first and second abscisse of the radius

about equal in length, combined equal to the third; the third

cubital cell no longer on the radius than on the cubitus; the

nervures are fuscous. But excepting in these points I cannot

detect any difference, and do not consider that small differences

of neuration can be held to be of specific importance, considering

how much variation occurs in other groups of Scoliide in such

detail. The type of Austrotiphia kirbyi is similar to sabulosa.

734 MR. R. BE. TURNER ON

10. ANTHOBOSCA UNICOLOR Sm.

Myzine unicolor Sm. Cat. Hym. B. M. i. p. 75 (1855), 2.

Dimorphoptera unicolor Sm. Trans. Ent. Soc. London, p. 238

(1868), 9.

Cosila (Callosila) minuta Sauss. in Grandidier, Hist. Madagascar,

xx. p. 233 (1892), 9.

Q. Radial cell rounded at the apex; second abscissa of the

radius as long as the first, the two combined a little longer than

the third; first recurrent nervure received a little before the

middle of the second cubital cell, second before one-third from

the base of the third cubital cell, which is much longer on the

cubitus than on the radius; second cubital cell not very strongly

produced towards the base. Basal joint of hind tarsi with a few

small spines beneath ; basal lobe of tarsal ungues rounded; lobe

on the apical third of the hind femora beneath not very strongly

prominent. Shining and sparsely punctured, the front closely

and strongly punctured; median segment subopaque, very finely

and closely punctured.

Black, with white pubescence; mandibles and scape beneath

fuscous; calcaria white. Wings hyaline, faintly tinted with

fuscous.

Length 10 mm.

Hab. Kastern Australia ; Tasmania to Cairns.

11. ANTHOBOSCA LEVIFRONS Sm.

Dimorphoptera levifrons Sm. Deser. new spec. Hymen. p. 188,

(1879), Q.

Myzine levifrons D, T. Cat. Hymen. viii. p. 124 (1897).

Q@. Radial cell broadly rounded at the apex; second abscissa of

the radius shorter than the first, the two combined longer than the

third ; first recurrent nervure received at two-fifths from the base

of the second cubital cell, second just before the middle of the third

cubital cell, which is much longer on the cubitus than on the radius.

Basal joint of hind tarsi with a row of fine hairs beneath, in which

are two or three longer spines; the lobe beneath the apical third

of the hind femora broadly rounded. Basal lobe of tarsal ungues

small and blunt. Shining, head and thorax very sparsely punc-

tured, the front smooth; median segment and abdomen closely

and minutely punctured.

Black, with sparse white pubescence; mandibles, flagellum

beneath, and legs beneath fusco-ferruginous. Wings hyaline,

tinged with fuscous ; nervures fusco-ferruginous.

Length 7 mm.

Hab. Adelaide, S. Australia.

12. ANTHOBOSCA OCCIPITALIS, sp. n.

2. Nigra, vertice macula utrinque supra oculos fusco-sanguinea ;

unguiculis bifidis ; alis subhyalinis, venis fusco-ferrugineis.

Long. 14 mm

TUDIES IN THE FOSSORIAL WASPS. 735

@. Radial cell rounded at the apex; third abscissa of the radius

shorter than the second ; third cubital cell as long on the radius as

on the costa; first recurrent nervure received just beyond one-

third from the base of the second cubital cell, second at one-fifth

from the base of the third cubital cell; second cubital cell not

very strongly produced towards the base. Tarsal ungues bifid,

without a basal lobe; basal joint of hind tarsi with rather long

hairs beneath ; lobe beneath the apical half of the hind femora

broadly rounded. Closely punctured; rather strongly on the

head, finely on the thorax, coarsely on the scutellum, rather

sparsely and shallowly on the abdomen; sixth dorsal segment

rather broadly truncate at the apex.

Black, with grey pubescence, a fusco-sanguineous spot on the

vertex on each side close to the summit of the eyes; sixth dorsal

segment fuscous at the apex, calcaria whitish.

Hab. S. Australia; probably from Adelaide.

This species is remarkable for the very long second abscissa of

the radius, and may be easily distinguished from the majority of

Australian species by the bifid tarsal ungues.

13. ANTHOBOSCA NUBILIPENNIS Turn.

Anthobosca nubilipennis Turn. Proc. Zool. Soc. London, p. 307

(ESTO) 90".

@. Radial cell detached from the costa for fully half its length,

slightly angular at the apex ; second abscissa of the radius longer

than the first and about equal in length to the third; first

recurrent nervure received beyond two-thirds from the base

of the second cubital cell, second before one-quarter from the

base of the third cubital cell, which is nearly twice as long

on the cubitus as on the radius. Tarsal ungues bifid, without a

basal lobe ; hind femora convex beneath, the lobe not apical but

almost medial; basal joint of hind tarsi with three or four short

spines beneath. Deeply but not very closely punctured ; meso-

notum and scutellum very sparsely punctured; sixth dorsal

abdominal segment coarsely striated at the base, rounded at the

apex.

Shee with white pubescence; calcaria black. Wings fusco-

hyaline, nervures fuscous; the stigma small. Median segment

abruptly truncate.

Length 16 mm.

Hab. Perth district, S.W. Australia.

A very distinct species in the characters of the hind legs,

neuration, and median segment.

14, ANTHOBOSCA CLYPEATA Sm. (Pl. LX XXI.fig.2; Pl. LX XXII.

fig. 5; Pl. LX XXIII. fig. 7.)

Dimorphoptera clypeata Sm. Trans. Ent. Soc. London, p. 239

(1868), 2.

2. Radial cell rounded at the apex, detached from the costa

736 MR. R. E. TURNER ON

at the extreme apex only; second abscissa of the radius twice as

long as the first and about equal in length to the third; second

cubital ceil receiving both recurrent nervures, the first at one-

third from the base, the second just before the apex; third

cubital cell as long on the radius as on the cubitus, Tarsal

ungues bifid; lobe beneath the apical half of the hind femora

strongly rounded : basal joint of hind tarsi with a row of short

hairs beneath, in which are a few longer spines. Closely punc-

tured; the thorax very closely and finely punctured, with larger

and sparser punctures intermixed ; abdomen shining and sparsely

puuctured.

Black, with whitish pubescence; sixth dorsal segment densely

clothed with long golden hairs; clypeus except at the apex and

the margins of the eyes broadly interrupted on the summit,

yellow ; basal two-thirds of dorsal segments 2-4 and of ventral

segments 2-3 emarginate in the middle posteriorly, rufo-

testaceous. Wings subhyaline, nervures fusco-ferruginous.

Length 22 mm,

Hab. Swan River, W. Australia.

An obscure scar runs from the base of the stigma to close to

the base of the first transverse cubital nervure. I have little

doubt that the male of this species will prove to be A, crassicornis

Sm., which corresponds with it in the position of the recurrent

nervures, the bifid tarsal ungues, and also to some extent in the

colour of the clypeus and abdomen,

*15, ANTHOBOSCA FASCICULATA Sichel.

Cosila (Colobosila) fascicylata Sichel, Sauss, et Sichel, Cat, spec.

gen. Scolia, p, 263 (1864), 9,

Hab, Australia.

I fail to recognise this species with any certainty, but do not

consider that the subgenus Colobosila can stand, the truncation

of the apex of the radial cell being insufficient as a subgeneric

character unless supported by others,

*16, ANTHOBOSCA INORNATA Sauss.

Cosila inornata Saussure in Grandidier, Hist, Madagascar, xx.

p. 233 (1892), Q.

T do not recognise this species as a synonym of any known to

me, though it seems to be related to A. anthracina, differing in

the position of the first recurrent nervure, Considering how

unreliable the details of neuration are in this genus, it is quite

possible that it is only a variety of anthracina,

*17, ANTHOBOSCA MELANARIA Cam,

Plesia melanaria Cam. Rec. Albany Mus, i. 5, p. 297 (1905), 2,

@. From the description I have no doubt that this is an

Anthobosca, the characters “‘ apex of radial cell sharply pointed,”

and “pygidium piceous red, its base fringed with long, bright

rufous hair,” and “ metapleure smooth,” agreeing much better

STUDIES IN THE FOSSORIAL WASPS. 737

with Anthobosca than Plesia. Although the two genera belong

to different subfamilies of the Scoliide, the females are likely to

be confused by a beginner. Cameron places male Scoliide of the

subfamily Anthoboscine in the Thynnide, and females of the same

group with the Scoliidz of the subfamily Elidine.

*18. ANTHOBOSCA DONALDSONI Fox,

Cosila donaldsoni Fox, Proc. Acad, Philadelphia, p. 549

(1896), 2.

Q. Third transverse cubital nervure received close to the apex

of the radial cell. Clypeus tridentate on the anterior margin.

Head and thorax very coarsely punctured, pronotum and scu-

tellum scabrous; median segment finely striato-punctate, the

sides obliquely striated ; abdomen strongly but sparsely punc-

tured, sixth dorsal segment striato-punctate; tarsal ungues

cleft.

Black, with greyish pubescence ; two apical abdominal segments

red; calcaria whitish. Wings black, strongly violaceous.

Length 18 mm, .

Hab. Somaliland.

I have not seen this species, but the description corresponds

almost exactly with Hlis alicie described in this paper, though

the clypeus in that species is not distinctly tridentate. But I do

not believe that Fox would have confused the genera.

19. AnrHoBoscA INSULARIS Sm. (Pl. LXXXIIT. fig. 6;

Pl. LX XXIII. fig. 8.)

Myzine insularis Sm. Descr. new spec. Hymen. p. 178

(1879),..9.

Cosila insularis Saussure in Grandidier, Hist. Madagascar, xx.

p. 231 (1892), 2.

2. Radial cell subacute at the apex, receiving the third trans-

verse cubital nervure very near the apex, in some specimens

almost at the apex; second abscissa of the radius longer than the

first, the two combined not quite as long as the third; first

recurrent nervure received at two-fifths from the base of the

second cubital cell, second at one-quarter from the base of

the third eubital cell, which is nearly or quite as long on the

radius as on the cubitus. Tarsal ungues bifid; hind femora

very broadly rounded beneath on the apical two-thirds, basal

joint of hind tarsi with three long spines beneath. Shining and

sparsely punctured, closely on the front and closely and very

finely on the median segment.

Black, with black pubescence; calcaria brown. Wings flavo-

hyaline, nervures ferruginous.

Length 23-29 mm.

Hab. Madagascar.

The type has a rufous spot in the middle of the first dorsal

segment, and obseure reddish shading on the head and thorax.

738 MR. R. E. TURNER ON

20. ANTHOBOSCA NATALICA Turn.

Anthobosca natalica Turn. Trans. Ent. Soc. London, p. 895

(1908), 2.

Q@. Radial cell narrowly rounded at the apex, subacute; third

abscissa of the radius twice as long as the second, the fourth

shorter than the first ; first recurrent nervure received just beyond

the middle of the second cubital cell, second before the middle of the

third cubital cell, which is as long on the radius as on the cubitus.

Tarsal ungues bifid; basal joint of hind tarsi with a scopa beneath.

Shining and sparsely punctured, more closely on the front and

pronotum ; median segment very finely and closely punctured ;

abdomen finely but more sparsely punctured.

Black ; pubescence on the apical dorsal segment brown and

long ; tegule and extreme apical margin of the abdominal segments

testaceous brown; calcaria whitish ; fore tarsi fusco-ferruginous.

Wings fusco-hyaline, nervures fusco-ferruginous.

Length 12-17 mm.

Hab. Malvern, Natal.

The neuration given is as in the type, but in another specimen

the second abscissa of the radius is much more than half as long

as the third, and both recurrent nervures are received nearer the

base of the cells.

21. ANTHOBOSCA ERYTHRONOTA Cam. (Pl. LX XXI. figs. 5, 6;

Pl. LX XXII. fig. 4.)

Plesia erythronota Cam. Rec. Albany Mus. i. 5, p. 320 (1905).

Q. Nigra; prothorace. mesothorace, tibiis tarsisque rufo-ferru-

gineis ; segmentis abdominalibus 2-4 macula laterali alba utrinque ;

alis fusco-hyalinis, venis nigris.

3. Niger, gracilis; mandibulis basi, clypeo, oculis margine

interiore, pronoto late postice, tegulis, mesonoto macula, mesopleuris

macula, scutello macula mediana, postscutello macula mediana

transversa, segmentisque abdominalibus 2-4 macula magna trans-

versa utringue albido-flavis ; pedibus nigris albo-variegatis ; alis

hyalinis, venis nigris.

Variat 2 prothorace mesothoraceque nigris.

Long., 2 10-11 mm., g 8-11 mm.

@. Clypeus shining in the middle and somewhat flattened,

narrowly truncate at the apex. Head convex, broader than long,

not much broader than the thorax; rather sparsely punctured,

the space round the base of the antenne closely punctured and

clothed with fulvous hairs. Eyes touching the base of the

mandibles, the line of the interior margin slightly undulating ;

posterior ocelli much nearer to each other than to the eyes.

Thorax rather sparsely punctured ; pronotum slightly emarginate

anteriorly, the posterior margin widely but not strongly arched ;

the projection of the mesosternum between the intermediate cox

very deeply bilobed. Median segment very finely and closely

STUDIES IN THE FOSSORIAL WASPS. 739

punctured, the posterior slope very steep. Abdomen rather

closely punctured; the apical margins of the segments rather

broadly depressed and smooth, more broadly in the middle than

at the sides, the apical segment testaceous and thickly covered

with fulvous bristles. Radial cell bluntly rounded at the apex,

not detached from the costa; second abscissa of the radius a

little longer than the first, but scarcely more than half as long as

the third, the third cubital cell longer on the radius than on the

cubitus, second recurrent nervure received at about one-quarter

from the base of the third cubital cell. Ungues cleft.

3. Clypeus slightly convex, rather narrowly truncate at the

apex and armed with a row of three very minuteteeth. Antenne

no longer than the thorax and median segment combined, tapering

slightly towards the apex. Eyes convergent towards the clypeus,

the inner margin almost straight. Head and thorax finely and

closely punctured ; pronotum strongly narrowed anteriorly, the

anterior margin slightly emarginate, posterior margin strongly

arched. Abdomen shagreened, very slender, slightly tapering

to the extremities, the basal segment distinctly longer than

the second. Hind tibie finely serrate, all the tarsal ungues

cleft. Second abscissa of the radius half as long again as the first

and only a little more than half as long as the third, second

recurrent nervure received at about one-quarter from the base of

the third eubital cell.

Hab. Willowmore, Cape Colony; November to January

(Dr. Brauns).

The male comes very near Cameron’s genus Odontothynnus,

which I have elsewhere treated with some doubt as a synonym of

Anthobosca; but Cameron states that the posterior tarsal ungues

in his genusare simple. Even if he is correct as to this character

it would not be sufficient to justify the formation of a genus on

one sex only, and his remarks show that he has no knowledge of

the genus Anthobosca. I suspect that the present species may

prove. to be identical with Plesia leucospila Cam., with the

description of which it agrees fairly well, but the mesopleure

are rather strongly though not very closely punctured, not almost

smooth as in Cameron’s description. In the broadly rounded

apex of the radial cell this species differs from dA. natalica

Turn., in which the cell is subacute, but agrees with A. arabica

Turn.

*92. ANTHOBOSCA LEUCOSPILA Cam.

Plesia leucospila Cam. Rec. Albany Mus. i. 5, p. 319

(1905), @.

@. It is almost certain that Cameron has misplaced this

species. It is possibly identical with A. erythronota Cam., which

has a similar colour variety, in which case the name leucospila

should be used for the species. But Cameron states that the

pleure are almost smooth, whereas in erythronota the mesopleure

740 MR. R. E. TURNER ON

are strongly though not closely punctured. But this is possibly

an error in Cameron’s description.

*23. ANTHOBOSCA SAUAKINENSIS Magr.

Myzine sawakinensis Magr. Ann. Mus. Civ. Genova, xxi. p. 560

(1884), 9

I think it probable that A. arabica Turn. is only a colour variety

of this species, with the abdomen black instead of ferruginous.

The difference in size between the two forms is considerable, but

both size and colour vary much in this genus. But as I have not

seen sauakinensis I think it better to keep the two separate at

present.

24, ANTHOBOSGA ARABICA Turn.

Anthobosca arabica Turn. Trans. Ent. Soc. London, p. 397

(USHUG) e

@. Radial cell broadly rounded at the apex ; second abscissa

of the radius longer than the first, the two combined equal in

length to the third, first recurrent nervure received a little before

the middle of the second cubital cell, second before one-third from

the base of the third cubital cell, which is as long on the radius

as on the cubitus. Tarsal ungues bifid; the lobe beneath the hind

femora occupying nearly the whole length of the joint and scarcely

rounded ; basal joint of fore tarsi with a comb of nine rather short

spines on the outer margin and a row of short fine spines on the

inner margin, the outer angle strongly produced and almost

reaching the apex of the second joint. Shining, finely and sparsely

punctured, more closely on the front and pronotum.

Black, with grey pubescence, a narrow band on the inner margin

of the eyes, continued and arched on the vertex, a spot on the

front, hind margin, and anterior angles of the pronotum, a curved

band on the scutellum, a median spot and the posterior angles on

the median segment, and a transverse band on each side on dorsal

segments 1-4 pale yellow; mandibles, tegule, tarsi, and pygidium

testaceous brown.

Length 9 mm.

Hab. Aden district.

25. ANTHOBOSCA MINIMA Turn.

Anthobosca minima Turn. Trans. Ent. Soc. London, p. 398

(1910), 9.

©. Radial cell narrowly truncate at the apex; second abscissa

of the radius more than twice as long as the first, but much shorter

than the third. Tarsal ungues bifid. Shining, sparsely and finely

punctured ; apical dorsal segment strongly punctured and covered

with stiff fulvous hairs.

Black ; mandibles, antenne, and legs testaceous aerate ;

STUDIES IN THE FOSSORIAL WASPS. 741

abdomen dark reddish brown; yellow marks as in arabica, but

the yellow band on each side of the first dorsal segment is

absent.

Length 5 mm.

Hab. Mombasa.

26. ANTHOBOSCA CHILENSIS Guer.

Cosila chalensis Guér, Voy. ‘Coquille,’ Zool. ii. p. 249 (1839), @;

Spinola, in Gay’s Hist. Fis. Chile, Zool. vi. p. 312 (1851), g 9.

Myzine flavicornis Sm. Descr. new spec. Hymen. p. 183

(1879) .52.

@. Radial cell rounded at the apex; second abscissa of the

radius twice as long as the first, but distinctly shorter than the

third; first recurrent nervure received close to the middle of

the second cubital cell, second at about one-fifth from the base

of the third cubital cell. Tarsal ungues bifid; lobe beneath the

apical third of the hind femora scarcely rounded, basal joint of

hind tarsi with a thinly-set row of very short spines beneath.

Finely and rather closely punctured, median segment finely

rugulose ; abdomen shining, very finely and sparsely punctured.

Basal joint of fore tarsi not strongly produced at the outer apical

angle.

Black, with long black pubescence ; calearia black; flagellum

bright orange. Wings fusco-violaceous.

3. Third abscissa of the radius twice as long as the second in

some specimens, shorter than the second in others ; first recurrent

nervure received by the second cubital cell at a distance from the

base slightly less than the length of the first transverse cubital

nervure, second either interstitial with the second transverse

cubital nervure or received a little before the apex of the second

eubital cell. Apical joint of antenne no longer than the

penultimate. Basal abdominal segment nearly twice as broad

at the apex as long. Very finely and closely punctured, minutely

on the abdomen.

Colours as in the female.

Length, 2 22mm., ¢ 16 mm.

Hab. Chile.

27. ANTHOBOSCA CARBONARIA Burm.

Myzine carbonaria Burm. Stett. ent. Zeit. xxxvii., p-. 168

(1876); 2:

Anthobosca carbonaria Turn. Trans, Ent. Soc. London, p. 83

(1908), @. |

Cosila carbonaria Bréthes, Ann. Mus. Buen. Aires, xx. p. 256

(GEE) Sees

@. Radial cell narrowly rounded at the apex; second abscissa

of the radius longer than the first and third combined; first:

recurrent nervure received at about two-fifths from the base of

742 MR. R. E. TURNER ON

the second cubital cell, second before one-third from the base

of the third cubital cell, which is nearly twice as long on the

cubitus as on the radius. Basal joint of hind tarsi with a row of

fine spines beneath; basal joint of fore tarsi with seven spines

above and a row of short spines beneath; tarsal ungues cleft ;

apical half of hind femora broadly rounded beneath ; tarsal ungues

cleft. Shining, finely and sparsely punctured; front and pro-

notum more closely and coarsely punctured; median segment

subopaque, very finely and closely punctured, the posterior slope

rather indistinctly transversely striated. Pygidium densely clothed

with fusco-ferruginous pubescence. Basal joint of fore tarsi not

very strongly produced at outer angle.

Black, with grey pubescence ; calearia pale brownish. Wings

rather light fuscous, fusco-hyaline at the apex.

Length 20 mm.

Hab. Nova Fribourg, 8. Brazil.

28. ANTHOBOSCA BIPUNCTATA Perty.

Tiphia bipunctata Perty, Delect. anim. artic. Brasil. p. 139

(1833), 2.

Myzine bipunctata Sm. Cat. Hym. B. M. iii. p. 76 (1855).

Anthobosca bipunctata Turn. Trans. Ent. Soc. London, p. 83

(1908).

Q@. Radial cell narrowly rounded at the apex; first and fourth

abscisse of the radius about equal in length, second and third also

nearly equal to each other and twice as long as the first; the first

recurrent nervure received just beyond two-fifths from the base of

the second cubital cell, second at the middle of the third cubital

cell, which is shorter on the radius than on the cubitus. Tarsal

ungues bifid; lobe beneath the posterior femora extending over

the apicai two-thirds of their length and scarcely rounded. Shining

and very sparsely punctured, the front smooth, apical dorsal seg-

ment covered with long fuscous hair. Basal joint of fore tarsi

strongly produced at the outer apical angle.

Black ; a yellow spot on each side of the third dorsal segment.

Wings fusco-hyaline, nervures fuscous; calcaria whitish.

Length 16 mm.

Hab. Minas Geraes, Brazil.

Allied to carbonaria Burm., but has the lobe beneath the hind

femora longer, the basal joint of the fore tarsi is more produced,

and the puncturation is sparser. A. bipustulata Turn. is probably

the male of this species.

29. ANTHOBOSCA ALBOMACULATA Sm. (PI. LXXXT. figs. 7, 8.)

Myzine albomaculata Sm. Descr. new spec. Hymen. p. 181

(1879), Og.

Myzine lecointet Diicke, Rev. entom. p. 146 (1907), 9

@. Radial cell narrowly rounded at the apex; second abscissa

STUDIES IN THE FOSSORIAL WASPS. 743

of the radius a little longer than the first, the third longer than

the first and second combined ; first recurrent nervure received

just beyond two-fifths from the base of the second cubital cell,

second close to the middle of the third cubital cell. Tarsal ungues

bifid ; lobe beneath the basal half of the hind femora not very

prominent and very feebly rounded; basal joint of hind tarsi

with a row of short hairs beneath; basal joint of fore tarsi

strongly produced at the outer apical angle. Shining and sparsely

punctured ; median segment opaque, closely and minutely punc-

tured; apical dorsal segment coarsely punctured and _ thickly

clothed with long fuscous hairs.

Black ; a spot on each side near the anterior angles of the pro-

notum, a spot on the mesonotum, a spot at the base of the median

segment and one at each of the posterior angles, and a spot on each

side of the second and third dorsal segments yellowish white ;

calearia whitish. Wings pale fusco-hyaline, nervures fuscous.

3. Second abscissa of radius twice as long as first, third about

half as long again as second; first recurrent nervure received a

little beyond the middle of the second cubital cell, second at two-

fifths from the base of the third cubital cell. Tarsal ungues bifid ;

hind tibiz serrate on the outer margin. Basal abdominal segment

nearly half as long again as the breadth at the apex. Finely and

rather closely punctured, abdomen finely shagreened.

Black ; mandibles, clypeus, scape beneath, a spot on each side

of the pronotum, a spot on the mesonotum, a spot on the

scutellum, one on the postscutellum, another at the apex of

the median segment, the base of the tibiz, and the tarsi pale

yellow. Wings hyaline iridescent, nervures nearly black.

Length, 9 12 mm., ¢ 9 mm.

Hab. Amazon, from Para to Ega.

30. ANTHOBOSCA ANTENNATA Sm.

Anthobosca antennata Sm. Descr. new spec. Hymen. p. 174

(1879), o.

Cosila jheringi Saussure in Grandidier, Hist. Madagascar, xx.

p. 234 (1892), ©.

Thynnus antennatus D. 'T. Cat. Hymen. viii. p. 101 (1897), 3.

@. Radial cell very narrowly rounded at the apex; third

abscissa of radius half as long again as the second, which is

nearly twice as long as the first, the fourth shorter than the first ;

first recurrent nervure received a little before the middle of the

second cubital cell, second at one-fifth from the base of the third

cubital cell. Tarsal ungues bifid ; lobe beneath the hind femora

commencing near the base and extending to the apex, broadly

rounded ; basal joint of bind tarsi with a row of very short fine

spines beneath ; basal joint of fore tarsi strongly produced at the

outer apical angle. Shining, very finely and rather sparsely

punctured ; median segment subopaque, very closely and minutely

744 MR. R. E. TURNER ON

punctured; apical dorsal segment densely ciothed with long fulvous

hairs.

Black ; mandibles at the base, antenne, and legs ferruginous ;

an interrupted band on the posterior margin of the pronotum, a

spot on the mesonotum, one at the base of the median segment

and another at each of the posterior angles, and an interrupted

band on the four basal dorsal segments, more broadly interrupted

on the second than on the other segments, yellow. Wings very

pale flavo-hyaline, nervures ferruginous.

3d. Third abscissa of the radius half as long again as the second,

which is about equal to the fourth and twice as long as the first ;

first recurrent nervure received at or a little beyond the middle

of the second cubital cell, second at one-third from the base of the

third cubital cell. Tarsal ungues bifid; hind tibie serrate on the

outer margin. Antenne short and stout, tapering to the apex;

pronotum rounded anteriorly ; first abdominal segment a little

longer than the breadth at the apex, the apical dorsal margin of

the segment broadly rounded. Finely and closely punctured ;

abdomen finely shagreened.

Black; mandibles, clypeus, seape beneath, posterior margin of

the pronotum broadly, a spot on the mesonotum, one on the

seutellum, another on the postscutellum, and the greater part of

the tibize and tarsi yellow. Wings hyaline, nervures fuscous.

Length, 9 13-16 mm., ¢ 12-14 mm.

Hab. S. Brazil; Rio Grande do Sul.

As I have noticed before, the antennz in the type of antennata

are a little shorter and stouter than in the males sent with jhering?.

This difference may possibly prove to be specific, but I do not think

it is.

-*31. ANTHOBOSCA ERYTHROPYGA Burm.

Myzine erythropyga Burm. Stett. ent. Zeit. xxxvii. p. 169

(L876) On Sus

Anthobosca erythropyga Turn. Trans. Ent. Soc. London, p. 83

1908).

Costa erythropyga Bréthes, Ann. Mus. Buen. Aires, xx. p. 256

(1910).

I have not seen this species. Schrottky suggests that it is

identical with A. gheringi Sauss., but Burmeister’s description

gives the legs of the male as red, of the female black, and the five

basal abdominal segments in both sexes with lateral yellow spots,

whereas in jheringi the four basal segments are banded with yellow

and the legs red in the female, and the male has the abdomen

without yellow marks and the legs yellow and black. I do not

think there is sufficient ground for considering the two to be

identical, especially as Schrottky had not seen specimens of

jheringi. ;

STUDIES IN THE FOSSORIAL: WASPS, 745

*32. ANTHOBOSCA APICALIS Sichel.

Cosila apicalis Sichel, Sauss. et Sichel, Cat. spec. gen. Scolia,

p- 262 (1864), 9.

I have not seen this species, which may possibly be identical

with carbonaria Burm., but the dorsal segments of that species

are not reddish laterally.

Males.

1. Species from the Old World........5:......c0ce0ssec0bssetieoe 2.

Species from South America... ........0..6...csccceceeeeesseee 18.

awe AUSUUAlTANE SPECIES Ln siaeeateanustsatorostcie tac cteme eee Si

PAI ICATIESDECICR Mas ret tenee tte ree ice ie ee 14,

3. Tarsal ungues with a blunt lobe at the base; second :

and third cubital cells each receiving a recurrent

INSAALLIG) ces ood dhn Nebronaee Ihde Aadtsiadnaro nee SOORERODAR ARE HE OE rse 4.

Tarsal ungues bitid, without a basal lobe ; second cubital :

cell receiving both recurrent nervures....... A, crassicornis Sm.

Highth and three following joints of the flagellum sub-

tuberculate beneath, apical joint longer than penulti-

mate and slightly curved. Wings fusco-violaceous... 4. australis Sichel

None of the joints of the flagellum subtuberculate (=nigripennis Sw.)

beneath, apical joint no longer than penultimate.

Wings hyaline or subhyaline .....0......5.. 0000s. s0eccs eee 5.

5. First abdominal segment distinctly longer’ than the

second ; fully half as long again as its breadth at the

GI DDC ndrmenog aapaog sod nadbod endedeebnndneds ORGEBEs ceBDAE MNAnPRE alm lA CUTE A air icra XG nutcst-

First abdominal segment not distinctly longer than the :

second; scarcely, if at all, longer than its apical

buen are feeicc. at cae Nun ABU. cy ceneealioe oy Gln 6.

6. Thorax and abdomen wholly black ..............0ce0 se ece eee Cry

Pronotum at least marked with yellow .......0...0...000005 i).

Pap VES StErMPS MOUS Pe, Fei suslereyssetlssnvarse-reesdsstenensseisdeen 145 varipes Sin:

MOB SHDIAC ie Pease cecienn yee Ment AR MR I a OL gale :

8. Median segment transversely rugulose, very short and

broad ; anterior tibize black beneath ..................... A. @thiops Sm.

Median segment finely punctured, not very short or

broad ; anterior tibiee ferruginous beneath ............ A. nigra Sm,

DERcoROrnUSINOUS am an etn nk TPS dE 10.

Legs black, more or less marked with yellow............... 11.

10. Postscutellum yellow, legs marked with yellow; first

transverse cubital nervure as long as the second or

longer and strongly bent near the cubitus ............ A. torresensis Turn, —

Postscutellum and legs without yellow markings ; first i j

transverse cubital nervure much shorter than the ;

second, and not bent near the cubitus.................... 4. gilest Turn.

11, Five basal abdominal segments with yellowish lateral

spots; cubitus of hind wing originating just before

the transverse median nervure, almost interstitial ... 4. moderata Turn.

Five basal abdominal segments immaculate; cubitus

of hind wing originating very distinctly before the

transverse median nervure ............cceceesec cee seeeee ues 12.

12. Seventh dorsal segment with yellowish lateral spots ;

postscutellum yellowish .........0....cesesese-sscrerecctere 13.

Seventh dorsal segment and postscutellum immaculate . A. JSrenchi Turn.

13. Clypeus black ; mesonotum without a yellow spot ; hind

Liptoa ches blivg SPINS.) tcc cups -paeea nphwasten a mdteo ree A. longipalpa Turn,

Clypeus yellow ; mesonotum with a yellow spot; hind

tibieo feebly serwaties 3..\.s45.5.s0.csuecs hoes Mates A, lagardet Turn.

14, Apical margin of clypeus not toothed ............ 0.000600, 15.

Apical margin of clypeus with two small teeth

Proc. Zoou, Soc,—1912, No, XLIX.

746 MR. R. E. TURNER ON

15. Legs wholly ferruginous; mesonotum with a yellow

‘spot ; tegulee black... .. A. errans Sm.

Legs ferruginous marked with “yellow; “mesonotum

3 ‘immaculate ; begullae ty ellOW: i500... casts -ce seeseeecteeeieee A. flavopicta Turn.

17.

16. Wings hyaline’ ba:

Apical half of wings “with a distinct fusco-violaceous

tinge .... SEAN Ende pene co cacuenasbancerMenanencnanse, wala Uilaonertan Orban

17. Abdomen immaculate... A. lacteipennis Cam.

Dorsal abdominal segments ‘2-4 with | large yellowish

lateral spots livers sect castonsste cece aoc teete aadeet meeoate A. erythronota Cam.

18. Antennze orange; wings fusco-violaceows .................. A. chilensis Guér.

Antennee black; wings hyaline ...............0..ceecee eee eee 19.

19. Abdomen spotted with yellow ............... ccc ceeeeeseeee ees *A. erythropyga Burm.

Abdomen nmmiac ull ate Wee seeceet eee tise teats eee eee 20.

BON Megs TerrusimoOus) tie i ecresay Baek ane ta Bees nea se decease A bipustulata Turn.

Legs black, marked with yellow ........ brs 21.

21. Length 13mm. Yellow band on pronotum entire ...... A. antennata Sm.

Length 10 mm. Yellow band on pronotum interrupted. A. albomaculata Sm.

33. ANTHOBOSCA AUSTRALASIZ Guér. (Pl. LX XXIII. fig. 13.)

Anthobosca australasie Guérin; Duperry, Voy. ‘ Coquille,’ Zool.

li. p. 237 (1839), ¢

Anthobosca crabroniformis Sm. Cat. Hym. B. M. vi. p. 59

(1859), 3.

Thynnus cathreinii D. T. Cat. Hym. viii. p. 103 (1897), 3.

3. Pronotum much narrowed anteriorly, nearly as long as the

scutellum. First dorsal abdominal segment slender, more than

half as long again as the apical breadth, much longer than the

second segment. Seventh dorsal segment very broadly rounded

at the apex. Hind tibie feebly spined. Third abscissa of the

radius longer than the first and second combined ; first recurrent

nervure received close to the middle of the second cubital cell,

second at one-quarter from the base of the third cubital cell ;

first transverse cubital nervure curved, not sharply bent at the

base. Cubitus of the hind wing originating before the transverse

median nervure, separated from it by a distance exceeding half

the length of that nervure.

Black ; mandibles, clypeus, margins of the eyes interrupted on

the summit, pronotum very broadly posteriorly, tegule, a

quadrate spot on the mesonotum, a spot on the mesopleure, the

middle of the scutellum and postscutellum, a curved transverse

band at the apex of the median segment, an interrupted trans-

verse band on dorsal segments 2—4, sometimes also a spot on each

side of the first and fifth segments, a spot on the anterior cox

and a line on the anterior tibie yellow; legs ferruginous. Wings

hyaline, tinged with yellow ; nervures ferruginous.

Length 16 mm.

Hab. New South Wales, Queensland, as far north as Cairns.

The female is unknown, but the range of the species is about

the same as that of signata Sm.

STUDIES IN THE FOSSORIAL WASPS. TAT

34. ANTHOBOSCA GILESI Turn.

Anthobosca gilesi Turn. Proce. Zool. Soc. London (1910),

p. 308, ¢.

¢. Antenne shorter than the thorax and median segment

combined ; pronotum much narrowed anteriorly, shorter than

the scutellum. First dorsal segment as broad at the apex as

long, a little shorter than the second segment. Seventh dorsal

segment subtruncate at the apex. Neuration similar to that of

A. australasie, but the position of the first recurrent is variable—

it is received beyond the middle of the second cubital cell in the

type, but at the middle in another specimen.

Black; the base of the mandibles, part of the clypeus, the

pronotum very broadly posteriorly, and the tegule yellow;

legs ferruginous. Wings hyaline, nervures fuscous. Calcaria

white.

Length 11-14 mm.

Hab. Perth, West Australia.

35. ANTHOBOSCA TORRESENSIS Turn.

Anthobosca torresensis Turn. Proc. Linn. Soc. N.S.W. xxxii.

p- 518 (1907), ¢.

3. Pronotum narrowed anteriorly, nearly as long as the

scutellum. First dorsal segment as broad at the apex as long,

no longer than the second segment; seventh dorsal segment

narrowly truncate at the apex. Second abscissa of the radius

shorter than the first, the two combined much shorter than the

third. First transverse cubital nervure longer than the second,

very sharply bent near the cubitus; first recurrent nervure

received close to the middle of the second cubital cell, second

beyond one-quarter from the base of the third cubital cell.

Black; legs ferruginous; mandibles at the base, scape beneath,

pronotum very broadly posteriorly, postscutellum, and a line on

the anterior tibiz yellow. Wings hyaline, nervures fuscous.

Calearia whitish.

The hind tibiz are almost smooth, the spines nearly obsolete.

Neuration of hind wing as in A. australasie.

Length 7-8 mm.

Hab. Cape York, Queensland.

36. ANTHOBOSCA VARIPES Sm.

Anthobosca varipes Sm. Cat. Hym. B. M. vi. p. 59 (1859), 3.

Thynnus fischeri D. T. Cat. Hym. viii. p. 106 (1897), 3.

g. Pronotum shorter than the scutellum, not very strongly

narrowed anteriorly ; scutellum with a delicate carina from the

base to the apex. First dorsal segment nearly as broad at the

apex as long, very little longer than the second segment;

seventh dorsal segment broadly subtruncate at the apex. Hind

tibie very feebly serrate. First transverse cubital nervure

748 MR. R. E. TURNER ON

oblique, not sharply bent near the cubitus, longer than the

second. Second abseissa of the radius about twice as long as

the first, the two combined shorter than the third. First

“reat nervure received distinctly beyond the middle of the

second cubital cell, second beyond one-third from the base of

the third cubital cell. Hind wing as in A. australasie. Apical

joint of maxillary palpi as in A. australasie, not filiform and

scarcely longer than the penultimate joint.

Black ; the legs ferruginous. Wings hyaline, nervures

fuscous.

Length 12-15 mm.

Hab, Adelaide, 8. Australia; Ararat and Ringwood, Victoria.

_ 87. ANTHOBOSCA NIGRA Sm.

Anthobosca nigra Sm. Cat. Hym. B. M. vii. p. 59 (1859), 3

Thynnus reischu D, T. Cat. Hymen. viii. p. 114 (1897), 3

3. Clypeus truncate at the apex; antenne nearly as long as

the thorax and median segment combined. Pronotum short,

scarcely more than half as*long as the scutellum, very slightly

narrowed anteriorly. First dorsal segment longer than the apical

breadth ; seventh dorsal segment broadly rounded at the apex.

Hind tibie rather feebly serrate ; hind femora without a lobe at

the apex beneath. Second abscissa of the radius longer than the

first, the two combined much shorter than the third; first

transverse cubital nervure oblique, sharply bent close to the

cubitus; first recurrent nervure received at the middle of the

second cubital cell, second at two-fifths from the base of the third

cubital cell. The distance between the cubitus of the hind wing

and the transverse median nervure is greater than half the length

of that nervure.

Black ; the fore tibie ferruginous within; calcaria whitish.

Wings hyaline, nervures fuscous.

Length 9-12 mm.

Habs Victoria and South Australia.

Differs from varipes in the absence of a carina on the scutellum,

in the colour of the legs, and the.more distinct serration of the

hind tibie.

38. ANTHOBOSCA ZTHIOPS Sm.

Anthobosca cthiops Sm. Descr. new spec. Hymen. p. 175

(1879), ¢.

Thynnus stolzii D. T. Catal. Hymen. viii. p. 116 (1897), 3

g. Antenne stouter than in other Australian species ;*pro-

notum very slightly narrowed anteriorly, shorter than the

secutellum. Head and thorax very closely and: finely punctured,

median segment transversely rugulose, short and not as strongly

convex as in other species. First: dorsal segment very little:

longer than the apical breadth ; seventh dorsal ‘Segment. narrowly

STUDIES IN THE FOSSORIAL WASPS. 749

truncate at the apex. Hind tibie spinose; hind femora without

a rounded lobe at the apex beneath. Second abscissa of the

radius equal to the first, the two combined shorter than the third ;

first transverse cubital nervure oblique, sharply bent close to the

cubitus; first recurrent nervure received close to the middle of

the second cubital cell, second just before one-third from the

apex of the third cubital cell. Cubitus of the hind wing

separated from the transverse median nervure by a distance equal

to one-third of the length of that nervure.

Black; the inner margin of the eyes narrowly whitish; calcaria

whitish ; wings hyaline, tinged with fuscous, nervures fuscous.

Length 14 mm.

Hab. Champion Bay, W. Australia.

39. ANTHOBOSCA MODERATA Turn. (Pl. LX XXI. fig. 4.) :

Anthobosca moderata Turn. Ann. & Mag. Nat. Hist. (8) i.

p- 482 (1909), 3.

g. A minute tubercle in the middle of the clypeus; pronotum

not much narrowed anteriorly, not more than half as long as the

scutellum ; first dorsal segment a little narrower at the apex than

long, a little longer than the second segment. Hind tibie very

distinctly spinose ; hind femora with a very distinct rounded lobe

at the apex beneath. Second abscissa of the radius longer than

the first, the two combined about equal to the third; first

recurrent nervure received at the middle of the second cubital

cell, second beyond one-third from the base of the third cubital

cell. Cubitus of the hind. wing originating just before the

transverse median nervure.

Black; mandibles at the base, clypeus, posterior margin of |

pronotum, tegule, a spot on the mesonotum, postscutellum, a

spot on each side at the apex of the median segment, a small

lateral spot on each side of dorsal segments 1—5, a large spot on

each side of the seventh segment, the base of the tibiz and

the base of the tarsi whitish yellow; the apex of the seventh

dorsal segment testaceous. Wings hyaline, tinged with fuscous ;

nervures fuscous.

Length 12 mm.

Hab. Townsville, Queensland.

40. ANTHOBOSCA FRENCHI Turn.

Anthobosca frenchi Turn. Proc. Linn. Soc. N.S.W. xxxii.

p- 518 (1907), 3.

3. Clypeus truncate at the apex; pronotum much shorter

than the scutellum, very slightly narrowed anteriorly. First

dorsal segment no longer than the apical breadth ; seventh dorsal

seement broadly rounded at the apex. Hind tibie very feebly

serrate; hind femora without a lobe at the apex beneath.

Second abscissa of the radius longer than the first, the two .

759 MR. R. E. TURNER ON

combined as long as the third; first transverse cubital nervure’

oblique, strongly bent near the cubitus, longer than the second :

first recurrent nervure received at the middle of the second

cubital cell, second at one-third from the apex of the third cubital

cell; the distance between the cubitus of the hind wing and the

transverse median nervure equal to less than half the length of

that nervure.

Black; base of the mandibles, a small spot round the base of

each antenna, the hind margin of the pronotum very narrowly,

tegule, anterior tibiz in front, and the base of the hind tibiz pale

yellow ; apex of the mandibles and fore tarsi fusco-ferruginous.

Wings hyaline, nervures dark fuscous.

Length 7-8 mm.

Hab. Victoria.

41, ANTHOBOSCA LAGARDEI Turn.

Anthobosca lagardei Turn. Trans. Ent. Soc. London (1908),

p: 86, 3.

3. Differs from A. frenchi in the yellow clypeus, yellow spot

on the mesonotum, postscutellum, apex of median segment, and

on each side of the seventh dorsal segment ; in the rounded lobe

at the apex of the hind femora beneath, which is well developed

in lagardet, and in the yellow basal joints of the tarsi.

Length 8 mm.

Hub. Sydney, New South Wales.

42. ANTHOBOSCA LONGIPALPA Turn.

Anthobosca longipalpa Turn. Proc. Linn. Soc. N.S.W. xxxil.

pole ds0n). Se

6. Clypeus very feebly emarginate at the apex; the apical

joint of the maxillary palpi filiform, longer than the penultimate

joint, the three apical joints much more slender than the basal

ones. Pronotum short, very slightly narrowed anteriorly. First

dorsal segment a little longer than the apical breadth ; seventh

dorsal segment narrowly truncate at the apex. Hund tibie feebly

spinose; the lobe at the apex of the hind femora beneath not

distinct. Second abscissa of the radius nearly twice as long

as the first, the two combined longer than the third. First

transverse cubital nervure curved, much longer than the second,

and rather sharply bent near the cubitus; recurrent nervures

received a little before the middle of the second and third cubital

cells. Cubitus of the hind wing originating before the transverse

median nervure, the distance between it and that nervure equal

to about one-third of the length of the transverse median

nervure.

Black; posterior margin of the pronotum very narrowly, a

transverse line on the postscutellum, a spot on each side of the

seventh dorsal segment, and the base of the hind tibize whitish

STUDIES IN THE FOSSORIAL WASPS. 751

yellow ; tibize fusco-ferruginous ; calearia white. Wings hyaline,

nervures fuscous.

Length 12 mm.

Hab, Cairns, Queensland.

43. ANTHOBOSCA CRASSICORNIS Sm. (Pl. LXXXTI. fig. 1.)

Tachypterus crassicornis Sm. Cat. Hym. B. M. vil. p. 64

(1859), ¢.

Trachypterus crassicornis D. T. Cat. Hymen. viil. p. 120

(iso); 'C..

Anthobosca crassicornis Turn. Proc. Linn. Soc. N.S.W. xxxil.

p. 519 (1907), 3.

3g. Clypeus produced in the middle and rather narrowly trun-

cate at the apex. Antenne slightly longer than the thorax and

median segment combined. Head and thorax very closely and

not very finely punctured. First abdominal segment scarcely

longer than the second ; the apex of the dorsal segment rounded

and a little, but not very distinctly, narrower than the first

segment is long. Posterior tibie very distinctly serrate, tarsal

ungues bifid. Second abscissa of the radius longer than the

third; both recurrent nervures received by the second cubital

cell, the first at two-fifths from the base, the second at about

nine-tenths or more from the base. Third cubital cell as long on

the radius as on the cubitus.

Black, with long grey pubescence; mandibles, clypeus, lower

portion of the inner margin of the eyes, anterior coxe, trochanters

and femora beneath, and a line beneath the intermediate femora

yellow ; second and third abdominal segments and the base of

the fourth, tibie, tarsi, hind femora, and intermediate femora

above ferruginous. Wings hyaline, nervures fusco-ferruginous,

stigma fuscous.

Length 13 mm.

Hab. Australia.

A variety has the ferruginous colour much obscured. As

mentioned under that species, I believe this to be the male of

A. clypeata Sm , owing to the similarity of the structure of the

tarsal ungues, the position of the recurrent nervures, the length

of the second abscissa of the radius, and some similarity of

colour.

44, ANTHOBOSCA ERRANS Sm.

Anthobosca errans Sm. Descr. new spec. Hymen. p. 174

(1879), ¢.

3. Clypeus broadly truncate at the apex, not toothed; an-

tenn short and stout. Pronotum shorter than the scutellum,

distinctly narrowed anteriorly ; scutellum and median segment

obscurely longitudinally carinated in the middle. First dorsal

segment longer than the apical breadth; seventh dorsal seement

752 MR. R. E. TURNER ON

narrowly truncate at the apex. Hind tibie serrate ; hind femora

with a rounded lobe at the apex beneath. Second abscissa of the

radius nearly twice as long as the first, the two combined shorter

than the third; first transverse cubital nervure oblique, strongly

bent close to the cubitus, longer than the second; the distance

between the cubitus of the hind wing and the transverse median

nervure equal to about one-third of the length of that nervure

Tarsal ungues bifid.

Black ; “base of the mandibles, clypeus, a spot on each side of

the pronotum, a spot on the mesonotum, one on the scutellum

and a transverse line on the postscutellum yellow; legs ferru-

ginous. Wings hyaline, nervures fusco-ferruginous.

Length 13 mm.

Hab. Zululand.

45. ANTHOBOSCA FLAVOPIOTA Turn.

Anthobosca flavopicta Turn. Trans. Ent. Soc. London, p. 399

(1910), 3d pl. L. fig. 4.

3. Clypeus subtruncate at the apex, not toothed. Posterior

tibie serrate, tarsal ungues bifid.

Black; clypeus with a black spot in the middle, inner margin

of the eyes narrowly, a small spot behind the eyes near the

summit, a broad band on the posterior margin of the pronotum,

tegule, a large spot on the scutellum, one on the postscutellum,

the apex of the fore femora, and the anterior and intermediate

tibie and tarsi above pale yellow; tibie and tarsi ferruginous

brown. Wings hyaline, nervures black.

Length 11 mm.

Hab. Zanzibar.

46. ANTHOBOSCA BIDENTATA Cam.

On bidentatus Can. Eee Albany Mus. i. p. 162

(1904), 3

3. Apex of clypeus bidentate. Hind tibie serrate. First

transverse cubital nervure rounded. Wings hyaline, the apical

half fusco-violaceous.

Black ; lower half of inner orbits of the eyes, face, clypeus,

base of mandibles, base and outer side of fore and intermediate

tibiz, calearia, tarsi, base of hind tibi, and the posterior margin

of the pronotum broadly yellow.

Length 11-12 mm.

Hab. Grahamstown, 8. Africa.

47. ANTHOBOSCA LACTEIPENNIS Cam.

Odontothynnus lacteipennis Cam. Rec. Albany Mus. i. p. 162

(1904), 3

3. Apex of clypeus bidentate. Hind tibiew serrate. First

STUDIES IN THE FOSSORIAL WASPS. 753

transverse cubital nervure oblique.

white.

Colours as in bidentata,

Length 10 mm.

Hab. Grahamstown, 8. Africa.

I have seen neither bidentata nor lacteipennis. Cameron’s

statement that the ungues of the hind tarsi are simple, not bifid,

needs confirmation.

Wings hyaline, nervures

48. ANTHOBOSCA BIPUSTULATA Turn.

Anthobosca bipustulata Turn. Ann. & Mag. Nat. Hist. (8) iv.

p. 843 (1909), 3.

d. Slender; antenne stout, tapering towards the apex;

pronotum scarcely more than half as long as the scutellum,

moderately narrowed anteriorly. First dorsal segment nearly half

as long again as the apical breadth; seventh dorsal segment

rather narrowly rounded at the apex. Tarsal ungues bifid ;

hind tibie serrate ; hind femora without a distinct lobe at the

apex beneath. Apical joint of maxillary palpi filiform. Second

abscissa of the radius twice as long as the first, the two combined

nearly as long as the third; first transverse cubital nervure

oblique, sharply bent near the cubitus, longer than the second;

the distance between the cubitus of the hind wing and the

transverse median nervure equal to half the length of that

nervure.

Black: mandibles, clypeus, scape beneath, a spot on each side

on the posterior margin of the pronotum, the base of the tegule,

a small spot on the mesonotum, one on the scutellum, another on

the postscutellum, the two latter spots sometimes absent, and the

anterior coxe beneath yellow; legs ferruginous. Wings hyaline,

nervures fuscous.

Length 8-12 mm.

Hab. Barbacena, Minas Geraes.

A variety has the legs black, marked with yellow.

EXPLANATION OF THE PLATES.

Prats LXXXI.

Fig. Fig,

1. Anthobosea crassicornis Sm. 3. 10. Braunsomeria quadraticeps Turn.

2. Anthobosca clypeata Sm. 2. :

3. Anthobosca fastuosa Sm. 2. ll. Myzine stigma Turn. ¢.

4. Anthobosca moderata Turn. 3. 12, Elis (Mesa) alicie Turn. 2.

5. Anthobosca erythronota Cam. ¢. 18. Elis (Mesa) longiventris Turn. ¢.

6. Anthobosca erythronota Cam. 2. 14. Myzine braunsi Turn. 3.

7. Anthobosca albomaculata Sm. 6. 15. Elis (Mesa) dimidiaticornis

8. Anthobosca albomaculata Sm. 2. Bingh. ¢.

9. Braunsomeria quadraticeps Turn. | 16. lis (Mesa) tricolor Sm. 2.

Proc. Zoou. Soc.—1912, No. L,

To4 MR, ABEL CHAPMAN ON

Pratt LXXXII.

Neuration of Wings.

Fig. Fig.

1. Anthobosca australis Sichel. ¢. 9. Elis (Mesa) ruficeps Sm. 9.

2. Anthobosca australis Sichel. ¢. 10. Elis (Mesa) ruficeps Sm. 6-

3. Anthobosca anthracina Sm. &. 11. Elis (Mesa) tricolor Sm. &.

4. Anthobosca erythronota Cam. §. 12. Elis combusta Sm. @.

5. Anthobosca clypeata Sm. &. 13. Myzine stigna Turn. 6.

6. Anthobosca insularis Sm. 2. 14, Myzine braunsi Turn. 6.

7. Braunsomeria quadraticeps Turn. | 15. Myzine constrictiventris Turn. 3.

3. 16. Myzine abdominalis Guér. 9.

8. Elis (Mesa) alicie Turn. &.

Prate LXXXIII.

Exoskeletal Structures.

Fig. 1. Anthobosca australis Sichel. 2. Suture between two basal abdominal

segments.

2, Elis (Mesa) ruficeps Sm. 9. Suture between two basal abdominal

segments.

3. Anthobosca australis Sichel. 6. Apical ventral segment.

4, Elis (Mesa) ruficepsSm. $. Apical dorsal segment.

5. Myzine abdominalis Guér. 3. us 5 ‘S$

6. Anthobosca australis Sichel. 9. Tarsal unguis.

7, Anthobosca clypeata Sm. °. F ¥

8. Anthobosca insularis Sm. 2. Intermediate and hind coxee.

9. Hlis (Mesa) alicie Turn. 2. Basal joint of hind tarsus.

10. Elis (Mesa) ruficeps Sm. 2. 5s 53 #

11. Elis (Mesa) longiventris Turn. J. Two basal segments of abdomen.

12. Myzine constrictiventris Turn. 6.

13. Anthobosca australasie Guér. 6. a is Be

14, Anthobosca australis ae 3. 5

15. Elis (Mesa) ruficeps Sm. 6. a y

16. Elis (Mesa) ruficeps Sm. g: Intermediate and hind coxz.

bb) 39 bby

40. Notes on the Spanish Ibex.

By Apexi Cuapman, F.Z.S.

{Received April 13, 1912: Read May 7, 1912. ]

INDEX.

Page

ED vestorily wait Geshe eect he ste ec etwas ae eee ice one ee eee sae AON (EIS

In Professor Angel Cabrera’s most informative paper on Capra

pyrenaica,* two small points occur to me as worthy of brief

comment. In discussing the present distribution of the Spanish

Wild-Goat in some six isolated colonies, Dr. Cabrera takes

exception to the remark in ‘ Unexplored Spain’ (by Walter Buck,

C.M.Z.S8., and myself) that they had been so isolated “during ages.”

Well, the term used in our former book (‘ Wild Spain’) was

“during centuries,’ and that is certainly more definitive and

probably more accurate. Dr. Cabrera, however, goes on to state

that there exist ‘‘strong reasons for believing that in the past

* P.Z.8. 1911..p. 963.

THE SPANISH IBEX, 15

[inferentially as late as the middle of the seventeenth century—

say 250 years ago] Ibex inhabited every suitable point of almost

every mountain-ridge in Spain.” The only reason actually

adduced, however, is the prevalence of place-names based upon,

or compounded with, the Spanish word Cabra = goat. Such

names, it is true, are ubiquitous; but it would never have

occurred to me that those names necessarily refer to the wild

goat. Spain is a land of goats, and many localities bearing names

such as Sierra de las Cabras, Cabrales, Cebrero, and so on, are not

at all adapted to the nature and requirements of the wild Ibex.

I would suggest that, in many cases, the names merely indicate

the existence of suitable local pasturage for domestic goats, which

are herded everywhere.

Again, Dr. Cabrera translates the Spanish name of the Ibex,

Cabra montés, as equivalent to ‘‘ Mountain-Goat.” Now it would

be nothing less than presumptuous for me, a foreigner with but a

limited colloquial knowledge of the Spanish tongue, to question

his rendering. I donot doso. I accept that as the pure classic

Castilian of Madrid. But I do venture to say that, in wilder

Spain, the term monte, with its derivative adjective montés

(pronounced montéss), possesses quite a different signification.

Monte may occasionally, and in combination, be used to indicate

a hill or mountain ; but in its ordinary provincial sense, it signifies

serub or brushwood. Thus the wild-cat, which is equally common

on lowland or sierra, is known as Gato montés = Scrub-Cat: on

the low-lying plains of Andalucia or Estremadura, the expression

Reses montéses includes all the scrub-haunting animals—such as

deer, wild-boar, lynx, ete.

Now, viewed in this light, it has always appeared to me utterly

inexplicable and incongruous to apply the name Cabra montés, or

Scrub-Goat, to the Ibex of the higher ranges, such as the Sierras

de Grédos and Nevada, where the Ibex live exclusively amidst

rock-regions far above the topmost levels of scrub. But such

incongruity would disappear if Dr. Cabrera’s assumption were

correct, that the Ibex, up toa couple of centuries ago, occupied the

whole vast area shown in the map at p. 965, supra. Was such the

distribution, that name would become appropriate enough, since

an immense proportion of the dotted area consists, not of high

mountains at all, but of low scrub-clad hills. Such country

might appear, to preconceived ideas, totally unsuitable for Ibex :

but we have the fact before us (as fully explained in our books on

Spain) that in several of the lower Mediterranean sierras (some

of which are bush-clad to the summits) the local Ibex do to-day

take kindly to a bush-haunting habit. Indeed, in such situations,

it is obvious, they have no other option.

This latter point tends to support Dr. Cabrera’s assumption,

and equally, of course, undermines our own,

In our two books we had pointed out that the Ibex of the two

extreme ends of Spain (7. ¢., those of the Pyrenees and those of

756 ON THE SPANISH IBEX.

the Mediterranean sierras) most nearly assimilated to each other

in their more flattened and laterally compressed horns*. It is

gratifying to find that our rough field-observations are now

corroborated by Dr. Cabrera’s skilled investigation and careful

cross-sections. But again, it appears anomalous—assuming that

complete isolation only commenced some two centuries ago—that

the central group (now honoured with subspecific rank | as C. p.

victorie) should have developed the greater difference.

The females and young males of the Spanish Ibex are devoid

of the dark dorsal stripe, as is correctly shown in the plate in

‘Unexplored Spain,’ at p. 140. They are of a dun-brown, uni-

colorous in coat as the Spanish Red-Deer ; but Dr. Cabrera 1s quite

justified in criticizing the second plate (op. cit. p. 967), at p. 216.

That ship should be debited, in the first instance, to the artist,

Mr. E. Caldwell, but the fault is wholly mine, since I should have

detected the mistake and had it corrected before passing the

drawing for reproduction.

If permissible to express an opinion on the three beautiful

plates given by Dr. Cabrera, I would say that in life the Spanish

Ibex is rounder and bulkier in the barrel than can ever be

gathered from museum specimens, since skins shrink.

In conclusion, may I express a fervent hope (since interest in

her vanishing Ibex has been aroused in Spain) that further

protection may be extended to the few surviving colonies ?

Within my own time, Ibex have been exterminated in several. of

their earlier haunts. To-day they are at their last gasp in the

Pyrenees and in the Gerez (Portugal). Fortunately, in Grédos,

Morena, and Bermeja, their future has been assured—though

only at the eleventh hour. Can our Spanish friends not see to

safeguarding the much-menaced remnant that yet survives on the

main chain of Nevada 2

* Unfortunately, in ‘ Unexplored Spain,’ in an effort to be concise, and to avoid

repetition, we omitted the word “laterally”; but a reference to ‘ Wild Spain,’

p. 129, makes our meaning clear.

+ Page

maanial Phonsant-breeding. By Rosz Hare oe WAT ae LXIV.-LXVII.) 539

B. Pratt in Dutch New Guinea in 1909-10 ; with Descriptions of new be eget By

Sir Gzorer H, Kevyrice, F.Z.S. (El PAVILE SOS ose s Fp Sey eet he Re hee 546

Society by His Majesty King George. By RB. I. a F.R.S., F.LS., F.Z.S.,

Superintendent of the Gardens. (Text-figs. 66-71.) :.... Peisalerci ete ee eens eesese JOD

Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.—IV. On

_ a Species of Inermicapsifer from the Hyrax, and on the Genera Zschokkeella, Thy-

sanotenia, and Hyracotenia. By Franz E. Bepparp, M.A., D.Sc., F.R.S., F.Z.S.,

E-erosccior. to the Society. (Toxt-figs. 72-83)" ses. sesers cs sdeccecescceauccssecic SIG

82. “Additional Notes on the Living Spécimens of the Australian Lung-fish (Ceratodus

_ Sorsteri) i in the Collection of the Zoological Society of London. By Basurorp Dzay,

(Text-figs. B84 GBs) sonia s ow Sesiagaemraian fae Ste cits Weal fai’ s'alg Were clnrela ietdeattae eat) OOM

The Circulatory System of the Common Grass-Snake (Tropidonotus natriz). By

_ Cuas. H. O’Donocuvs, B.Sc., F.Z.S., Assistant to the Jodrell Professor of Zoology,

_ University College, London. (Pls. LXX.-LXXIT. and Text-figs, 86-91.) ......... » 612

s A First Aecount of the Courtship of the Redshank (Totanus calidris), By Juxtan S.

Be Hoxuey, Lecturer of Balliol College, Oxford ...1....escececeeeces ttececeeesess GAT

3 Some Brackish-water Amphipoda from the mouths of the Weser and the Elbe, and from

the Baltic. By E. W. Sexton, Marine Biological Laboratory, Plymouth. (Pls. LXXIII.

Me URRIY)) (5.5.5, eis cseeeses eat siccsvue oes Bea eibie leis erties Gattial ste eas 656

‘Descriptions of new Fishes of the Family Loricariide in the British Museum

_ Collection. By O. Tara Rucay, M.A.,F.Z.8. (Pls. LXXV.-LXXVIL) ...... péees 666

. i Sree see seoes Ce ee ee ee 672

ntributions to the Anatomy and Systematic Arrangement of the Cestoidea—V. On

a new Genus (Dasywrotenia) from the Tasmanian Devil (Dasywrus ursinus), the type

of a new Family. By Frank E. Bepparp, M.A., D.Sc., F.R.S., F.Z.S., Prosector to

ey: _ (Text-figs. 92-101.) vias bine oor es Sassen ens eles v'alalen as Seaway OLS

| dies in the Fossorial Wasps of the Family Scoliide, Subfamilies Elidine and

thoboscine, By Rowxanp E, Turnur, F.Z.S., F.E.S. (Pls. LXXXI.-LXXXIII.) 696

tes on the Spanish Ibex, By? Aap Ginn ASE, 25 ues abs tc or doen chase TOA

- LXVIII. Pyralidx from Dutch New Guinea! co's eiuein gs oo 2b aha eatin cae 546

“192, Pane Tt. a p- rma

_ Page

r. | Aleyonaria from Singapore . ee hi ar la Ms ch hoa Spi ‘ 505

LXIIt. ) pa ‘ \ Ba

eee ee eee tere es eees ee eetees

XVI | ? ALS 7 es a

UXVal.) : Bae pee Tas

LXIX. oun Coreg) ee ee 57

LXxXx. ; a ak ot

LXXI. } Circulatory System of the (Grsaeceoke Bn ere, 612

LEKI : : mares. Ge

-LXXIml.

| Amphipoda from Bremerhaven . titlate saa Vslciriaelye = ea 656

LXXIv. ea ae te

LXXV. 1. Chetostomus lepturus. = C. paucispinis, aire palmeri.\ oe ae

LXXVIL 1. Xenocara multispinis. 2. X. heterorhynchus, — 8, Pleco- | ‘ se s

: ; jena tac ema te AT

.. LXXVIL 1. Oxy yloricavia tamane. 2. O. leightont. 3. Otocinclus | A é

— Mmaculipinnis ....ereeveerree ceesnsernsneaetsen) ee

SUXXVIL. Leben porcini: See ee ee eee ee ee ee =) cat

Leas . 1. Barbus. argyrotenia. 2, ge minus. — i

PECHDAD vee eee crete ecsstetenest cscs 0

'UXxx, | | Amphilius oxyrhinus SarG © pre aiwin e's Sip ferniey ts retetenet ei

LXXXI. _Fossorial Wasps - Stare iopiecia nitanal slate aie csteletetelftatargis cre

~LXXXIT. Wing Neuration of Fossorial Wasps

oe ee OY SPOS

Ete) that i Ae eee ie now P.

is as follows :— ;

Part yt: issued i in tee

a pelle site Ae ;

rie Sit. ve

"December,

Ive

4a be te ee eke vega ng als Sait

PROCEEDINGS

Aap e

ius ee a

DECEMBER

Pent PRINTED FOR THE SOCIETY,

- SOLD AT THEIR HOUSE IN REGENT'S PARK.

MEL AO x LONDONE han :

“MESSRS. LONGMANS,: GREEN, “AND as

PATERNOSTER ROW, —

LIST OF CONTENTS.

1912, Parr IV. (pp. 757-913).

EXHIBITIONS AND NOTICES.

The Secretary. Report on the Additions to the Society’s Menagerie during the month

of April 1912 eee ee ey @ eer etoescescse ee ® Cer aves Ce ee eC ie ere ey

Mr. A. Buaynzy Prrcrvat, F.Z.S8. Exhibition of piatperbes and lantern-slides of Game

Animals from British East Africa ...... cece ccc c eee ee seen e tee ce te ce ee seaene

Mr. D. Seru-Sarrn, F.Z.8. Exhibition of two living specimens of the Lory, Calliptilus

solitarius, from Biji .. 6... ce cece ee cece e eee e cee cette eee ee tees at etencenees

Mr, G. A. Boutencer, F.R.S.,F.Z.S. Notice of a paper entitled “ Second Contribution to

our Knowledge of the Varieties of the Wall-Lizard” .........-. weet ee steeee

Mr. E. G. Bovtencer, F.Z8. Exhibition of a cocoon of the African Lung-fish

(Protopterus GNNECteNs) ...2++sseeceeeees misis(aleteehetelore © cain tela sles cere sen ie oie \eeere ee

Mr. E. G. B. Meapr-WaAtpo and otuErs. Discussion on the Preservation of the Native

Fivaiitaes of: CLmeatt Epeira Ter hore eas Se Rane Sri octane eee ae herent ere a

The SEcrETaRy. Report on the Additions to the Society’s gi paeerie during the months

of May to September 1912 ..0e.. cece cece eee cece cette teen teen teen cette eee ;

Mr. R. Lypexxer, F.B.S., F.Z.8. Note stating that “ Gazella hayi” = Gazella fuscifrons .

The Rev. T. R. RB. Strssine, M.A,, F.R.S., F.Z.S. Notice of a memoir on the Crustacea

Isopoda collected by the ‘Porcupine’ Expedition in 1869-1870 ........-.+++-..+ .

Sir Epuunp G. Lopzr, Bt., V.P.Z.S. Demonstration of the capacity of the electric lantern

presented by him to the Society ...... Searcy cece tec teens Terese ie BRIS ANC

Mrs. Rost Hata Tnomas, F.Z.S. _ Exhibition of the Eggs of Phasianus formosanus,

P. versicolor, and their F, 1 and F. 2 onepElue: (Text-fig. 126.) .

PAPERS.

41. The Local Races of Burchell’s Zebra. By Major J. Srrvenson-Hamitton, C.M.ZS.,

Game Warden of the Transvaal. (Text-figs. 102-106.) ........-.+00 cada cueereiate

42. On Dipteropeltis, a New Genus of the Crustacean Order Branchiura. By W. T.

Caran, D.8e., FBR. (Pl. UERXIV;) | ccacaues eae Ree bt Seinen ary

43, On two new Larval Trematodes from the Striped Snake (Tropidonotus ordinatus

sirtalis). By Wit1am Nicorn, M.A., D.Se.. M.D., F.ZS8., Lister Institute of

Preventive Medicine, London, (Text- fig. 107.) . SQOTLMOONH Coco DO Oa aa dome cIC

913

912

767

Contents continued on page 3 of Wrapper.

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MEETINGS

OF THE

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FOR

SCIENTIFIC BUSINESS.

1913.

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sp RC Ped INE cso a eal tae ee 3

x OCTOBER: AAwet gute: 28

be ae NGVEMB ERAS tee Uh ares)

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i a

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December, 1912. =

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se tian

ZOOLOGICAL SOCIETY OF LONDON.

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b> @

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tq)

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P. CHALMERS MITCHELL,

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December, 1912.

ON BURCHELL’S ZEBRA. 757

41. The Local Races of Burchell’s Zebra. By Major J.

STEVENSON-Hamiuton, U.M.Z.S., Game Warden of the

Transvaal.

[Received April 1, 1912: Read May 21, 1912.]

(Text-figures 102-106.)

INDEX.

Page

WaiiWbiOm:.))) 5 eee ny amr terre Oe TRI

Burchell’s Zebra (Lquus burchelli) south of the Zambesi is

usually divided by naturalists into several local races—wahlbergi,

transvaalensis, chapmannt, selousi. These derive their subspecific

distinctions from certain assumed differences in their leg and body

striping as observed in museum specimens. Thus :—

“In wahlbergi the body stripes meet the ventral stripe

inferiorly, while the legs are more or less fully striped ; the shadow

stripes on the hind quarters are strongly developed and not much

narrower than the main stripes, which are narrower than the

intervening spaces, while the fetlocks and pasterns are devoid

of stripes and spots. A female zebra from the Transvaal

differing from the typical wahlbergi by the extension of the

shadow stripes to the neck, has been named JZ. 6. transvaalensis.

In £. 6. chapmanii the shadow stripes have become faint and

narrow, the legs are marked to the hoofs, but the stripes on the

lower portions tend to break up into spots, and the inferior part

of the pasterns 1s not wholly black. This race inhabits the

country between Damaraland and Matabeleland. The last repre-

sentative of the species with distinct shadow stripes is the

Mashonaland bonte quagga (/. 6. selousi), which differs from the

last in that the striping of the legs is complete right down to

the hoofs, the pasterns being striped on both sides, and their

lower portion, owing to the fusion of several stripes, wholly

black. ‘The sides of the tail are also striped.” (Lydekker.)

Long observation of the herds of Burchell’s Zebra, running

in the Transvaal Game Reserves, has convinced me that all or

nearly all the above distinctions are found among them, frequently

in the same herd. I have often discussed the matter with

Dr. Gunning, of the Transvaal Museum, and have with him

examined skins and live specimens obtained from the Western

and Northern Transvaal. There is no doubt that any of these

might, from their markings, have come from the Eastern part of

the Province. Among the herds found in the latter occurs a

very great variety of striping. Some animals show heavy, wide,

and deeply tinted shadow stripes, while others display only the

slightest indications of them. Some are strongly ringed down

to the fetlocks, while others have no signs at all of any markings

Proc. Zoou, Soc.—1912, No. LI. 51

758 MAJOR J. STEVENSON-HAMILTON ON

below the knees and hocks. In some cases the body stripes are

continued right round the barrel to the ventral stripe, in others

Text-fig. 102.

Burchell’s Zebra, ¢, six years old, from south of the Sabi River,

Eastern Transvaal.

they stop far short of it. The striping is usually carried right

round the buttocks, but in I should say 20 per cent. at least of the

BURCHELL’S ZEBRA. 759

animals in each herd this is not the case. In fact there appears

to be no special distinctive marking common to any locality or

Text-fig. 103.

Burchell’s Zebra, g, ten years old, from south of the Sabi River,

Eastern Transyaal.

51*

760 MAJOR J, STEVENSON-HAMILTON ON

even herd. The shadow stripes on the neck, said to be dis-

tinetive of ZH. b. transvaalensis, have been occasionally, but very

rarely, noticed in individuals, and I cannot conceive their presence

to be more than an individual eccentricity, occasionally perhaps.

transmitted to offspring, but by no means constant.

Text-fig. 104.

Burchell’s Zebra, 2, eight years old, from south of the Sabi River,

Eastern Transvaal.

During the year 1911 [I was asked to obtain a couple of skins

for the Transvaal Museum, and selected two staliions from the

sane herd at a point a few miles south of the Sabi River (roughly

25 degrees 8. lat. by 31 HE. long.). I chose them merely as being

typical of the darker and lighter forms of striping respectively,

neither of them being in any way exceptional. Some animals.

present displayed deeper striping than the specimen shown in

text-fig. 102, while others showed more white than that in text-

fig. 103. These figures are from photographs of the skins. Text-

BURCHELL’S ZEBRA. 761

fig. 102 is that of a stallion six years old, and text-fig. 103 is of

one about ten years of age. A little later I obtained the skins of

a mare and her foal killed at a spot less than ten miles from

where I had shot the two stallions. The photographs of these are

reproduced as text-figs. 104 and 105,

Text-fig. 105.

Burchell’s Zebra, ¢, foal, trom south of the Sabi River, Eastern Transvaal.

In the first stallion (text-fig. 102) both the ordinary and the

shadow stripings are exceptionally strong, and the former is con-

tinued down and nearly all round the legs to the pasterns, which

are quite black. The body stripes meet the ventral line through-

out. The dorsal stripe is wide and very strongly marked. The

neck stripes are exceptionally deep, and set very close together.

In the second stallion (text-fig. 103) the shadow striping is not

nearly so well defined and the black stripes also are narrower

The body stripes do not extend all round the barrel, and there is

nractically no connection between them and the not very distinct

762 ON BURCHELL’S ZEBRA.

ventral stripe. The dorsal stripe is a very narrow black line, and

there is a distinct gridiron pattern on the back. The legs are ©

very slightly striped below the knees and hocks, and there are only

a few black hairs on the pasterns.

In the case of the mare (text-fig. 104), while tinh is a well-

marked ventral and a wide dorsal stripe, and complete union

between the former and the barrel stripes, there is an entire

absence of any black markings from all the pasterns, and from

the fore legs below the knees.

Text-fig. 106.

Burchell’s Zebra, ¢ ; the same animal as that shown in text-fig. 103,

photographed just after death.

Tn the foal again (text-fig. 105), while the legs all bear faint

indications only of stripes, the pasterns are all quite black, and

there is some indication of a gridiron pattern on the back.

In none of the specimens is there the smallest indication of

any shadow striping on the neck. It will be noticed also that in

the younger stallion and the mare (text-figs. 102 & 104) the

frontal stripes on the face run straight from forehead to nose,

while in the older stallion (text-fig. 103) (and to some extent in

the foal (text-fig. 105), though it does not show up well in the

figure) the frontal stripes tend to form concentric rings.

LXXXIV.

Po Ze Sele 2 Pip

Bale & Danielsson, L*? sp.

ID G2 IMAI ONS IL WIS) IeiUENOINIO) ©) .

del.

’

G.M. Woodward

ON A NEW CRUSTACEAN GENUS. 763

Jt would be interesting to know whether this absolute want of

uniformity found among Burchell’s Zebras in the Transvaal is

peculiar to this country, or whether it is the case throughout

Africa south of the Zambesi. Obviously it would be very

dangerous to attempt any classification from isolated museum

specimens obtained from the Transvaal at any rate.

Dr. Gunning proposes, I understand, to deal more fully in

the forthcoming ‘ Annals’ of the Transvaal Museum with a matter

which I here only lightly touch upon.

42. On Dipteropeltis *, a New Genus of the Crustacean

Order Branchiura. By W. T. Caumay, D.Sc., F.Z.S.f

[Received April 13, 1912: Read May 21, 1912.]

(Plate LX XXIV.f)

INDEX.

Page

Structure or Morphology .....0.....0.0...cecereeee 768

Hiholosva(arasitism)) -ese.c-ere see hens aes Feehan AOS

Geographical 200 ona ta cceaunitnnt cece Melee a GS

Systematic: Dipteropeltis hirundo ..................... 763

The specimens described in this paper were presented to the

British Museum twenty years ago by Spencer Moore, Esq., by

whom they were collected in Southern Brazil. They were found

among a number of specimens of Dolops longicauda Heller bearing

the label “ Parasite on the gills and body of the fish known

all up the river as ‘Dorado’ from its golden colour. From

Corumba and neighbourhood, Matto Grosso.”

DIPTEROPELTIS HIRUNDO§. (Pl. LXXXIV. figs. 1, 2.)

Calman, Abstract P. Z. 8. 1912, p. 34 (May 28th).

Description of female :—The most striking feature of the species

is the form of the carapace, the lateral lobes of which are drawn

out into narrow lanceolate wings directed backwards and ex-

tending far beyond the tips of the long abdominal lobes. The

wings are fleshy in texture, traversed by numerous branching

blood-channels, but without spines or other armature on the

under surface.

The head is small, defined on each side from the carapace wings

* The complete account of the new form described in this paper appears here,

but since the name and a preliminary diagnosis were published in the ‘Abstract,’ it

is distinguished by bemg underlined —Hprror.

+ Published by permission of the Trustees of the British Museum.

{ For explanation of the Plate see p. 766.

§ For definition of the genus see p. 766.

764 DR. W. T. CALMAN ON A.

by a shallow antero-lateral sinus from which two converging

grooves run in on the dorsal surface. These grooves unite into

one on each side, but the resulting groove dies out before reaching

the middle line. <A pair of inter-ocular chitinous rods are well-

marked, closely approximated in front, diverging posteriorly, and

united by an indistinct articulation with a nearly parallel posterior

pair. The paired eyes are very small and close together near the

front margin. The unpaired eye could not be detected.

The free thoracic region is not distinctly segmented ; it is of

equal width throughout and more than three times as long as

wide.

The abdominal lobes are very long, narrowly lanceolate, and

cleft nearly to the base. No trace of furcal rami can be found.

The antennules and antenne (fig. 3) are very minute and nearly

hidden from view under the in-turned front margin of the head.

The antennules, in particular, are very easily overlooked; they

are not divided into segments but have a blunt lobe at the base

carrying a short apical tooth. The antenne are about the same

length, and consist of a stout basal part and a subglobular terminal

part separated from it by a constriction.

The mouth-parts form a prominent cone (fig. 4). ‘The two large

and well-separated lobes of the upper lipare unarmed. The lower

lip bears a pair of slender conical papille (“maxille ” of Claus)

which project slightly from the opening of the mouth. The

mandibles are crescentic in form, with the concave edge very

finely serrated and the convex edge bearing a few teeth near the

tip.

In front of the oral cone and continuous with it at the base is

a papilla, directed forward, having at its tip the opening of a duct.

This, no doubt, represents the sheath of the preoral “sting” of

Argulus, but no trace of the spine or sting itself could be detected

in the dissection of two specimens. At the base of this papilla

and of the mouth-cone are groups of very large cells, presumably

glandular, with deeply-staining nuclei.

The suckers (first maxille, second maxille, or first maxillipeds

of various authors) are placed close together in front of the

mouth-cone, generally concealing altogether the preoral papilla.

In place of the usual radial supports, the whole of the membranous

border of the sucker is covered with discoidal scales (fig. 5).

The maxillipeds (fig. 6) (maxille of some authors) are very short,

very stout at the base, and tapering rapidly to the tip. They are

each composed of five segments, of which the second and third

have numerous pectinate or branched spines on the ventral and

anterior surfaces. The terminal segment bears two minute claws

and a process which lies alongside of them.

The successive pairs of legs (figs. 7-10) are set wide apart from

each other along the sides of the thoracic region. The rami are

in all cases shorter than the protopodite and carry only a few

short sete ; there is no flagellum on any of the legs. In the first

NEW CRUSTACEAN GENUS. 765

pair the endopod may be a good deal shorter than the exopod, or,

as in the specimen figured, nearly equal to it; both rami are un-

segmented. In the second pair also the rami are unsegmented,

and the endopod is from about one-half to two-thirds as long as

the exopod. In the third and fourth pairs the rami are subequal

and the endopod is divided into two subequal segments. The

basal lobe of the protopodite in the fourth pair is tongue-shaped,

with a slight protuberance at the base of its distal edge.

Dimensions of Holotype in millimetres.

Length of body to tip of abdominal lobes............ 20-0

Total length to tip of carapace wings ............... 26-0

Br ecoree Nordea 4 c.eueuelel S, sean oceans 2°5

ihength of thoracic resiomy Jysc.)-Lisechh heubose.aeelces 8:0

Bueadthlvof dhoracie resiony):0). wrk Acyl glares 2°09

iBucagihy ofewangearbibaser,.|..chj:..kardeon ed. 21... -Amee 1-8

Greatest breadth of wing, at about 8mm. from base 4:8

dienethvofiabdominalJobessyes, ise hhise: ee beeen. 6°8

Greatest breadth of abdominal lobes.................. 13

Length of antennules and antenne, about ......... 0-13

diene phyottonral comes jo scseacius aids: Os Bhan beds us. 0-5

Greatest diameter of sucker ..............0..cc.eeeee es de

enothomsecomdvles 5. i. sik ue. es eee Re:

Miametery/ Gt eves: aWOUb sted sauces we Matedeees oee =< 0-1

Wistance apart oltemes abouts. 6 la Aenea e ek 0-45

Locality.—Corumba, on the Paraguay River, Matto Grosso,

Brazil. Four female specimens taken (in company with Dolops

longicauda) on the fish known as ‘“‘ Dorado” (probably a species

of Salminus).

Holotype.—Female, No, 92.10.24.2 in British Museum Register

of Crustacea.

A ffinities.—In having the so-called maxille or first maxillipeds

modified into suckers this species differs from the genus Dolops,

and in possessing antennules and a preoral papilla it differs from

Chonopeltis. From all the species hitherto referred to Argulus it

differs in (1) the remarkable form of the lateral wings of the

carapace ; (2) the length of the abdominal lobes and the absence

of fureal rami, in which characters it resembles some species of

Dolops ; (3) the entire absence of conspicuous spines or hooks on

the under side of the body and appendages, in which it resembles

Chonopeltis ; (4) the vestigial condition of antennules and

antenne ; (5) the absence of a spine or sting on the preoral

papilla; (6) the absence of the usual radial supports on the disc

of the suckers. Some of these characters, especially Nos. 1, 3,

and 6, are possibly not of great systematic importance, but

together they seem to show that the new form is less closely

related to any of the species included in the genus Argulus than

766 ON A NEW CRUSTACEAN GENUS.

these are to one another. I have proposed, therefore, to establish

for it a new genus, which may be defined as follows :—

DiIPTEROPELTIS.

Calman, Abstract P. Z.8. 1912, p. 34 (May 28th).

Argulide with the so-called maxille modified as suckers ; with

the preoral papilla present, but without a spine ; with antennules

and antenne very minute, imperfectly segmented ; without large

spines or hooks on under side of carapace, body, or appendages :

without furcal rami on the abdominal lobes; with the lateral

wings of the carapace greatly elongated.

Genotype.—D. lirundo Calman.

It is to be noted that the single species of Chonopeltis,

(. inermis Thiele *, is known only from a single specimen (from

Lake Nyasa). It is not at all impossible, therefore, that the

antennules, if they are as small as in Dipteropeltis, may have

escaped observation, and it is even possible, though less probable,

that an unarmed preoral papilla may be present. Should this

prove to be the case it is doubtful whether, in spite of the great

difference in the form of the carapace, the separation of Diptero-

peltis from Chonopeltis could be maintained.

EXPLANATION OF PLATE LXXXIV.

(The magnifications given are only approximate.)

Vig. 1. Dipteropeltis hirundo. Female (holotype), from the dorsal side. x 3.

2. Anterior part of body of same specimen from ventral side. The suckers

have been drawn apart and slightly backwards. In the preserved

specimens they meet in the middle line and entirely conceal the preoral

papilla. x 6. .

3. Antennule and antenna. x 450.

4. Tip of oral cone from antero-ventral aspect. x 100.

5. Part of membranous margin of one of the suckers. x 160.

6. Maxilliped. x 20.

7. Leg of first pair. x 20.

8. Leg of second pair. x 20.

9. Leg of third pair. x 20.

10. Leg of fourth pair. x 20.

Figures 3-10 are drawn from one of the paratypes.

* Thiele, Zool. Anz. xxiii. p. 47 (1900); Mitth. Zool. Mus. Berlin, i. p. 44.

figs. 110-116 (1904).

ON NEW LARVAL TREMATODES. 767

43, On two new Larval Trematodes from the Striped Snake

( Tropidonotus ordinatus sirtalis). By Wi1LL1am NIcoLL,

D.Sc., F.Z.8., Lister Institute of Preventive Medicine,

London.

[Received April 9, 1912: Read May 21, 1912. |

(Text-figure 107.)

INDEX.

Page

Ethology: Two larval Trematodes in the mesenteric fat of

Striped Snake (Tropidonotus ordinatus sirtal is). Adult

form probably in a bird ..........-. 6.0.0. ee ee 767

Geographical Zoology: North America ; Striped Snake

infected with two larval Trematodes ...... 767

Systematic: Cercaria ordinata, sp. ., and Diplostomum

sirtale, sp.n., from cysts in themesentery of the Striped

The occurrence of encysted larval parasites in snakes is evidence,

if such were wanting, that some snakes are eaten by other

animals. What is more important, the character of the parasites

may indicate what variety of animal is in the habit of eating the

snake in question. Conversely, the presence of any particular

species of adult parasite in an animal is almost always a sure

proof that such animal eats the snake in which the larval stage

is found. It is unfortunately in many cases a matter of difticulty

to diagnose the systematic characters of a larval parasite. Ina

number of cases, however, it is possible to assign it to a definite

genus, rarely to a particular species.

The two cases to be dealt with here present a certain amount

of difficulty. The first larva is evidently a Distomate Trematode,

but beyond that it is impossible to go; the second larva is just

as obviously a Holostomid, and almost certainly belongs to the

genus Hemistomum. Such a diagnosis does not lead very far,

but it at least enables one to say that in all probability the

Striped Snake (Zropidonotus ordinatus, var. sirtalis) is eaten by

some bird, for adult Holostomata are known to oceur only in

birds. This will possibly be confirmed by direct observation.

Both forms were met with together in each of three Striped

Snakes from North America, which died in the Society’s Gardens

on the bth and 20th December, 1910, and on 10th March, 1911.

They occurred in enormous numbers in the mesenteric fat along

the whole length of the intestine. Each was enclosed in a small

spherical or ovoid cyst with unusually thin and soft walls.

Unlike what is generally found in a Trematode cyst, the wall

gave the impression of being a thin membrane instead of the

more usual tough chitinous investment. On this account, not

only could the larve be extracted from the cysts without dif-

ficulty, but when placed in water they escaped readily of their

own accord. When a piece of the cyst-infested mesentery was.

768 DR, WILLIAM NICOLL ON

suspended in water, a continuous shower of larve was observed to

fall to the bottom of the vessel.

The first form, which I name CERCARIA ORDINATA, sp. n. (text-

fig. 107 a), was much more numerous than the other. It isa

typical tailless encysted cercaria, about *5 mm. (‘455 mm.) in

length and -2-"25 mm. in greatest breadth. In shape it is ovoid

and flattened dorso-ventrally. The entire surface of the body 1s

covered with minute regularly-arranged spines. The oral sucker

is almost terminal and has a diameter of ‘07 mm. (‘06-08 mm.).

Text-fig. 107.

See ee oe

ea ee

wes

Sse

A. Cercaria ordinata, sp.n. Ventral view, x 150.

B. Diplostomum sirtale, sp.n. Ventral view, X 150.

The length is usually slightly greater than the breadth, and the

sucker has a somewhat characteristic funnel-shaped appearance.

The globular ventral sucker is situated rather-in front of the

centre of the body, and has a diameter of -083 mm. (‘075—-097mm.).

Its distance from the anterior end of the body is on an average

-22 mm. (17-25 mm.). On the dorsal lip of the oral sucker are

the two symmetrical apertures of the cystogenous ducts. The

cystogenous glands are conspicuous structures, and consist of four

NEW LARVAL TREMATODES. 769

large cells situated mm a transverse row immediately in front of

the ventral sucker. There are two pairs,a right and a left. The

ducts from each pair unite almost at once, and the united ducts

then pass forward in an irregular course; but just before they

reach the oral sucker each makes a characteristic twist, following

which there is a gradual increase in calibre until near the term1-

nation, when they contract slightly again. The cystogenous

glands have an irregular rectangular outline, and measure

"048 x:037 mm. In direct contact with the oral sucker is the

small muscular pharynx measuring ‘024 mm., which is continued

by a short, somewhat dilated cesophagus of the same length as

the pharynx. The intestinal bifurcation is about midway between

the two suckers. ‘The diverticula are simple, somewhat wide

tubes, which terminate not far behind the ventral sucker (i. e. a

third of the distance from the sucker to the posterior end of the

body). The excretory vesicle is V-shaped. A common trunk is.

practically absent, and the limbs extend forward to near the

terminations of the intestinal diverticula. The excretory tubules

are very fine; the main tube on each side extends forward to

near the oral sucker, where it turns back. The excretory aperture

appears to be slightly dorsal.

No trace of other organs could be made out, so that no accurate

idea can be obtained of the systematic position of this larva.

The peculiar configuration of the excretory vesicle may, in con-

junction with the shape of the alimentary canal, eventually lead

to the identification of its adult form, but at present, so far as I

am aware, there is no known adult Distome to which this larva

can be ascribed.

The second form, which I name DipLosromum siRTALe, sp. n.

(text-fig. 107 B), differs markedly from the first. It is about the

same size, and occurs in somewhat similar cysts, but its shape and

colour are entirely different. The body of Cercaria ordinata is

light and transparent, whereas that of Diplostomum sirtale is

dark and almost opaque. The opacity is due to the presence of

innumerable small granules distributed throughout the whole

body. The shape is that of a typical Diplostomum larva, being

scoop-like with a short handle. The shape is due to the rolling

over of the postero-lateral margins of the body. In life, however,

these margins are capable of more or less eversion, so that on

occasion the body may appear almost flat. The dimensions of

this larva are "48-55 x -28-"32 mm., the short stumpy tail being

-06 mm. long.

The oral sucker measures ‘(039 mm. in diameter; the ventral

sucker ‘042-045 mm. The latter is situated a little in front

of the middle of the body, -25--28 mm. from the anterior end.

Midway between it and the posterior end of the body occurs

the characteristic Holostomid fixing disc, which appears as

a transparent disc about the same size as the ventral sucker.

Of the internal organs only the alimentary canal and excretory

vesicle were visible. The former comprises a pharynx contiguous

770 SIR CHARLES ELIOT ON A RARE NUDIBRANCH.

with the oral sucker and measuring °024x°020 mm. This is

followed by an cesophagus about twice as long as the pharynx.

The intestinal bifurcation takes place rather nearer the ventral

sucker than the oral sucker, and the simple diverticula extend

a little beyond the posterior border of the fixing disc. The

excretory vesicle consists of a wedge-shaped sac, which extends

forward as far as the ends of the intestinal diverticula.

This is in all probability the larval stage of some species of

Hemistomum, parasitic in a bird.

44. A Note on the rare British Nudibranch Hancockia

eudactylota Gosse. By Sir Caartes Extor, K.C.M.G.,

CB. I Z.8.

[Received April 18, 1912: Read May 21, 1912. ]

(Plate LXXXY.)

See Gosse, On Huncockia eudactylota, Ann. Mag. Nat. Hist. ser.

4, xx. 1877, pp. 316-318; Gamble, On two rare British Nudi-

branchs, Lomanotus genet and Hancockia eudactylota, ib. ser. 6, ix.

1892, pp. 378-385; Trinchese, Ricerche anatom. sul genere

Govia ( = Hancockia), Mem. della R. Accad. delle Sci. dell’

Istituto di Bologna, ser. 5, vii. pp. 183-191, 1886; Eliot, Sup-

plement to Alder and Hancock’s Monograph of the British

Nudibranchiate Mollusca, Ray Society, 1910, pp. 17, 72,

118-120, 163.

No coloured figure of this rare British Nudibranch has yet

been published so far as I am aware. | endeavoured to include

one in my Supplement to Alder & Hancock’s Monograph, but

no specimen of the animal could be obtained before the time fixed

for publication. Shortly afterwards a single individual was

captured at Plymouth and drawn by a local artist under the

supervision of the naturalists who were working in the Laboratory

of the Marine Biological Association. I have not seen the animal

alive, but these drawings agree with what is known of its struc-

ture, and I have no doubt that they faithfully reproduce its

appearance.

For an account of the genus and species see the references to

my Supplement given above.

EXPLANATION OF PLATE LXXXV.

Hancockia eudactylota.

Fig. 1. Dorsal view of animal.

2. Ventral view of animal.

3. Side view of head.

4, A rhinophore.

5. .

at Different views of lateral processes.

Pansy MONA, IL IL MOK NOW.

London Stereoscopic Co. imp.

HANCOCKIA EUDACTYLOTA.

eZeohy WO), IPAS WOO.

MANS

London Stereoscopic Co. imp.

GIRAFFA CAMELOPARDALIS THORNICROPFT1.

ON THE NORTH RHODESIAN GIRAFFE. Viral

45, The North Rhodesian Giraffe.

By R. LyprxxKer, F.R.S., F.Z.8.*

[Received April 25, 1912: Read June 4, 1912.]

(Plate LXX.XVI. +)

InDEX,

Page

uhodesvan GuraiterOb ys qasd essere on con aren alee en erage

Giraffa camelopardalis thornicrofti, Description of ... 771

On the 20th of October, 1910, the British Museum (Nat. Hist.)

received from Mr. H. Thornicroft, Native Commissioner, Petauke,

North-east Rhodesia, the skin, skull, and limb-bones of an adult

male Giraffe shot by himself in that district. Mr. Thornicroft

had previously called on me in London, and expressed his willing-

ness to shoot and present to the Museum a Giraffe from the

single herd in this part of Rhodesia, if the necessary permit could

be obtained. This was in due course procured, and was followed,

after an interval, by the arrival of the above-mentioned skin and

bones.

The skin was soon afterwads set up by Rowland Ward Ltd.,

and the mounted specimen placed in the big case at the head of

the staircase leading to the Hast Corridor of the Museum, along-

side the male and female of the Hast African Giragfa camelo-

pardalis rothschildi. As it is mounted with the neck bent, it is

difficult to ascertain the exact height, but I estimate this at close

on 18 feet, or possibly rather more.

When the specimen was installed in its case, it became essential

that it should receive a distinctive name; and I accordingly

communicated the following preliminary note to ‘ Nature’ +:—

“This Giraffe is characterised by the low and conical frontal

horn, the grey colour and scattered spotting of the sides of the

face, the chestnut-brown forehead, deepening into black on the

tips of the horns, the absence of a distinctly stellate pattern in

the neck and body spots, which are light brown on a yellowish

fawn ground, and the uniformly tawny colour of the lower portion

of the limbs. This Giraffe, which I propose to call Giraffa

camelopardalis thornicrofti, appears to be related to the Kiliman-

jaro G. c. tippelskirchi, but differs by the more compact frontal

horn, the brown, in place of grey, forehead, and the uniformly

fawn lower part of the legs, the latter being whitish in adult

bulls [of tippelskirchi|, but fawn and spotted in cows and young

bulls.”

The last statement rests on the authority of Messrs. M. de

* Published by permission of the Trustees of the British Museum.

+ For explanation of the Plate see p. 773.

ft Vol. Ixxxvii. p. 484 (1911).

172 ON THE NORTH RHODESIAN GIRAFFE.

Rothschild and H. Neuville*, who state that in the East African

Giraffe which they describe as rothschildi, but which—despite the

locality whence it is stated to come—is certainly tippelskirchi,

these age and sex differences are observable. I have, however,

doubts whether they hold good in all cases; and it is still possible,

in spite of what I have previously written, that there may be one

form (schillingsi) in which the shanks of adult bulls are white and

another (tippelskirchi) in which they are fawn and spotted T, and

further, that these two types may intergrade.

That the nearest relative of the Rhodesian Giraffe is G’. c. tip-

pelskirchi, may be considered certain. Of the latter I have had

tor comparison the mounted head and neck of an adult male, a

mounted immature female, and the mounted head and neck of

a calf, as well as a coloured plate in Messrs. de Rothschild and

Neuville’s memoir f.

Elaborating to a certain extent the foregoing brief diagnosis,

attention may be directed to the fact that tippelskirchi and

thornicrofti agree (and thereby differ markedly from rothschildi)

in having the triangular space between the eye and the nostril

devoid of spots. In the adult male of tippelskirchi, however, the

eround-colour of the whole head is dirty greyish white, whereas

in thornicrofti the forehead is chestnut or umber-brown, deepening

into black at the tips of the horns, which are grey in the

Kilimanjaro race.

In the Rhodesian Giraffe the spots on the region behind the

eye and the side of the lower jaw are very faintly marked, and

blackish grey in colour; whereas in the Kilimanjaro bull they

are larger, more numerous, and chocolate-brown in colour, being

deeper in tint than the neck-spots (this feature being also shown

in the immature female and the calf).

Tn thornicrofti the spots on the neck are burnt-umber in colour

and markedly elongated in form, with their terminal ends jagged.

There are about eight of them in the longitudinal row which

starts immediately in advance of the point of the shoulder. In

tippelskirchi they are more numerous (ten or eleven in what

appears to be the corresponding row), less elongated, and much

more irregular in shape.

Compared with the young cow tippelskirchi, the spots on the

body of thornicrofti are less numerous, more especially on the

hind-quarters, while many of them are more deeply incised on one

side, although they are less jagged in general contour. The

spotting on the inner side of the thighs and of the upper part

of the fore-legs is also much less pronounced. In the original

description (which was drawn up when the specimen was in the

basement of the Museum) it is stated that the shanks of the

legs are uniformly fawn, but, as a matter of fact, they are

* Ann. Sci. Nat., Zool. ser. 9, vol. xili. pp. 124, 129 (1911).

+ See Proc. Zool. Soc. 1904, vol. i. p. 219.

{ Op. cit. pl. ii. fig. 1, lettered G. c. rothschildi.

ON ANTLER-GROWTH IN THE CERVIDA. AUS

rufous-fawn with very faint traces of spotting nearly down to

the fetlocks; while from the latter to the hoofs they are dirty

greyish white.

The foregoing evidence clearly establishes the right of the

North Rhodesian Giraffe to rank as a distinet local race; and if

it be true that the one herd is completely isolated, there is

probably no intergradation with the Kilimanjaro race.

EXPLANATION OF PLATE LXXXVI.

Adult bull of Giraffa camelopardatis thornicrofti.

46. On Antler-Growth in the Cervide, with special reference

to Hlaphurus and Odocoileus (Dorcelaphus). By

R. I. Pococg, F.R.S., F.L.S., F.Z.8., Superintendent

of the Gardens and Curator of Mammals.

[Received and Read June 4, 1912.]

(Text-figures 108-112.)

InDEX.

Page

Classifications of the Cervide 00.0... 778

Mode of growth of Antlers in typical Old World Deer ....., 775

Interpretation of the Antlers of Elaphurus davidianus ...... 777

Interpretation of the Antlers of Odocotleus sp. incert. ...... 780

Date of Antler-change in American and Old World Deer ... 783 (Note)

Introduction.

Most, if not all, the attempts that have hitherto been made to

understand the antlers of Deer and arrive at correct conclusions

regarding the homology of the tines have been based upon com-

parisons between the fully formed antlers of different species.

This, in my opinion, is the reason why there has been failure in

some cases to detect homologies which study of the growth of

individual antlers reveals.

The importance of this question depends upon the cireumstance

that twenty years ago Mr. Gordon Cameron * proposed a classi-

fication of the Cervidee, based upon the antlers, as a substitute

for the classification, founded upon the skeletal structure of the

fore feet, which Sir Victor Brooke had suggested 7.

To make clear the purpose of the present paper, it is necessary

to summarise briefly the rival classifications put forward by these

two authors. Sir Victor Brooke divided the Cervide into two

* “The Field,’ 1892, pp. 265, 703, 74.1, 860.

+ P.Z.S. 1878, pp. 883-928.

Proc. Zoou. Soc.—1912, No. LIT. 52

774 MR, R. I. POCOCK ON

sections. The first, which he called Telemetacarpi (Telemeta-

carpalia) because the distal ends of the lateral metacarpals

persist, comprises the Roe (Capreolus), the Chinese Water-Deer

(Hydropotes), the Reindeer (Kangifer), the Elk (Alce), and all the

exclusively American deer with the single exception of the typical

Wapiti (Cervus canadensis); the second, called Plesiometacarpi

(Plesiometacarpalia) because the proximal ends of the lateral

metacarpals are usually present, whereas their distal ends are

suppressed, comprises all the deer of the Old World, except:

the four genera mentioned above, but none of those of the

New World apart from the Wapiti. Amongst the Old World

forms the most important species for the moment figuring

amongst the Plesiometacarpalia is Pére David’s Chinese Deer

(Llaphurus davidianus).

Mr. Cameron’s classification was widely different. Dismissing

as unimportant the character relied upon by Brooke, he divided

the Cervide into three sections: one for the Reindeer with

antlers in both sexes, the second for the Elk with laterally

extended antlers, the third for the remaining species with antlers

restricted to the male and erect or suberect. This third section,

which alone concerns us now, was subdivided into two categories

of species, one comprising those in which the antlers consist, as

in the typical Old World deer and the Wapiti, of a “ brow-tine”

and a ‘“‘ beam,” to use Gordon Cameron’s terminology, and the

other those in which the antler has, as he thinks, no brow-tine

but consists of a “forked beam,” as in all typical American deer

(except the Wapiti) and in the Roe and Pére David’s Deer

amongst the Old-World species.

Now with regard to the affinities of the species composing

Cameron’s third division, there is only one point in which there

is complete divergence between him and Brooke. This concerns

Pere David’s Deer, a species classified by Brooke with the Red

Deer, Sambar, and other Hlaphine stags, and by Cameron with

the American forms allied to the Virginian and Mule Deer, the

correct name of which seems to be Odocoileus *.

So far as I can see, the only @ priori objection to be raised

against Mr. Cameron’s system, if we accept his premises, is that.

it is based upon a secondary sexual character. But although it.

cannot be justifiably consigned to oblivion on that account, it may

be doubted if it would ever have come into sufticient prominence

for serious discussion had it not been for the unqualified acceptance.

accorded it by Mr. Lydekker. However that may be, it is clear

that if Mr. Cameron’s assumption that there is a fundamental

difference in structure between the antlers of the groups of deer

mentioned above is wrong, his classification, based on that claim,

goes by the board.

In the following pages I shall endeavour to show that his

classification is untenable, because a study of the seasonal growth

* Dorcelaphus and Cariacus are better known but superseded terms.

ANTLER-GROWTH IN THE CERVID. (WES

of an antler of Pére David's deer and of an American deer, allied

to the Virginian, proves that the homologue of the brow-tine of

the Elaphine stage is present in both—a conclusion which is by

no sete evident from an examination of the fully-formed

antlers.

Antler-Growth in typical Old-World Deer.

In the Zoological Gardens I have repeatedly watched, year

after year, the growth of the antlers of deer belonging to the

Elaphine, Sikine, and Rusine types without finding any variation

Text-fig. 108.

Wry WE

Karly growth-stages of Antlers of some Old-World Deer.

A & B. Successive stages observed in Cervus hanglu. C. Cervus canadensis.

D. Rusa aristotelis.

a, the anterior branch or “ brow-tine” ; p, the posterior branch or “ beam” ;

6, the rudiment of the bez-tine arising from the posterior branch.

of moment in the method of their development. The antler

starts as an undivided bud. This bud then shows signs of

division into two buds, an anterior and a posterior. Thesetbuds

52*

776 MR. R. I. POCOCK ON

gradually, and with nearly equal rapidity, increase in length, the

anterior growing forwards and the posterior backwards. In the

Sambar (2sa, text-fig. 108, D) and some other species they show

a marked inclination upwards ; so that at one stage the antler may

be likened somewhat to a short-stalked Y, and at this or evenata

later stage in deer like the Sambar (/twsa) and others which have

no “ bez”-tine, the antler may be indifferently described as an

‘‘unbranched beam with a brow-tine” or as a “ forked beam” or

as a biramous antler. The anterior and posterior branches some-

times, as in Cervus eldi, grow at approximately the same speed

until the anterior has almost attained its limit; but usually the

growth of the posterior tine is from the first more rapid. However

that may be, the equivalence of the two branches in the early

stages is plain enough ; but afterwards this becomes less and less

evident as the posterior branch continues to lengthen and

develops its accessory tines.

Text-fig. 109.

Pp a is a 2

DIN yy ii

WW)

Five stages (4 to #) in the growth of an antler of Rucervus duvaucelli.

a, anterior branch or “ brow-tine” ; p, posterior branch or “ beam.”

These facts are shown in the annexed figure (text-fig. 109),

representing five stages in the growth of an antler of a specimen of

the Swamp Deer or Barasingha (Rucervus duvaucelli), These were

ANTLER-GROWTH IN THE CERVIDA, (id

sketched on May 13, 16, 22, June 6 and 12. Similar stages may

be observed in other typical deer of the Old World*. In the

Elaphine stags, however, which normally grow a ‘bez’’-tine,

the biramous stage is early complicated by the appearance of the

bud of this tine. Now this tine has been regarded as a dupli-

cation of the brow-tine; and in Max Weber’s? diagram showing

suggested homologies of the tines in certain deer the brow- and

bez-tines are tinted alike, suggesting his adoption of this view.

Nevertheless I believe it to be quite incorrect, for in all cases

where I have watched its origin, the bud of the “bez”-tine

arises, not from the brow-tine at all, but from the “beam.” It

is, in fact, the basal or proximal tine of the posterior branch of

the antler. This is illustrated in text-fig. 108, d—C, showing the

early stages of the growing antler of the Hangul (Cervus hanglu)

and of the Wapiti (Cervus canadensis).

Antler-Growth in Pere David's Deer (Elaphurus davidianus).

There is no stag whose systematic position has troubled

zoologists so much as Hlaphurus. On the one side are those,

like Dr. Gray, Mr. Cameron, and Mr. Lydekker, who, relying

upon the structure of the antlers of the adult, placed the genus

with the American deer. On the other side are those, like

Sir Victor Brooke, Flower, Max Weber, and others, who, adopting

the skeleton of the foot as a basis, classified it with the typical

Old-World species.

The antlers of this stag have often been figured and described, -

and a good idea of their form in the adult may be gathered from

text-fig. 110, C, and text-fig. 111, 7. They typically consist of a

comparatively long basal portion from which two branches arise :

one long, slender, simple or divided, projects backwards parallel,

or nearly so, with the animal’s back; the other stout, erect, or

curved slightly forwards, terminates in a pair of strong tines.

At first sight, these antlers appear to have no trace of a brow-

tine. This was evidently Sir Victor Brooke’s opinion, and it was |

adopted by Mr. Cameron and Mr, Lydekker, who, on the strength

of this belief, boldly claimed that this stag belonged to the same

group as the American deer, also held to have no brow-tine,

despite the resemblances in other respects pointed out by Brooke

between Hlaphurus and the typical Cervidee of the Old World.

Prof. Garrod was more cautious, and frankly gave up the attempt

to interpret the antlers of Hlaphwrus when he remarked that they

‘Care at present quite beyond my comprehension.”

This, then, was the state of the case when my researches on

the specialised cutaneous glands of Ruminants ¢ showed that the

* My. J. G. Millais (‘Mammals of Great Britain and Ireland,’ iii. plate facing

p. 140, 1906) has published a series of figures of antler-growth in the Fallow Deer

(Dama) illustrating precisely the same phenomenon.

+ Die Saug. p. 667, 1904.

t P. Z. S. 1910, p. 840.

778 MR. R. I, POCOCK ON

absence of interdigital glands on the feet and the smoothness of the

integument between the hoofs in Hlaphurus corroborated Sir Victor

Brooke’s views as to the relationship between this animal and such

Old World deer as Rusa, Rucervus, and Cervus, and weakened to

a corresponding degree the claim for affinity between it and the

Text-fig. 110.

Diagram of the Antlers of four genera of Cervide, to illustrate the homologies

established in this paper.

A. Cervus. B. Rusa. C. Elaphurus. VD. Odocoileus.

a, anterior and p, posterior branch. In 4 and B the anterior branch is called the

“brow-tine” and the posterior branch the “beam.” B is somewhat inter-

mediate between A and ©. They differ collectively from D in having the

anterior branch well developed. In D it is small and concealed behind the

highly developed posterior branch.

Telemetacarpal species, in all the members of which examined by

me, belonging to the genera Mazama, Odocoileus (Dorcelaphus),

Capreolus, Rangifer, and Alce, the skin between the hoofs is

ANTLER-GROWTH IN THE CERVID. 779

thickly hairy and, in all but Alce, a large pouch-like interdigital

gland is present at least in the hind foot.

Text-fig. 111.

Skat ANI

8 A

Nine stages (4 to I) in the growth of an Antler of Klaphurus davidianus, showing

that the branches marked @ and p correspond precisely in origin with the

brow-tine and the beam of other genera of Old World Deer. Compare D and #

with fig. 108, D.

(From sketches made at Woburn and kindly supplied by Lord Tavistock.)

780 MR. R. I. POCOCK ON

Thinking, for these reasons, that there must be some flaw in

the claim that the antlers of Hlaphurus differ fundamentally from

those of, say, Rusa, I suggested the following homologies :—In

Rusa the antlers have a short base, a short undivided anterior

branch or brow-tine, and a large divided posterior branch or

beam; in Hlaphurus they have a longer base, a very large divided

and more erect anterior branch or brow-tine, and a correspondingly

reduced, comparatively slender, divided or undivided posterior

branch or beam.

This, however, was a mere guess, which I was unable to

substantiate by any evidence of much value. Believing, how-

ever, that the growth of the antlers in Llaphurus would finally

settle the question one way or the other, I asked Lord Tavistock

if he would kindly observe the process for me on one of the stags.

at Woburn. This he was good enough to do, and sent me in

addition the series of sketches reproduced in text-figure 111.

These sketches show, in my opinion, that my guess was, as I

expected, correct. In the first three stages the antler is little

more than an excrescence dividing into an anterior and a posterior

bud. In the fourth stage the base is beginning to lengthen, the

anterior bud to grow upwards, and the posterior bud nearly

straight backwards. This process continues during the succeeding

stages, the anterior branch gradually taking the lead in size and

importance, and becoming divided distally into two tines. I can

see no escape from the conclusion that the anterior and posterior

buds of the very young antler in this stag are the homologues of

the corresponding buds in the young antler of the Barasingha

(Rucervus duvaucelli) shown in text-fig, 109. That being so,

it is clear that the anterior branch of the antler of Hlaphurus is

homologous with the “ brow-tine” and the posterior branch with

the “beam ” of the antler in the Red Deer, Sambar, Barasingha,

and other deer characteristic of the Old World. ‘The differences

between them are mainly a matter of size and direction of growth ;

that is to say, they are differences of degree and not of kind*.

Antler-Growth in a Species of Odocoileus.

Writing of the antlers of the typical American Deer, Mr.

Lydekker said *:—‘“‘ A large amount of misconception has arisen

with regard to the structure of the antlers of this group. In

1872 the late Dr. Gray rightly termed the single upright prong

arising from the inner side of the lower part of the beam of the

antlers of the Virginian Deer the ‘subbasal snag’; but this snag

* The subdivision of the anterior branch of the antler in Hlaphurus is, of course,

no argument against it being the homologue of the “ brow-tine,”’ for the latter not

infrequently, though abnormally, produces an additional snag in Elaphine and allied

groups of deer. In some species indeed, as in the Irish Elk and Cervus eldi, it is

commonly and normally provided with supplementary processes.

+ ‘Deer of All Lands,’ p. 246.

ANTLER-GROWTH IN THE CERVIDA. 78t

Sir Victor Brooke incorrectly identified with the brow-tine of

the typical Old World deer. This error has been pointed out by

Mr, A. Gordon Cameron, [who stated that] these characteristic.

tines have nothing in common with the true brows of Old World

types, and rise vertically from the inner side of the beam between

the coronet and the main furcation, usually converging at the

apex. ‘They are subject, in common with the antlers that produce

them, to all kinds of eccentricities; are frequently forked or sub-

palmate.”

Mr, Lydekker writes as if Mr. Cameron’s dictum settled the

question at issue; but it does not appear to me that much weight

can be attached to the reasons adduced by the latter for his

dogmatic denial of the truth of Sir Victor Brooke’s interpretation

of what Gray called the ‘‘subbasal snag” in the Virginian deer.

Except that the tine in question is situated on the inner side of

the antler, there is no great difference between it and the brow-

tine of the Old World stags, which is highly variable in direction,

as a comparison between the antlers of, e. ¢., Cervus affinis and

Rusa aristotelis will show. Not less does it vary in size and

structure even in nearly allied forms, as is testified by Dama

dama, where it is large, by Dama mesopotamica, where it is

sometimes almost suppressed, and by the Ivish Elk, believed to

be a Damine stag, where it may be palmated and branched.

The question to be settled, then, is this :— Does the position of

this tine on the inner side of the antler in the Virginian deer

preclude its being the homologue of the brow-tine situated on the

front of the antler in the Old World deer? Study of the growth

of the antler justifies, in my opinion, a negative answer to this

question and shows that Sir Victor Brooke’s opinion was correct.

Karly last year the Society received from the northern part of

South America a male specimen of Odocoileus, which I cannot

determine accurately. It is smaller and browner than a Venezuela

specimen identified as O. savannarum, but is otherwise very like

it. Its antlers are short, with the beam curved forwards in the

upper portion and ending in two tines, an anterior and a posterior ;

while on the inner side, near the base, arises the so-called “ sub-

basal snag.”

The growth of these antlers was very instructive. They started

as a simple exerescence, which soon began to divide into an anterior

and a posterior bud, the only difference between the antlers at.

this stage and those of a typical Old World deer being that the

anterior bud was slightly internal and projected a little inwards

as well as forwards. Nevertheless the two buds were perfectly

visible in profile view. The appearance of the antler at this

stage is shown in text-fig. 112, A, takenon May 12th. Four more

stages of the growth are represented in the following figures, B-Z,

taken respectively on May 22nd, May 30th, June 6th, and

June 17th, which show very markedly the gradual assumption of

an apparently more internal position by the anterior branch, its

782 MR. R. I, POCOCK ON

Text-fig. 112,

ans

Five stages (4 to Z) in the growth of an Antler of an American Deer (Odocoileus

sp. incert.), showing that the “subbasal snag” (@) and the “forked beam”

(p) were respectively the homologues of the ‘‘ brow-tine ” and the “beam ” of

the typical Old World Deer. Compare B with fig. 108, D.

ANTLER-GROWTH IN THE CERVID, 783

point of attachment to the posterior branch being completely

concealed from the external aspect in the last three stages *.

In view of these facts, I do not think it can be doubted that

the anterior bud which develops into the “subbasal snag” in

Odocoileus is the homologue of the anterior bud which forms

the brow-tine in Cervus. In that case the “ subbasal snag ” and

the “brow-tine” are homologous structures passing under different

names, and to state that Odocoileus has no brow-tine is merely

playing with terminology.

If this interpretation of the structure of the antlers in Hlaphurus

and in the species of Odocoileus above referred to be, as I believe,

correct, it shows that these two genera are widely divergent in the

very point upon which relationship between them has been claimed

to exist, and that the likeness, such as it is, between the antlers

of Elaphurus and of the Mule Deer (0. hemionus), for instance,

which has the so-called forked antlers without a brow-tine or with

the merest vestige of it, is purely a question of parallelism in

development ; that is to say, it has been brought about by growth

and modification of fundamentally different parts of the antler.

In the Mule Deer the anterior branch or brow-tine is to all

intents and purposes suppressed, practically the whole antler

heing composed of the posterior branch or “beam,” which is

highly developed and heavily tined. In Hlaphurus, on the

contrary, the principal part of the antler is composed of the

anterior branch or “ brow-tine,” which attains a large size and is

divided into two prongs, while the posterior branch or beam

remains comparatively small and slender and projects straight

backwards as a long often undivided prong.

* Tn connection with the date of antler-change in this Stag, attention may be

directed to its approximate coincidence with that of the typical elaphine deer of the

Old World; that is to say, the antlers were in the velvet during the summer months

and functional during the autumn and winter. They were shed in the early spring

and at the time of writing (July 3) the new antlers are nearly full-sized though still

in the velvet, exactly as in our Wapiti, Red Deer, Japanese Deer, and other Old

World species. The same is true of a specimen of Odocoileus americanus. On

the other hand an example of Mazama bricenii which shed in April 1908, and again

in April 1909, did not repeat the process till May 1911. He then carried a pair of

antlers for 25 months ; and those that started to grow in May 1911 are still on his

head. Thus Dr. Scharff (‘Distribution and Origin of Life in America,’ p. 111) is

mistaken in saying that the antler-change in American deer takes place at a quite

different time of year from that of Old World deer. It is well known too that the

time of antler-change at all events in some tropical Old World deer is highly variable

within specific limits. For instance, one example of C. dwvaucelli in the Gardens

regularly carries his antlers till about the end of May, while another of the same

species has antlers at least half their full size at that time.

(

84 MISS HELEN PIXELL ON

47. Polycheta from the Pacitie Coast of North America.—

Part I. Serputip#, with a Revised Table of Classi-

fication of the Genus Spirorbis. By Henen L. M.

PIxeLL, B.Sc., F.Z.8., Demonstrator of Zoology and

Reid Fellow, Bedford College, University of London.

[Received May 6, 1912: Read June 4, 1912.]

(Plates LXXXVIT.-LXXXIX.*)

Systematic: InDEX. Page

APOMatAssTAMSiW SP. ARs A, A eke

‘Protulamactieapsp: nid A. ls ent ay (eee GET SS

Classification of Spir07rbis.c.... se. c.cccesscdeveaideresseaee C92

Protoleeospira. SUDREN Da Soren eke, ce kee ees OS

Spirorbis ambilateralis, sp. N...........60000ce ee 798

SS PA CONOSUS, SP. Nes coe. en eer a dame hve ae EO

Ms EQUUS USD. De, cutycosscseaec gir mice eee ae he EEO OU

General Characteristics of the Family Serpulide.

1. Tube calcareous, nearly always attached to rocks or other

substratum for some part of its length.

2. Generally one or more branchie on dorsal side terminated

by an operculum.

3. Thorax, generally provided with a thoracic membrane,

representing the fused cirri and having 3-9 (usually 7)

segments.

4, Gland shields in thorax only.

Genus Serputa Linné (12) 1767, Philippi (21) 1844.

Generic characteristics T :—

1. Collar setz of bayonet-shape, with spines at base of blade.

2. Operculum funnel-shaped, with numerous radii ending in

serrations on margin.

3. Uncini with only a few large teeth.

. SERPULA COLUMBIANA Johnson (9), 1901. (Pl. LXXXVILI.

fig. 1.)

Serpula splendens Bush (8), 1905.

Serpula columbiana Moore (19), 1909.

Specific characteristics :—

1. Anterior abdominal setz with flarmg fringed ends, short

and deeply embedded, posteriorly replaced by small fascicles

of very long stiff spines.

* For explanation of the Plates see p. 805.

y+ An attempt is here made to summarise briefly the generic and_ specific

characteristics in every case. Such a procedure has not previously been adopted, so

far as I know, and it will, no doubt, in some cases be necessary at some future time

to modify such characteristies, but in the present confused state of our systematic

knowledge of the Serpulids, this seems to be a course likely to eliminate some of the

difficulties.

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NORTH AMERICAN SERPULID&.

POLYCHATA FROM NORTH AMERICA, 785

2. Uncini generally 6 or 7 teeth, the anterior one being the

largest (figs. 202 & 203 Johnson, 9).

3. Very large size (fig. 1) with numerous abdominal segments,

30 to 54 branchiz and from 80 to 160 serrations on edge

of operculum. '

Numerous specimens from Departure Bay, Dodds Narrows,

and two from Puget Sound (Prof. Kincaid’s collection).

Isolated tubes are to be found attached to the undersides of

stones on rocky shores near extreme low-water mark, and small

colonies may be found above this level in rock-pools.

In other places attached to rocks near low-water level the

masses of large white calcareous tubes are very striking—they

are thick and often finely ridged, the lower parts being much

intertwined, the free distal ends often overgrown with Polyzoa,

Spirorbis, etc.

The brilliant red colour on the branchial crown may involve

the whole of the gills and operculum, or these may be colourless

except for the tips, or they may be barred and mottled in a

large variety of ways. The remainder of the body is generally

yellowish.

As regards collar, thoracic membrane, and operculum with its

tubercles, this species agrees very closely with S. vermicularis,

so fully described by St.-Joseph (24. pp. 6528-335). Johnson (9)

presumably has made it into a new species on account of its

very much greater size with corresponding greater number of

abdominal segments, branchize and serrations on operculum,

together with small differences in the sete and uncini. He

cannot, I think, have examined many specimens, for he states

that the functional operculum is on the right side (9. p. 432).

The position of this in the genus Serpula may be right or left as

shown by Zeleny (27. p. 34), but out of 50 specimens that I

have examined 28 had it on the left, 21 on the right, and the

remaining specimen had one on each side. Consequently, when

he says that there are about 100 serrations on the edge of the

operculum, I cannot think that Bush (8) is justified in recording

this as a distinction between this species and SS. splendens with

127 to 150 serrations.

Moore (19) gives 140 for the one specimen of S. columbiana in

which he counted the serrations. In my specimens they vary

from about 80 to 160—the number apparently increasing with

age.

peoples distinction given by Bush is that there are in

S. columbiana “but .250 abdominal segments in a length of

55 mm.,” whereas Johnson says “250 or more” (9. p. 432), and

she gives 313 as the number in a specimen of her so-called

S. splendens, of which she does not state the size, and 190 ina

specimen 35 mm. long. These figures speak for themselves I

think as creating nothing but confusion. In 15 of my specimens

the average number of abdominal segments was 236, including 79

in a specimen 10 mm. long, 142 in a specimen 41 mm. long, and

310 in one 81 mm, long.

786 MISS HELEN PIXELL ON

The third point of difference given by Bush is that S. colwm-

biana has more numerous branchize—54 in each lobe as given

by Johnson—than her S. splendens, for which she records 45 to:

50 pairs.

In the specimen of the former species counted by Moore (19)

there were only 38. One small specimen of mine had only 18,

but the general number was from 30 to 50 on each side, one or

two large specimens having as many as 54—the number given

by Johnson. These facts show clearly I think that S. splendens

is a quite unnecessary species and can be included in S. columbiana,

which in turn, as pointed out by Johnson (9. p. 433), may be

identical with S. jwkesit Baird, for which however no satisfactory

description was given.

Numerous large free Selenidia in the trophozoite stage were

found in the alimentary canal of nearly every specimen of

S. columbiana examined.

Genus Crucicnra Benedict (1), 1886.

Generic characteristics :—

1. Collar setze and uncini similar to those of Serpula.

2. Operculum with comparatively few radii forming a scalloped

margin to the funnel and with conspicuous basal pro-

cesses. :

2. CrucicERA zyGorHoRA Johnson. (Pl. LXXXVII. fig. 2.)

Serpula zygophora Johnson (9), 1901.

Orucigera zygophora Bush (8), 1905.

Specific characteristics :—

1. Branchiz about 30 pairs with long filamentous ends to

rachises.

2. Operculum thick, shallow, with about 30 radii and 3 rounded

processes at its base; attached by a long pedicle.

One specimen from Puget Sound was 45 mm. long. A smaller

incomplete specimen came from Victoria (fig. 2). Another

specimen was only 7 mm. long and had a much thinner operculum,

but seemed otherwise similar.

3. CRUCIGERA IRREGULARIS Bush (3), 1905. (Pl. LXXXVII.

fig. 3.)

Specific characteristics :—

1. Branchize much coiled and with comparatively short fila-

mentous ends to rachises.

2. Operculum irregular, apex of funnel displaced ventrally and

the posterior and lateral walls deeper and rolled over to

some extent. Not more than two basal processes which

may, however, be bi-lobed and attached by long stout

pedicle (fig. 3).

About 12 specimens from the Channel outside Departure Bay

POLYCHATA FROM NORTH AMERICA. 787

and one from Dodds Narrows. Depth 15 to 25 fathoms. Tubes

generally solitary, attached to stones, shells, old wood, ete. Only

one specimen is recorded by Bush from Juneau. Tubes much

coiled, with flaring ends and one or two other conspicuous ridges

at intervals indicating the flaring end of a younger tube. Young

tubes seem to develop with a centimetre or so attached more

or less straight along the substratum, then to coil indifferently

to right or left, and only at a much later stage, if at all, to ascend

and form the flaring end.

General colour pale orange, the branchie and operculum

variously mottled and barred with red. Pinne sometimes

golden; ova greenish. Length varies from about 14 mm. to

50 mm. (fig. 3).

Largest diameter of operculum 1:3 to 4:5 mm.; the latter had

32 radii forming a thick scalloped edge. The pedicle was bifid

and contracted at the top just before joining the basal processes

of the operculum (fig. 3). Sets as figured by Bush (3); in the

posterior region of the abdomen the ordinary setz are replaced

by small fascicles of long slender spines.

Genus Aromatus Philippi (21), 1844.

Generic characteristics :—

1. Opereulum globular, terminating a gill retaining its pinne.

2. Some thoracic setz bladed sickles (sete of Apomatus)

(fig. 4 ¢). :

3. Terminal dorsal gland present.

4, APOMATUS TIMSII, sp.n. (Pl. LXXXVII. figs. 4a-4f.)

Specific characteristics :—

1. Collar setz simple tapered blades (fig. 4 0).

2. Branchie about 40 pairs with pinne nearly to the ends of

rachises.

3. Uncinigerous tori begin on third setigerous segment.

4, Uncini with numerous small teeth, the posterior one larger

with terminal enlargement (fig. 4 f).

5. Abdominal setz more or less sickle-shaped with some long

filiform ones in the last segments.

Eleven specimens from the Channel outside Departure Bay in

about 20 fathoms. This is the first time apparently that the

genus has been recorded from the Pacific Coast of America.

The tubes are solitary, adherent, and sinuous—one was attached

along its whole length to a portion of the Hexactinellid sponge

A phrocallistes whiteavesianus.

When full-grown about 80 mm. with 150 abdominal segments

(fig. 4a).

The branchial rachises are much coiled and almost colourless,

with pairs of red spots up their outer surfaces and only short

filiform extremities—the pinne appear green due to the contained

blood. The thorax is more or less orange or red, the thickened

788 MISS HELEN PIXELL ON

ridges bearing the tori being especially deep in colour, Ab-

domen pinkish; there is a short anterior asetigerous region, and

posteriorly nearly covering the last centimetre or so of dorsal

surface is a chi ky white strap-shaped raised glandular area. This

is the terminal dorsal gland mentioned by St. -Joseph (24) as

possibly serving in the construction of the tube. It tapers

anteriorly and ends abruptly posteriorly just above the vertical

slit-like anus.* Along nearly the whole of this glandular region

the ordinary abdominal sete are replaced by fascicles of about five

slender capillary sete without blades and nearly equal in length

to the width of the abdomen at this place. The collar is entire

ventrally and has a deep incision on each side—the lateral lobes

being continuous with the wide thoracic membrane.

The functional operculum is large and transparent and carried

by the second branchia from the dorsal side. The pedicle is

generally coiled twice like the other gills; except in very young

specimens there is a small club-shaped one terminating the

branchia on the opposite side.

All the collar sete and most of the other thoracic sete have

simple narrow blades; amongst them in the posterior thoracic

fascicles are to be found a few of the typical bladed sickles

(fig. 4c). The abdominal sete are more or less sickle-shaped,

though they easily become folded (figs. 4d, e) or straightened

out.

Several of the specimens examined were much smaller and

obviously young forms, with 70, 83, 101, 114 abdominal segments

and fewer branchie than in adult specimens.

Genus Proruta Risso (23), 1826.

Generie characteristics :—

1. No operculum,

2. Collar sete simple tapered blades (fig. 5 8).

3. Terminal dorsal gland present.

5, PROTULA PACIFICA, sp.n. (Pl. LUXXXVIIT, figs. 5 a—5 f.)

Specific characteristics :—

1. Ventral lobe of collar notched.

2. Abdominal setz somewhat sickle-shaped in the anterior

region, narrow terminal bladed ones posteriorly.

3. Uncinigerous tori extend from segment 3 to the end of the

abdomen.

4. Uncini with numerous small teeth, the posterior one long

with a bulbous extremity (fig. 57).

5. Thoracic sete simple blades with sometimes a few sete of

Apomatus posteriorly.

* In section the gland is seen to consist of epithelial cells crowded with spherical

eri anules which stain easily with iron hematoxylin but not with Delafield’s. The

gland is apparently unchanged by the presence of acid in the preservatives and

therefore does not appear to be calcareous.

POLYCH ATA FROM NORTH AMERICA, 789

Three specimens from Fairway Channel outside Departure Bay

in about 30 fathoms; one from Puget Sound (Prof. Kincaid’s

collection). No tube.

The specimens were colourless except for the branchize, which

appear greenish due to the contained blood. Down the outer

side of each branchial rachis was a line of opaque white spots.

The branchial crown (10-15 mm. high) easily falls off, leaving a

scar; the remainder of the body is 38 to 60 mm. long, rather more

than one-third of which is thorax. This is broad and flattened

with nearly parallel sides (fig. 5a) and the usual 7 setigerous

segments. The thoracic membrane is very wide, with an entire

margin which can extend beyond the setw# but is generally

considerably crumpled. The collar is notched in the median

ventral line and has a deep fissure on each side. The inward

coil of the stout base of the branchial crown is seen on the right

side of the specimen in fig. 5a. It takes 14 to 2 turns inwards

and upwards, the short inner gills reaching about the same

height as the outer long ones. An interbranchial membrane

connects the lower third of the gills, of which there are about

60 pairs having small closely placed pinnze almost to their

extremities. At the junction of the branchiz with the stout

basal membrane which carries them there is a slight ridge

visible externally, and at this place on the inner side arises

the oral membrane, which is continuous across the median line

and up each of the spirals. On the dorsal side of the mouth is

another shorter membranous lip.

Uncini begin on the third setigerous segment. They have the

characteristic shape (fig. 5f). The first two millimetres or so of

the abdomen is achzetous, nearly round, and of smaller diameter

than the wide dorso-ventrally depressed part which follows.

The ventral surface has deep segmental grooves showing 83-110

segments, and a wide feecal groove which turns to the right on

reaching the thorax. lLaterally the short tori are raised on

distinct parapodial processes which extend to the posterior end

of the body. The last 30 or so. segments on the dorsal surface

are covered with the calcareous-looking gland which anteriorly

tapers off to two points, suggesting a paired origin.

In section the gland is seen to occupy nearly the whole

thickness of the dorsal body-wall, the longitudinal muscles being

pushed towards the sides. Nearly all the cells are crowded with

the spherical granules or globules which stain easily with iron

hematoxylin, and are apparently similar to those in the glandular

cells so frequently found in the epidermis.

This hind region of the abdomen has very long sete which

extend 2 mm. or more on each side but are easily broken. At

first sight they appear to be simple spines very slightly bent at

_the extremities, but, with high magnification, a narrow striated

wing may be made out (fig. 5¢), shorter sete found with these

are more distinctly winged (fig. 5d). The other abdominal

segments have ventral fascicles containing about 13 short, stout,

Proc. Zoon, Soc.—1912, No. LITI. 53

790 MISS HELEN PIXELL ON

somewhat sickle-shaped sete (fig. 5c). These sete were, un-

fortunately, not examined before being preserved, and as

St.-Joseph (24. p. 338) points out, a lengthened immersion in

alcohol tends to reduce the curve of the sickle.

In general structure this species resembles P. capensis McIntosh

(13) fairly closely, but differs in the shape of the sete and

uncini: in the latter respect and in some other points it differs,

too, from P. diomedee Benedict (1). There are many eheraeeer

istics distinguishing it from P. superba Moore (19) and other

Pacific species that have been described. P. at ypha Bush (8)

might possibly be a young specimen of this same species.

Genus Cuitivopoma Levinsen (10), 1883.

Generic characteristics :—

1. No thoracic membrane.

2. Collar sete with fin-like expansion at base of blade.

3. Some of the other thoracic sete are sickle-shaped.

4, Abdominal setze geniculate.

5. Uncini with 9 or 10 fine teeth, the anterior one being

larger and blunter than the others.

6. Operculum with horny plate.

6. CHITINOPOMA GREENLANDICA. (Pl. LX XXVIII. figs. 6 a—6 e.)

Serpula triqueter Fabricius (7), 1780.

Hydroides norvegica var. grénlandica Morch (20), 1863.

Hydroides (¢) grénlandica Malmgren (14), 1867.

Chitinopoma fabrictti Levinsen (10), 1883.

Specific characteristics :—

1. Bodies elongated, somewhat cylindrical.

2. About 6 pairs of branchie with ends free from pinne.

3. Operculum enclosing central stalked vesicle.

Numerous specimens from Departure Bay and neighbourhood,

one incomplete one from Victoria. In thick sinuous tubes

adherent to shells, stones, etc., and having a very conspicuous

dorsal keel generally ending in a spine overhanging the aperture.

One tube was U-shaped with the two ends close together. The

largest specimen was about 12 mm. long (fig. 6a). The whole

animal was practically colourless, the pedicle of the operculum

and the branchie sometimes having faint transverse bands, and

the contents of the alimentary canal were in some specimens

dark rea.

The branchiz varied very much in number (from 6 to 8 in each

lobe) and some were frequently found in a rudimentary state:

one specimen had 7 in the right, and only 3 fully-developed

functional ones in the left. The operculum was in every case

on the left dorsal side, and I have found no trace of a secondary

one on the other side.

POLYCHATA FROM NORTH AMERICA, 791

The full-grown branchie had about 17 pairs of ciliated pinne,

which stopped short some distance from the top of the rachis

leaving a fairly long filamentary end. In sections the rachises

are seen to be strengthened externally by a thick layer of chitin,

which also forms a protecting layer round the pedicle (fig. 6 d),

and is continuous over the operculum with the thick horny plate

on its top. This plate was often covered with sand, and many

specimens had infusorians, etc. attached to it. The central

coelomic space in the pedicle is lined with peritoneum provided

with very large conspicuous nuclei. There is a small vessel

running along its whole length and enlarging in the opercular

cup into a spherical vesicle (fig. 6¢). This is filled with a finely

granular precipitate, and from its wall and general appearance

seems to correspond with the branchial blood-vessels of the

ordinary gill rachises, though I do not see that it can have any

respiratory function. It is apparently suspended in a fine

reticulate connective-tissue which easily shrinks away from the

epithelial cells ; Levinsen, in his original description (10. p. 203),

suggested that it might be a new operculum forming in the old

one.

The collar is very wide—the entire ventral lobe being generally

reflexed—the latero-dorsal lobes are continued down the dorsal

side to between the second and third thoracic sete, where they end

abruptly, giving the appearance of a short thoracic membrane.

This was a constant characteristic in both small and large

specimens. I have not seen it referred to before, but do not

think it necessary on that account to separate this as another

species.

With the large collar sete (fig. 66) are a few shorter curved

forms with very narrow blades.

The abdominal set agree with those described by St.-Joseph

(24) for the genus (fig. 6c).

The abdomen is long and slightly dorso-ventrally flattened,

with 25-40 segments. The tori contain about 17 uneini, but

there are only a few sete to each segment. The dorsal longi-

tudinal muscles are greatly developed—there is a small fascicle

of ventral ones on each side of the wide fecal groove. The large

ventral nerve-cords are separated from one another. The

epithelium consists of low columnar cells with numerous gland-

cells containing the usual spherical masses which stain easily

with iron hematoxylin.

This species seems to agree very closely with that described

by Bush (3) as Hyalopomatopsis occidentalis. J cannot under-

stand why this form with no thoracic membrane and “ trumpet-

shaped” abdominal sete is put in a genus, of which St.-Joseph

writes (24. p. 264): “Il m’a fallu créer un genre nouveau

Hyalopomatopsis pour le Hyalopomatus langerhansi Ehl. et le

H. marenzelleri Lang., qui par la présence d’une membrane

thoracique ... ne pouvaient rentrer, comme l’avait du reste

prévu Langerhans, dans le genre Hyalopomatus tel que Vavait

792 MISS HELEN PIXELL ON

défini von Marenzeller.” St.-Joseph also gives as another

characteristic of his genus Hyalopomatopsis, the presence of

capillary setee in all the abdominal segments.

Genus Spirorsis Daudin (6), 1800.

(Pls. LXXXVITI., LXX XIX. figs. 7-16.)

Generic characteristics and Schemes of classification :—

1. Caleareous tubes coiled in a dextral or sinistral spiral.

The method of coiling and the markings on the tube have

been used by Bush (8) in drawing up Table I., but the

coiling of the tubes is variable (cf. figs. 8 a and 6), being

determined to a great extent by the nature of the sub-

stratum, and as Caullery & Mesnil (5) have already

pointed out there is no constancy in either coiling or

markings.

2, Branchie are constant in different species, the operculum

with terminal calcareous plate, always occurring as the

second on the concave side (7. e. the right in dextral forms

and left in sinistral; since the animal lies with its back

towards the substratum).

. Thoracic segments generally 3—-the first having only dorsal

setee—the two following have on each side an uncinigerous

torus as well. In the” subgenera given the prefix “Para

by Caullery & Mesnil an extra torus (and in Sp. can-

cellatus a fascicle of dorsal sete also) is developed on

the concave side of the animal; this condition has been

described in the following Table of Classification (p. 794)

as 35 setigerous segments.

In Sp. ambilateralis, sp. n., there are four complete setigerous

segments, although the fourth on the convex side is very reduced.

This specimen therefore approaches the hypothetical Prospirorbis,

described by Caullery & Mesnil (5. p. 233), who point out

that the genus Spirorbis has been evolved from other Serpulide,

which have the characteristic greater number of thoracic segments,

by a gradual reduction. I have therefore placed this species in a

new sub-genus Protoleospira.

4, Abdominal segments 8-40,

Between the thorax and the abdomen is a more or less long

asetigerous region—often crowded with ova. The spermatozoa

develop in the posterior setigerous segments.

The thoracic sete are distinctive and the differences are of

use as specific characteristics. Table II. given by Bush

(3. p. 261) is drawn up with regard to these and the

direction of the coiling alone.

As a rule, the first thoracic segment has some slender capillary

sete forming the inferior part of the fascicle (fig. 14 6)—the

POLYCH MLA FROM NORTH AMERICA. 793

superior ones, referred to as collar setw, are distinctive, they may

have simple blades (fig. 13) or there may be a distinct fin-like

expansion at the base of the blade (figs. 7, 10@, 11a). In the

case of sinistral forms Caullery & Mesnil referred the former

to their sub-genus Romanchella, putting only those with a distinct

fin in the sub-genus Lagspira. In the species S. verruca and

S. evolutus Bush and S. medius, sp. n., however, the sete are

intermediate between these two types, having blades which are

faintly notched (fig. 14a), showing an indication of a superior

blade and inferior fin. Consequently I propose to do away with

Caullery & Mesnil’s sub-genus Romanchella altogether—taking

Leospira to include all sinistral forms with three setigerous

segments: it thus comes into line with the other three sub-genera

proposed by Caullery & Mesnil (cf Table of Classification).

The second thoracic segment has only ordinary bladed sete,

differing very little from one species to another.

The third thoracic segment has some ordinary bladed sete,

but generally also some bladed sickles as found so generally in the

genus Apomatus (fig. 14 e).

In many Pacific species there are present instead of the

ordinary bladed sickles a peculiar shorter form which appears

almost fringed at the extremity (fig. 10).

6. Abdominal ventral sete generally geniculate.

7. Uncini similar in thorax and abdomen—plates with the free

edge provided with fairly numerous fine teeth, the anterior

_one being larger than the others.

The following Table of Classification has been adapted from

that given by Caullery & Mesnil in their excellent paper on

Spirorbis (5) to include such of the Pacific forms as have so far

been studied. These authors pointed out that such a modification

would possibly be necessary.

Unfortunately the majority of the new species described by

Bush (2, 3, 4) from California, Alaska, and Japan cannot be

included owing to absence of information as to the number of

thoracic segments. Two or three of the species are established

on details as to the tubes alone.

Sub-genus ParApExiospira Caullery & Mesnil (5), 1897

(modified).

Characteristics :—

1. Tube dextral.

2. Thorax with 34 setigerous segments.

7. SprrorBis virrEus Fabr., 1780. (Pl. LX XXVIII. fig. 7.)

Serpula vitrea Fabricius (7), 1780.

Spirorbis vitrews Morch (20), 1863; Malmgren (14), 1867 ;

Levinsen (10), 1883; Caullery & Mesnil (5) 1897;

Moore (17), 1902; Bush (3), 1905.

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796 MISS IELEN PIXELL ON

Specific characteristics :—

1. Last thoracic segment on the right has no dorsal sete.

2. Collar sete with fin-like expansions at base of deeply serrated

blades.

3. Some sete of third segment bladed sickles.

4, Embryos incubated in the tube.

This species was quite common on stones and roeks from the

Departure Bay region and from Victoria.

The tubes vary a good deal but are always translucent, with

the whorls piled on one another. ‘The whole tube measures

about 2°5 mm. in diameter and the aperture about 1 mm. across.

There are various markings on the exterior. Sometimes a

ridge along the median line ends in a sharp projection above

the aperture—in the grooves on either side of this there may

or may not be scalariform markings. Young specimens have

shells as figured for this species by Levinsen (10. fig. 11), older

ones were ridged more like Sp. cancellatus (10. fig. 18). The living

animals were bright pink in colour. Gills 7, each with six pairs

of long opposite pinne, opercular plate a shallow funnel. About

20 abdominal segments.

Sub-genus Dexrospira (Caullery & Mesnil, 1897).

Characteristics :—

1. Tubes dextral.

2. Thorax with 3 setigerous segments.

8. Sprrorpis spirrntuM Linné, 1767. (Pls DXXX Viti,

figs. 8a—8c.)

Serpula spirilaum Linné (12), 1767; Fabricius (7), 1780.

Spirorbis spirillum Malmgren (14), 1867; Levinsen (10),

1883; Caullery & Mesnil (5), 1897; Moore (17), 1902;

Bush (3), 1905.

Spirorbis lucidus Montagu (16), 1803; Morch (20), 1863 ;

Malmgren (14), 1867.

Circeis armoricana Saint-Joseph (24), 1894.

Spirorbis borealis Fewkes (8), 1885.

Specific characteristics :—

1. Collar setee geniculate (fig. 8 c).

2. Operculum Seon brood-pouch.

3. Concave plate of operculum has a slight projection (talon)

on under side.

Two varieties of this species were fairly common in the Departure

Bay region and at Victoria, growing on calcareous polyzoa,

seaweeds, ete. The discoid form grows only on smooth surfaces,

Laminarian thalli being often extensively covered with specimens.

These were very flat, regularly coiled tubes—the spiral with

14 to 3 coils measuring *5 to 2 mm. in diameter (fig. 8a). The

POLYCHZTA FROM NORTH AMERICA, 797

ascending variety in some localities is much more common than

the discoid one, and it grows toa much larger size (fig. 8 6), often

attaining a height of 5 mm. It was generally found with its

lower coils overgrown with an orange Bryozoan growing on

Chetopterus tubes.

The living animal had a reddish colour, with a colourless trans-

parent operculum. A string of pink ova extended along the tube

beyond the posterior end of the body in some specimens. There

were 3 thoracic and from 12 to 20 abdominal segments.

The collar setz attain a length of -27 mm., much larger than

those figured by Caullery & Mesnil (5. fig. 46), they resemble

more closely in shape those given for S. armoricanus (5. fig. 5 6).

The differences between these two species were pointed out

by Caullery & Mesnil (5. p. 199) to be quite unimportant. All

that they could summarise were that Sp. armoricanus (Circeis

armoricanus St.-Joseph (24. p. 350)) was slightly larger, had more

abdominal segments (16-20) and a reduced talon to the operculum.

They record having seen intermediate forms themselves, and the

above observations as to variation in size, the number of abdom-

inal segments, and the collar sete consequently confirm their

opinion that Sp. armoricanus should be considered as a variety

only of Sp. spirillum.

9. SPIRORBIS PUSILLOIDES Bush (3). (Pl. LX XXVIII. figs. 9a,

9b.

S. pusillus Caullery & Mesnil (5), 1897. )

Mera pusilla St.-Joseph (24), 1894.

Non 8. pusillus Rathke (22 a), 1836.

Specific characteristics :—

1. Collar sete of a more or less geniculate fone (fig. 9a).

2. Embryos incubated in operculum.

3. Sete of 3rd thoracic segment Belletanee

4, Hepatic pigment reddish brown.

On stones from Taylor’s Bay, Gabriole Island. This animal

agrees in all important points with the full description by

St.-Josoph (24. p. 351) for Mera pusilla. The collar sete are a

little more distinctly angulated at the base of the blade (fig. 9 a),

but they have not the typical geniculate form described by

Caullery & Mesnil (5. p. 202). They are much shorter than

the setz of the second segment and are decidedly hooked. The

extent to which the sete are hooked has been pointed out by

St.-Joseph to be reduced by preservation (24. p. 338), so that

this does not seem to be a very important point.

The opercular brood-pouch is somewhat cylindrical and bounded

proximally and distally by calcareous plates, the talon is reduced

to a smal] quadrangular projection.

The abdomen has eight segments, with large, more or less

sickle-shaped sete (fig. 96) and an asetigerous anal segment.

798 MISS HELEN PIXELL ON

Sub-genus PRoroLaosPiIRa, nov.

1. Tube sinistral.

2. Four complete setigerous segments to thorax.

10, SPIRORBIS AMBILATERALIS, sp. 0. (Pl. LUXXXVIII.

figs. 10 @-10 e.)

Specific characteristies :—

1, Collar set very large, conspicuously serrated blades with

fin-like expansion at base (fig. 10 @).

Operculum without brood-pouch.

Talon of operculum with large hook-like process (fig. 10 6).

- Some sete of 3rd segment have fringed ends (fig. 10).

Some set of 4th segment ordinary bladed sickles.

Abdominal sete brush-like (fig. 10 e).

» Dorso-lateral brood-pouch.

“I> Ol Co bo

Several specimens on the inner sides of shells of Balanus

nubilus from Dodds Narrows, 15-25 fathoms. Tubes forming

translucent sinistral spirals measuring 3 to 4 mm. across. The

surface is distinctly corrugated outside, highly polished inside ;

the aperture measures 1 mm. in diameter.

Branchie 12—6 on the right, and the operculum with 5

others on the left. The pinne extend upwards and reach the

same height as the rachises. The opercular plate (fig. 10d)

is very like that of S. cornw-arietis Marion & Bobretzky (15).

From close to the point of origin of the pedicle arises a wide tube

which passes across the dorsal surface beneath the left lateral

lobe of the collar, and enlarges into the thin-walled brood-sae

which les along the dorso-lateral surface on the right side. This

structure has never, so far as Iam aware, been described before :

it is figured and more fully described for 8. racemosus, in which

species it was first seen.

Collar wide and entire ; the collar set rather more than 1 mm.

in length (fig. 10@). Sete of 2nd segment very numerous, all

with simple blades; these appear again in the 3rd fascicle, and

with them some shorter bladed sete with their extremities more

or less curved and fringed (fig. 10). The setze of the 4th thoracic

segment are few in number; on the convex side of one specimen

there were three bladed sickles (fig. 10d) and three with plain

blades. The uncigerous tori on the 4th segment are also smaller

than on the two preceding, especially on the right (convex) side.

The thoracic uncini are about 90 « long and have 20-25 teeth.

Those on the abdomen are only 25 « in length. About 50 of

the latter make up each of the tori which are very distinct on

the 18-20 abdominal segments: they begin quite close to the

posterior end of the thorax—the anterior asetigerous region of

the abdomen being very short.

The collar sete in this form seem to have been specially

POLYCHATA FROM NORTH AMERICA, 799

developed, for according to Caullery & Mesnil (5) the possession

of blade and fin is not a primitive characteristic, and they are

also very large ; possibly they are specially developed in correla-

tion with its habitat in rapids where the tide runs from 7-9 knots

an hour. The presence of the specially developed brood-pouch

might perhaps receive a similar explanation.

Sub-genus PARALOSPIRA,

1. Tubes sinistral.

2. Thorax with 33 setigerous segments.

11. Sprrorsis RACEMosUs, sp. n, (Pl. LXXXIX. fig. 11.)

Specific characteristics :—

1. Collar sete with fin-like expansion at base of very coarsely

serrated blade (fig. 11 a).

2. Operculum with massive talon having lateral wings and a

hook on ventral surface (fig. 11 6).

3. Thin-walled dorsal or lateral brood-pouch attached to the

thorax at the base of the operculum (fig. 11 d).

4. Some setze of 3rd fascicle bladed sickles (fig. 11 c).

A few large specimens from San Juan Island on barnacle-

shells overgrown with calcareous sponge; smaller ones from

Channel outside Departure Bay. Depth in both places 15 to

25 fathoms.

The largest specimen measures 5 mm. across the spiral; the

tube is corrugated and generally rather loosely coiled, leaving a

central hole.

Branchize 13—6 functional ones on each side; rachises have

short tapered extremities without pinnae; operculum not unlike

that of S. ambilateralis, but the talon rather more massive and

its outer concavity almost hemispherical and filled with débris.

The collar sete are very large (fig. lla); the 2nd fascicle

consists of plain-bladed sete, and the 3rd contain some fringed

sickles as well. There is a distinct third uncinigerous torus on

the left side (fig. 11d); the uncini have the usual shape.

There are 21 well-marked abdominal segments, each with three

or four brush-like sete and a long asetigerous anal segment. The

brood-pouch with its distinct wide stalk seems to be peculiar to

this and the last described species of Spirorbis: it does not appar-

ently replace a gill, for the calcareous operculum is developed as

usual on the 2nd to the left. The sac has a very thin wall, the

large ova causing marked protuberances on its surface, the whole

somewhat resembling a bunch of grapes (fig. 11 d).

Sub-genus Laospira.

1. Tube sinistral.

2. Thorax with 3 setigerous segments.

800 MISS HELEN PIXELL ON

12. Sprroreis asperatus Bush (3), 1905; Sitka. (Pl. LXXXIX.

fig. 12.)

Specific characteristics :—

1. Collar sete with simple blades.

2. Operculum without brood-pouch.

3. Calcareous plate of operculum thin, with large lateral wings

on its talon.

4, Some set of 3rd fascicle short fringed sickles (fig. 12),

others with long narrow blades.

On an old Serpulid tube inhabited by a hermit crab. One

specimen only. The opaque sinistral tube shows conspicuous

growth-lines, and the whorls are somewhat piled on one another.

There are 12 or 13 reddish gills nearly hidden by the high

collar, above which the operculum projects for some distance.

The alimentary canal was greenish and the ova along the dorsal

surface salmon-pink. The abdomen consisted of about 20

segments.

13. SprrorBis vALIDuS Verrill (26), 1874. (Pl. LXXXIX. fig. 13.)

S. verruca Levinsen (10), 1883, figs.+Caullery & Mesnil

(5), 1897 ; Moore (17), 1902.

S. validus Bush (8), 1905.

Specific characteristics :-—

1. All thoracic setz with long, finely serrated, narrow blades

(fig. 13).

2. Opereulum with brood-pouch.

3. Opercular plate (Bush (8), pl. xliv. figs. 11-14).

4. Branchie 13.

On old shells of Balanus nubilus from Dodds Narrows, 15 to

28 fathoms.

The tubes were smooth, sinistral, and opaque, measuring 3 mm.

in diameter. The operculum was colourless and transparent,

with a large sac-like brood-pouch which was, however, empty in

specimens collected on September Ist. The gills and thorax were

of a bright orange-red colour, the rachises being thick with a double

row of small pinne extending inwards at right angles. Abdomen

with 25 segments having a few sete, some being somewhat

geniculate, others small hooks (Bush, pl. xxxvil. figs. 5 & 6).

14, SprrorBIs. MEDIUS, sp.n. (Pl. LXXXIX. figs. 14a-14e.)

Specific characteristics :—

1. Some collar setee with shallow posterior notch (fig. 14 a).

2. Operculum without brood-pouch.

3. Calcareous plate of operculum very large and of character-

istic shape (figs. 14c & d).

4. Some sete of 3rd segment serrated bladed sickles (fig. 14 e).

From Channel just outside Departure Bay, Large flat tubes

POLYCHATA FROM NORTH AMERICA, 801

thick and opaque, slightly roughened, but without definite growth-

lines. A slight median ridge and sometimes one on either side ;

the aperture has, however, an entire margin and measures 2 mm.

ACLOSS.

The animal is of a uniform brick-red colour, and its total

length is 455 mm; there are 14 branchie—7 being joined at

their bases and situated on the right side; the other 6 func-

tional ones are joined with the operculum on the left. Each

rachis has a thin membranous projecting flap which wraps round

the outside of the next, and thus gives rise to a series of imbri-

cating semilunar membranes just inside the base of the collar.

The total height of the gills is 1 mm. There is a very wide

thoracic membrane on the right side which almost envelops the

whole animal. The calcareous plate of the operculum is 1-5 mm.

along its long axis; the outer side is concave and generally

covered with sand ; the talon projects obliquely inwards and has

large wing-like expansions at the sides (figs. 14¢ & d). The

collar set (fig. 14 a) appear to be in an intermediate stage between

plain bladed form and that with a distinct fin-like expansion at

the base of the blade—in one specimen I could find none of these,

only forms with the ordinary blades. In the 2nd segment there

were numerous setz of the normal kind, and the 3rd fascicle was

made up of bladed sickles (fig. 14e) and ordinary bladed sete.

The two tori in the thorax consisted of uncini of the ordinary

shape with about 20 teeth. In the abdomen were counted 20 to

25 segments, and the setz were of the ordinary geniculate type.

15. SPrRORBIS LANGERHANSI Caullery & Mesnil (5), 1897, and

Bush (3), 1905, from Panama. (Pl. LX X XIX. fig. 15.)

Specific characteristics :—

1. Collar sets with fin-like expansion at base of coarsely

serrated blade (fig. 15).

2. Embryos incubated in operculum.

3. Operculum of characteristic shape (5. fig. 22).

On tubes of Serpula columbiana from Departure Bay. Tubes

small, marked with lines and ridges.

The structure of the animals agrees with the description given

by Caullery & Mesnil (5. p. 217). They are 1-2 mm. long,

the abdomen being broad, with a long asetigerous region followed

by 9 short setigerous segments.

16. SprrorBis mMorcui Levinsen (10), 1883 (Greenland); Caullery

& Mesnil (5), 1897; Bush (8), 1905. (Pl. LXXXIX.

figs. 16 a-16 d.)

Specific characteristics :—

1, Collar sete about 12, with fin-like expansion at base of

coarsely serrated blade (fig. 16 a).

. Embryos incubated in operculum (fig. 16 3),

. Branchiz 8.

Cs ho

802 MISS HELEN PIXELL ON

Numerous specimens on tubes of Serpula columbiana from

Departure Bay. The opaque tubes are pressed together with

their ends standing erect, their bases being overgrown with an

encrusting sponge ; the surfaces are free from regular markings, the

apertures being circular and measuring about 1 mm. in diameter.

The animals vary in colour, some being almost colourless, but

were generally some shade of red or brown.

The sete of the 2nd and 3rd thoracic segments have long, very

narrow, delicately serrated blades (fig. 16c). A few of the 3rd

are shorter, with small posterior blades; these probably represent

straightened sickles (fig. 16 ¢).

There is a deep collar and a wide thoracic membrane on the

right side only-—this traverses the ventral surface obliquely

towards the end of the thorax and covers over several segments

of the abdomen. The gill rachises are thick and have long

pinne, the seven normal respiratory ones are rather taller than

the one bearing the operculum. ‘The latter is protected at

its extremity by a convex calcareous cap which extends nearly to

the base of the brood-pouch on the ventral side. The top is

quite opaque and slightly bilobed. Fifteen or more large reddish

ova were contained. Abdominal segments 20-29 with normal

geniculate sete.

17, SPrRoRBIS VARIABILIS Bush (3), 1905; Sitka,

Specific characteristics :—

1. Collar setz coarsely serrated blades with posterior fins.

2. Operculum without brood-pouch.

3. Talon of operculum without projections.

On stones and tubes of Serpula columbiana from Departure

Bay and neighbourhood. Thick tubes about 1 mm. across with

the outer whorl spreading over the others to some extent ; surface

markings were not very distinct.

General colour brick-red, with cerise ova along the dorsal

surface.

Branchiw 8 (including operculum), are nearly hidden by the

high collar. The thoracic membrane on the right side is very much

developed. The opercular plate had a marked concavity on its

outer side, and a small almost central talon projecting inwards.

The sete agree with those described by Bush (3). The abdominal

segments varied a great deal in number, from 15 to 28.

18. SPrroRBIS QUADRANGULARIS Stimpson (25), 1853.

Spirorbis quadrangularis Morch (20), 1863, and Bush (3),

1905.

Spirorbis fabriciti Malmgren (14), 1867.

Spirorbis carinatus Levinsen (10), 1883 + (11), 1886 +

Montagu (16), 1803.

Spirorbis affinis Levinsen (10), 1883.

Spirorbis granulatus Moore (17), 1902 + (¢) Canllery &

Mesnil (5), 1897,

POLYCH £TA FROM NORTH AMERICA, 803

Specific characteristics :—

1. Collar sete finely striated blades with basal fin.

2. Operculum with large brood-pouch.

3. Convex calcareous cap with long cylindrical projection

(figs. 14, 15, pl. xliii., Bush (3) 1905).

Victoria ; two specimens only obtained.

Tubes measure 2 and 3 mm. across respectively ; each has a

very conspicuous median ridge and two lateral ones, the outer

one being at the top of the perpendicular outer wall. The stria-

tions on the blades of the collar setee could hardly be distinguished

at all. Some sete in the third fascicle were serrated, bladed

sickles. The collar and thoracic membrane were well developed

on the right side, and there was a long asetigerous region

following the thorax. There were 12 abdominal segments with

geniculate sete.

Most of the specimens described in this paper were obtained

during a stay at the Marine Biological Station, Departure Bay,

Vancouver Island, in the summer of 1911. Mr. F. A. Potts

kindly collected those from Puget Sound and some others: I

should like to take this opportunity of thanking him, also the

Rev. G. W. Taylor, F.R.S.C., Curator of the Marine Laboratory,

Departure Bay, for his kindness and help with regard to dredging

arrangements, and Dr. Marett Tims for his advice and assistance

throughout.

REFERENCES,

1. Benepict, J. E.—Description of ten species and one new

genus of Annelids from the dredgings of the U.S. Fish

Commission Steamer ‘ Albatross.’ Proc. U.S. Nat. Museum,

ix. p. 547, pls. xx-xxv. Washington, 1886.

2. Busu, K. J.—On Spirorbis from Japan (see under Moore

(18).

3. Busu, K. J.—Sabellide and Serpulide. Harriman Alask:

Expedition, vol. xii. 1905.

4. Busy, K. J.— Description of new Serpulids from Bermuda,

with notes on known forms from adjacent regions. Proc.

Acad. Nat. Sci. Philadelphia, vol. Ixii. 1910.

5. Cautuery, M., et Mesniz, F.—Etudes sur la morphologie

comparée et la phylogénie des especes chez les Spirorbis.

Bull. Sci. France et Belgique, xxx. pp. 185-233. Paris,

1897.

6. Daupin, F. M.—Recueil de mémoires et de notes sur les

espéces inédites ou peu connues de Mellusques, Vers et

Zoophytes, Paris, 1800, p. 38. Nouveau genre de Ver a

tube caleaire, voisin des Serpules et des Dentales. Bull.

Sci. Soc. Philom., Paris, ii. p. 145, 1800.

7. Fasricius, O.— Fauna Greenlandica, Copenhagen anid

Leipzig, 1780,

804 MISS HELEN PIXELL ON

8. Fawkes, J. W. On the Larval form of Spirorbis borealis.

Am. Naturalist, xix. p. 247. Philadelphia, 1885.

9. Jonnsoy, H. P.—-Polycheta of the Puget Sound Region.

Proc. Boston Soc. Nat. Hist. xxix. pp. 381-437. 1901.

10. Levinsen, G. M. R.—Systematisk-geografisk Oversigt over

de Nordiske Annulata, Gephyrea, Chetognathi, og Balano-

glossi. Vidensk. Medd. Natur. 4, 1882-3, pp. 92-350.

Copenhagen, 1883.

11. Levinsen, G. M. R.—Kara-Havets Ledorme (Annulata).

Dijmphna-Togtets Zoologisk-botaniske Udbytte, pp. 289-

303. Copenhagen, 1886.

12. Linné, C. von.—Systema Nature, xir. 1767.

13. McInrosn, W. C.—Report on the Polycheta. ‘ Challenger’

Reports, xii. London, 1885.

14. Matmeren, A. J.—Annulata Polycheta. Ofvers. Vet.-Akad.

Forh. pp. 1-127. Stockholm, 1867.

15. Marion, A. F., et Bosrerzxy, N.—Annélides du golfe de

Marseille. Ann. Sci. nat. (6) ii. pp. 1-106. Paris, 1875.

16. Montacu, G.—Testacea Britannica. 1803.

17. Moors, J. P.—Descriptions of some new Polynoide, with a

list of other Polycheta from North Greenland Waters.

Proc. Acad. Nat. Se. Philadelphia, liv. 1902.

18. Moors, J. P.—Sabellide and Serpulide from Japan, with an

appendix on Spirorbis by K. J. Bush. Proc. Acad. Nat.

Se, Philadelphia, Ivi. p. 157. 1904,

19. Moors, J. P.—Polychetous Annelids from Monterey Bay

and San Diego, California. Proc. Acad. Nat. Sc. Philad.

si p-2od.) 1909:

20. Morcu, A. L.—Revisio critica Serpulidarum et Bidrag til

Rorormenes Naturhistorie. _Naturhist. Tidsskrift, i.

pp. 347-470. Copenhagen, 1863.

21. Puiipp1, A.—On the genus Serpula, enumeration of Medi-

terranean species. Ann. Mag. Nat. Hist. xiv. p. 153.

London, 1844.

22. QUATREFAGES, De.—Histoire naturelle des Annelés, Tome ii.

p. 488. (Suites 4 Buffon, 1865.)

22a. RatHKe, H.—Zur Fauna der Krym. Mém. Acad. Imp.

Sci. St. Pétersbourg, iii. pp. 407-429. 1836.

23. Risso, A.—Histoire naturelle des principales productions

de Europe méridionale, iv. Paris, 1826.

24. Sarnt-Josepu, Baron de.—Les Annélides polychétes des cétes

de Dinard. Ann. Sci. nat. (7) xvii. pp. 1-395. Paris, 1894.

25. Stimpson, W.—Synopsis of the Marine Invertebrata of Grand

Manan. Smithsonian Contributions to Knowledge, vi.

pp. 29-36. Washington, 1853.

26. Verritt, A. E.—Explorations of Caseo Bay by the U.S. Fish

Commission in 1873. Proc. American Assoc. Adv. Sci.

pp- 340-395. 1874.

27. Zuteny, C..—Experiments on the control of asymmetry in

the development of the Serpulid Hydroides dianthus.

Journ. Morph, vol. xxii, No. 4. Philadelphia, 1911.

Fig.

Fie.

Fig.

ile

POLYCHEATA FROM NORTH AMERICA. 805

EXPLANATION OF THE PLATES.

Pratt LXXXVII.

Serpula columbiana from left side, X 2.

Crucigera zygophora, anterior end from right side, X 2.

Crucigera irregularis, whole animal trom right side, X 2.

Apomatus timsii. 4a, whole animal from the right side, X 38, d.g., dorsal

terminal gland; 4, collar seta, X 300; 4c, bladed sickle-shaped seta

from thorax, X 300; 4d, crumpled anterior abdominal seta, X 300;

4 e, short posterior abdominal, seta X 300; 4 J, thoracic uncinus, X 300.

- Protula pacifica. 5a, ventral view, X $; 6b, thoracic seta, X 300;

5c, anterior abdominal seta, embedded up to the dotted line, X 200;

5d, short posterior abdominal seta, X 300; 5e, tip of long posterior

abdominal seta, X 300; 5f, abdominal uncinus, X 300.

Pruatre LXXXVILII.

. Chitinopoma greenlandica. 6a, dorso-lateral view, X 2; 65, collar seta,

X 460; 6c, abdominal seta, X 460; 6d, transverse section through

pedicle of operculum, v.s. stem of vesicle, X 55; 6e, transverse section

through operculum showing vesicle (v) suspended in a loose connective

tissue, X 48.

Spirorbis vitreus, collar seta, X 220.

Spirorbis spirillum. 8a, tube of discoid variety, X 48; 8b, tube of

ascending variety, X 10; 8c, collar seta, X 200.

Spirorbis pusilloides. 9a, collar seta, X 460; 96, large abdominal seta,

xX 460.

Fig. 10. Spirorbis ambilateralis. 10a, collar seta, X 220; 105, calcareous plate

of operculum, lateral view, X 48; 10c, seta from 3rd thoracic segment,

showing fringed sickle-shaped extremity; 10d, ordinary bladed. sickle,

from 4th segment (twisted end); 10e, brush-like abdominal seta. c-e,

X 220.

Pratt LXXXIX.

Fig. 11. Spirorbis racemosus, sp. u. 11a, collar seta, X 220; 118, calcareous plate

of operculum, X 48; Ile, seta from 3rd thoracic segment, X 460;

11d, dorsal view of whole animal showing stalkéd brood-pouch, < 12.

Fig. 12. Spirorbis asperatus, seta from 3rd thoracic fascicle, X 460.

Fig. 18. Spirorbis validus, collar seta, X 460.

Fig. 14, Spirorbis medius, sp. n. 14a, collar seta showing shallow posterior

notch, X 460. 144, ordinary capillary seta from 1st fascicle, < 460.

14, calcareous plate of operculum, dorsal view, showing talon and

lateral wings, X 16. 14d, latero-ventral view of opercular plate,

showing flat talon with lateral wing-like projections, X 16. 14, ser-

rated bladed sickle, x 460.

Fig. 15. Spirorbis langerhansi, collar seta, X 4.60.

Fig. 16. Spirorbis mérchi. 16a, collar seta, X 220. 160, operculum containing

ova, lateral view, X 48. 16c, narrow seta of 2nd and 8rd fascicles,

X 220. 16d, seta from 3rd fascicle, X 220.

Proo. Zoou. Soc.—1912, No. LIV. 54

806 MR. D. SELTH-SMITH ON A RARE LORY.

EXHIBITIONS AND NOTICES.

May 21, 1912.

Sir Epmunp G. Lover, Bt., Vice-President,

in the Chair.

The Srcrerary read the following report on the additions

made to the Society’s Menagerie during the month of April

1912 :—

The number of registered additions to the Society’s Menagerie

during the month of April last was 291. Of these 165 were

acquired by presentation, 61 by purchase, 17 were received on

deposit, 8 in exchange, and 40 were born in the Gardens.

The number of departures during the same period, by deaths

and removals, was 189.

Amongst the additions special attention may be directed to :—

1 Riippell’s Guereza (Colobus abyssinicus occidentalis) $, from

the Southern Cameroons, deposited on April 29th.

1 Black Jaguar (Felis onea) 2, from Para, purchased on

April 12th.

1 Capped Langur (Semnopithecus pileatus), 2 Elands (Tawro-

tragus oryx), 1 Common Camel (Camelus dromedarius), and

1 Ursine Tree-Kangaroo (Dendrolagus ursinus), born in the

Menagerie.

A collection of birds from Columbia, presented by W. K.

Pomeroy, Esq., F.Z.S., on April 6th, containing amongst others,

2 Purple Jays (Cyanocoraa affinis), new to the Collection; 1 Harpy

Eagle (Zhrasaétus harpyia); 2 Severe Macaws (Ara severa); and

2 Golden Hangnests (Iceterus xanthornus).

2 Limmergeiers (Gypaétus barbatus), from Russian Turkestan,

purchased on April 19th.

1 large Reticulated Python (Python reticulatus), from the East

Indies, presented by Rowland Ward, Esq., F.Z.S., on April 19th.

Mr. A. Buayney Percrvat, F.Z.S., exhibited a number of

photographs and lantern-slides of Game Animals from British

East Africa, including a fine series of the Reticulated Giraffe.

Mr. D. Sera-Smirn, F.Z.S., Curator of Birds, exhibited two

living specimens of a rare Lory, Calliptilus solitarius, from Fiji,

and remarked that Dr. Philip H. Bahr had recently brought

home two specimens which had died. The specimens exhibited

were from a collection of eight brought home alive by Mr. Rood

Tarte, of Taviuni Island, one of the Fijian group, where this very

beautiful species was still abundant, its numbers having been

very considerably reduced in the other islands by the introduced

ON THE PRESERVATION OF OUR NATIVE FAUNA, 807

mongoose. The exhibitor referred to a recent note on the species

by Dr. Bahr in ‘ The Ibis’ for April 1912, p. 298.

Mr. G. A. Boutencer, F.R.S., F.Z.S., read a paper entitled

*‘ Second Contribution to our Knowledge of the Varieties of the

Wall-Lizard.” This paper was a continuation of one published

in the Society’s ‘ Transactions’ in 1905, and dealt chiefly with the

variations of Lacerta muralis in South-Eastern Europe and South-

Western Asia. It also contained a supplement to the first part,

thus completing an account of the varieties, of which about thirty

were regarded as more or less definable, the author endeavouring

to show the inconstancy of the characters adduced by some her-

petologists in assigning specific rank to a number of these forms,

connected by many gradations. Mr. Boulenger hoped to support

his statements by a number of photographic figures of specimens

selected out of the enormous material which had passed through

his hands in the course of his study of this polymorphic and widely

distributed lizard.

This paper will be published in the ‘Transactions’ in due

course.

June 4, 1912.

E. G. B. Mzapr-WAtpo, Esq., Vice-President,

in the Chair.

Mr. E. G. Boutencer, Curator of Reptiles, exhibited a clay-

ball containing a cocoon of the African Lung-fish (Protopterus

annectens) presented to the Society by Capt. J. A. M. Vipan, F.Z.8.

He briefly alluded to the habits of the fish and the formation of

the cocoon, and gave an account of the method to be employed in

order to release the fish.

The Preservation of the Native Fauna of Great Britain.

Mr. E. G. B. Meape-Watpo, Vice-President of the Society,

introduced a discussion on the Preservation of our Native Fauna,

in which Mr. A. Heneage Cocks, Dr. F. G. Dawtrey Drewitt, and

Mr. Stewart Blakeney (who sent a written contribution) joined.

The necessity of creating public opinion on the matter was urged.

Tt was agreed that the laws with regard to birds were sufficient,

if administered strictly. With regard to mammals, it was the

opinion of those present that the use of steel traps, instead of

snares, for catching rabbits was chiefly responsible for the ex-

termination of wild cats, martens, and polecats in many parts of

the country, and ought to be suppressed.

54%

808 DR. HANS GADOW ON OVARIES

PAPERS.

48. The One-sided Reduction of the Ovaries and Oviduects in

the Amniota, with Remarks on Mammalian Evolution.

By Hans Gavow, M.A., Ph.D., F.B.S., F.ZS.

[Received May 31, 1912: Read June 4, 1912.]

INDEX.

Anatomical Structure and Development............... 803

Sanguine morphologists reckon that it takes about ten years

for their discoveries to find their way into a text-book. It

takes a generation to eradicate erroneous statements, especially

generalisations, out of such books, since most of them are

repeated from others without consultation of the immense host

of original papers. And it is apparently hopeless to expect the

enthusiast or amateur to appreciate the difference between a

generalising text-book of comparative anatomy and a zootomical

account. It means progress for a branch of science if we can

inscribe upon its statute-book a few lines of true generalisation,

which, if there be no hedging, require no longer any concrete

examples to be mentioned. If only partial generalisations are

possible, of course the exceptions are to be recorded and every

new case is welcome, until their accumulation in turn permits of

being summarised. ‘Then let there be drawn a line, and let the

discoverer of further cases keep his peace unless he has something

new to say.

The condition of the bird’s ovaries and oviducts is a case in

point. The main facts have by now become ancient history and

general knowledge to the zoologist, so ancient that the original

workers have been forgotten, as much as the name of the

originator of the term morphology.

That the ovaries and ducts of birds are one-sided was probably

known since time immemorial. Perrault * described and figured

them in the Ostrich without further comment. In the year 1810

Wolf mentioned that he had usually found two ovaries in the

Sparrow-hawk, a fact duly incorporated by Tiedemann } in his

excellent work, which reveals him as a zoologist far ahead of his

time. Next, Spangenbergt figured the right ovary in a Duck.

Barkow § described the occurrence of right-sided rudiments of

the female generative apparatus in various other birds. Emmert ||

observed equally large right and left ovaries in the Sparrow-

* PrERRAULT: Mémoires pour servir & Vhistoire naturelle. Amsterdam, 1736.

+ TrEDEMANN: Anatomie und Naturgeschichte der Vogel. 1810.

{ SPANGENBERG: Disquisitiones circa partes genitales foemineas Avium. Got-

tinge, 1813, 4to.

§ Barkow: Von der Kloake verschiedencr Vogel. Meckel’s Archiv f. Anat. u.

Phys., 1829.

|| KEumErr: Reil u. Authenrict’s Archiv.

AND OVIDUCTS IN THE AMNIOTA. 809

hawk and some other Accipitres. When Nitzsch made his

anatomical contributions to Naumann’s ‘ Naturgeschichte’ he

knew already of quite a» number of cases of double ovaries.

R. Wagner * added to them in his Lehrbuch, and still more in his

Beitriige. He described the cases of double ovaries accompanied

by double ducts; he knew that vestigial ducts are much rarer

than ovaries, and was well aware of the fact that even in Accipitres

with double ovaries there may be no traces of a second duct, ete.

Stannius f added further cases in his Lehrbuch. It was, how-

ever, mainly owing to Wagner’s lists that Duméril could give a

general summary in Cuvier’s posthumous edition of his ‘Anatomie

comparée,’ published in 1836. Owen also felt justified in

summarising without referring any more to special cases. On

p. 247 of his work he says:—‘‘The symmetry |of the ovaria] is

soon disturbed by concentration of development in the left

ovarium, the right one remaining stationary, and ultimately, in

most birds, disappearing.” On p. 249: ‘*Subsequently the left

oviduct alone proceeds to grow; the right is stationary, or

shrivels ; occasionally it may be discovered as a rudiment in the

mature bird, but usually all trace of it has disappeared.”

Still further instances of abnormalities have been described

and even figured without adding to or upsetting the generalisa-

tions quoted above. In most of the general text-books and

others, which have occasion to mention these organs, the

quotation of certain birds has become a regular stock in trade,

always the same, copied from one author by the next, who in

turn is quoted as the original authority by the more popular

writer.

So far as I know, I was the first to give a possible explanation

of the one-sided reduction, and I translate here what I wrote in

Bronn’s ‘ Thierreich,’ p. 842, published in 1890. “Tn all birds

only the left ovary is completely formed and functional; the

right is present in most cases, and may even produce unripe eggs,

but these degenerate later and seem never to become free. In

correlation herewith the right oviduct undergoes early reduction ;

at most it persists towards the cloaca as a ligamentous strand,

may be even asa tube which opens into the cloaca, ‘This one-

sided development of ovary and oviduct may be referable to

saving of space. Two completely developed, hard-shelled eggs

would scarcely have room in the belly, and we may add that even

in the left oviduct two complete eggs seem to be very exceptional,

if such ever occur. In Reptiles, however, both ovaries and ducts

are equally developed.”

This idea could not have been expressed more guardedly than

by the following sentence in the ‘ Dictionary of Birds,’ p. 783:

“This one-sided suppression of the organs may possibly be

* Wagner; (1) Lehrbuch der vergleichenden Anatomie, 1834.

(2) Beitrage zur Anatomie der Vogel. Abhandl. Miinch. Akad.

Wiss. ii. 1837, pp. 271-283.

- + Srawntrus’s Lehrbuch der vergleichenden Anatomie. 1846.

+ OwEn: Anatomy of Vertebrates, vol. ii. 1866.

810 DR. HANS GADOW ON OVARIES

referable to the inconvenience that might be caused were each

oviduct to contain an egg ready to be deposited.” Mr. Gunn *

takes a reporter's liberty by substituting for ‘ inconvenience ”

various gruesome calamities, as fracture of the egg-shell, rupture

of the oviduct, and even peritonitis! After thus having tried to

throw ridicule upon my generally accepted notion, he assures us

that such evil sequences need not be assumed at all because

of “the frequency with which Falconide are found with paired

ovaries, Which are obviously functional.” Then he proceeds to

distinguish between several theoretical possibilities besides the

only actual condition, namely, that in which the functional organs

are those of the left side.

Not a single case is known of a completely developed right

duct, whilst the left is vestigial. Further, I withdraw the state-

ment made in the ‘ Dictionary of Birds,’ that “but with rare

exceptions only that [ovary | on the left side becomes functional.”

‘“‘ Functional,’ I regret to say, was there used in a loose way,

since the right ovary not unfrequently forms rather large eggs.

But strictly it should be called functional only if any of those

eggs ever became ripe, 7. ¢e. burst from the ovary. We know

that even relatively large ovarial eggs, even those of the left

side, can undergo complete reduction ft.

Mr. Gunn, however, taking for granted that growing eggs in

both ovaries mean that both are functional, and that, although

two ovaries may not be a necessity, they must be better than one

(a principle which has produced the double-barrelled gun), has to

face the question whether one oviduct can serve two ovaries.

We are told that ‘‘ there is not much evidence for or against this

supposition,” and that ‘‘ there seems no physical objection to the

open end of the tube swinging across the mid-line of the spine,

and grasping the right ovum of the opposite ovary with nearly

the same facility as the ovum of its own side.” One physical

objection to this amazing trick-performance may be the gut with

its loops and mesenteries, and it is at least doubtful whether the

agile tube (the infundibulum of which is most carefully anchored

opposite its own ovary) can overcome these obstacles, in spite of

the best-intentioned regulating nerve-stimulus.

Let us enquire further into the meaning of the one-sided

reduction of the female bird’s reproductive organs.

Gegenbaur { favours the size of the egg, the complete egg with

albumen and shell, as the primary cause :—“ Bei den Végeln

gelangt nur das linke Ovar zu seiner vélligen Ausbildung dh. nur

* T. E. Gunn.—“ On the Ovaries in certain British Birds,” P. Z. 8. 1912, p. 63.

Mr. Gunn and the Rev. F. C. R. Jourdain credit the late Prof. Newton with the

authorship of the article “ Reproductive Organs” in the ‘ Dictionary of Birds,’ and

they suggest as his “ German source” of original information Taschenberg’s Intro-

duction to Naumann’s ‘ Naturgeschichte,’ re-edited in 1905, eleven years after the

+ Cf. A. von Brunn: “Die Riickbildung nicht ausgestossener Hierstockseier bei

Végeln.” Beitrage zur Anatomie und Embryologie, als Festgabe fiir Jakob Henle.

Bonn, 1882. 1 ; : n

{ Vergleichende Anatomie der Wirbelthiere, ii. 1901, p. 503.

AND OVIDUCTS IN THE AMNIOTA, 811

in ihm kommen Hier zur Reife, und das rechte erhiilt sich nur bei

einzelnen (manchen Accipitres, Schwimmvégeln und einzelnen Gat-

tungen verschiedener A btheilungen) fort, indess es bei den iibrigen

verkiimmert. Dies steht in Zusammenhang mit der Ausbildung

nur eines (des linken) Oviductes und mit dem bedeutenderen

Volum der Kier, wodurch jeweils nur einem einzigen ein lingerer

Aufenthalt in der engen Beckenhéhle gestattet ist. Hs ist also

hier die Riickbildung einer Hiilfte des gesammten weiblichen

Geschlechtsapparates von der Ausbildung des EHivolums abhingig,

und dadurch an einen héheren Zustand gekniipft, dass das mit

reichlichem Eiweiss und Dotter ausgestatte Ei das sich in ihm

entwickelnde Junge zu einer bedeutenderen Ausbildung gelangen

lisst.” The correlations mentioned in these cumbrously involved

sentences are valid enough, but they do not carry the question

any further than where I had left it.

Since the ovary is the prime organ and the duct merely

auxiliary, it might be assumed @ priori that the primary cause

of the reduction was the cessation of the production of ripe

eggs on one side, whereupon degeneration of the corresponding

duct would follow, as there would be no longer any work for it.

Illicit, undesirable traffic is stopped best by cutting off the

supply, in the present case by the stoppage of ripening eggs at

the source. But, as we have seen, this does not agree with the

facts, considering the frequent activity of right ovaries, whilst

right ducts are much rarer. Further, we know that even com-

paratively far advanced ovarial eggs can be resorbed. Lastly,

we should have every right to expect birds with right, and others

with left functional organs. It cannot well be assumed that the

one-sided reduction is an inheritance from reptilian ancestors, of

which unfortunately we know nothing. Of recent reptiles only

Crocodiles and Tortoises can be studied for the sake of analogy.

These produce for one clutch a considerable number of hard-

shelled eggs which pass through both ducts. The eggs are well

protected, and there is ample room for them in the broad body

of Chelonians, and there is likewise space and safety in the long

belly of a Crocodile.

Presumably therefore the cause of the asymmetry should lie in

the typical organisation of the bird. In proportion to its body

the eggs are enormous, especially in some of the nidifugous

groups which represent the lower conditions. They could not

well produce the whole clutch at once; and they incubate

their eggs, not merely because they require several days,

even weeks, to produce the full number, but because as warm-

blooded creatures they have reached a higher state of reproductive

organisation. There is no room within the pelvis for more than

one complete hard-shelled egg, leaving aside the inconvenience

of a right and a left egg which, for argument’s sake, might be

overcome, as is actually the case with reptiles. The available

space in the bird’s belly is limited ; the longitudinal distance is

relatively much shorter than in the majority of reptiles which

are devoid of a sacrum, and the peculiar pelvis of birds is as

812 . DR. HANS GADOW ON OVARIES

broad as is compatible with the upright walking and with the

flying organisation. Most likely the broadness and the absence

of symphyses have been produced in adaptation to the eggs, but

even the distention of the belly downwards must be limited in

the bird, which is essentially and primarily a flying creature.

Well, then, let us take it that it is advantageous that one of

the ducts and the activity of the corresponding ovary should be

suppressed. Instances of asymmetry, brought about by sup-

pression of one of originally paired organs, are common enough

in the Vertebrata, and they can in most cases be fairly explained

by mechanical factors. To refer them to mere accident, to a

toss up, which then becomes established, is too shallow a mode,

although not unprecedented in morphology. The tadpoles of

some Anura have paired “spiracles,” others a median, the majority

a left hole. The reduction of one of the lungs of Snakes and

snake-shaped Lizards is of course directly correlated with the

shape of the body, and it appears almost optional whether the

right or the left lung should be affected, since both cases have

become established in the various groups.

If we apply the principle of elimination of all those unfortunate

hen-birds which happened to produce eggs in either side, whilst

only those birds propagate the race which happen to have only

one side in working order, this would not explain the universal

right-sided suppression, which according to Gegenbaur is a

weighty argument for the monophyletic origin of the class*. If

we assumed this as a proof of their monophylism, we should

logically arrive not only at the imaginary pair of ‘“ Urvogel” but

also at the Eve of hens, which in her case would relegate the

establishment of the asymmetry to a toss up. During the

presumably long period of dawning bird-life such a one- “sided

incipient suppression must have taken place over and over again

before it was firmly established. Inheritance, if not swamped

by panmixis, might have established asymmetry, but once more

we are groping in the dark for a cause which favours the left

side. It must be a factor which is very ancient and yet does

not interfere with the symmetry of the male organs, neither the

testes nor the vasa deferentia. Since the ovaries are strictly

homologous, or rather homogeneous, with the testes, this may be

taken as another hint that the ovaries are not the parts primarily

affected. But the male ducts are not the same as the female ducts,

therefore the latter are indicated. No factor causing the

asymmetry can be derived from the vascular system, nor even

from the vestiges of the renal portal system, by the suppression

of which birds and mammals differ from their common ancestors,

the reptiles. To refer the enlarged left ovary and duct to the

* The suppression of one lung in Snakes, etc. stands on a different footing. It

may be due to an accideut or sport, as much as right- and left-clawed crabs, or

right- and left-twisted shells. The remaining lung enlarges and shifts its position

so as to occupy most of the space originally intended for both.—H.

AND OVIDUCTS IN THE AMNIOTA. 813

stronger arterial supply would of course mean mistaking effect

for cause *,

A sufficient cause, however, may be the fact that a full ovi-

duct is less liable to disturb the other intestines in the left half

of the body-cavity than in the right. The primary intestinal

loops are so arranged or packed, that their bases begin on the

‘right whilst their apices extend towards or into the left side of

the abdomen. This is especially the case with those loops which

starting to the right of the stomach (itself mostly shoved to

the left) fill the space between stomach and vent with their

distal halves. It needs no further comment that it is the

free or apical end, and not the base of a loop where the

mesenteric vessels enter, which is displaced easiest and which

will easiest resume its original position. But this packing

from right to left is not an adaptation to, and is not produced

by the preponderance of the left oviduct. It can be traced

to a much more primitive condition, namely to the fact that

the bird’s embryo comes to rest with its left side upon the yolk,

with its curved back towards the blunt pole. In all probability

this is a truly cenogenetic feature, essentially ontogenetic; one

of those numerous phenomena which, like the gills of tadpoles,

the allantois and placenta, are originally incidental to embryonic

life, although they may by correlated after-effects profoundly

influence even the adult organism. Obvious results of this left-

sided position of the embryo are the increasing preponderance of

the left vitelline vein ; the yolk-stalk causes the first loop of the

midgut; the stomach itself sinks in, turning the pylorus to the

right, upon which side the duodenal loop descends, and further

secondary loops of the midgut follow suit. If there are large

cxca, they likewise make their way towards the right and back

of the stomach. The allantoic bag, containing fluid only, rises

and comes to lie upon the embryo, 7. e. upon its right side.

Consequently there is asymmetry introduced at an early date,

which affects the viscera, notably the gut, and introduces a bias

in their mutual behaviour within the belly. During the growth

of the embryo, by shrinking of the yolk room becomes available

for extension of the gut towards the left side. The permanent

organs will soon—speaking from the point of evolution—establish

an equilibrium, whilst it is clear that any occasional or contingent

requirement of space, or disturbance, can be met with easiest in

the left half. Such a disturbance is caused by the periodic

growth and passage of the eggs which brook no delay. The

slightest bias will turn the scales, and now we may apply the

censorship of natural selection to its fullest extent. Left eggs

* The suppression of the right oviduct has had an effect upon the male copulatory

organ. Where such is present it is asymmetrical, although unpaired, and stowed

away in a left-sided recess of the cloaca. The act invariably takes place from the

left side, and the same applies to those birds which are now devoid of such an intro-

mittent organ.

814 DR. HANS GADOW ON OVARIES

and a left duct will be the least liable to set up complications.

Two canals may be good, but one improved way is better, and if

the traffic goes in one direction only, the other duct falls into

abeyance. If goods are still produced at the terminus of the

obsolete line, they will deteriorate, but this does not matter if

the output of the opposite factory is equal to the demand.

So far so good, and the enquiry need not be carried further

back, if it were not for the Monotremes. Although these archaic

creatures show no appreciable difference in the size of their

paired ovaries and ducts, only those of the left side are functional.

According to Semon, Ornithorhynchus invariably produces two

eggs, always in the left side; Hchidna lays only one egg, also

left-sided. The right ovary forms numerous large eggs which

never ripen, and the respective duct and uterus are swollen and

much vacuolised during the season. In short, Monotremes behave

exactly like certain abnormal birds, e.g. the famous Sparrow-hawk,

by the retention of an ancestral feature which is now normally

lost. Since the reduction in the Monotremes has made so little

progress, it looks as if it were but of comparatively recent date,

but at the same time so ancient as not to have interfered with

the inheritance of the full symmetry by the Meta- and Eutheria.

The Monotremes are no longer quite primitive, not even in these

organs. Their eggs have lost much of the yolk; they continue

to grow in bulk within the uterus after they have received their

keratine shell. Indeed, we cannot well imagine that, compared

with oviparous reptiles and birds, the very small egg of the

Monotremes, and the imperfect, almost larval condition of

the new-born represent truly ancestral conditions, unless—

and this is well worth further enquiry—we are prepared to

assume that in all Vertebrata the viviparous condition was

primary to one in which the fcetus is surrounded by a shell and

then hatched outside the mother. If this should be the case, we

should further have to distinguish between primordial viviparity

(of which recent examples are unlikely) and secondary, pseudo-

primitive viviparity, the numerous instances of which have been,

and are still being, acquired independently: many Sharks and

Teleosts; many Urodela, even one or two of the Anura, and

many scattered cases among the reptiles, as some Chameleons

and Lacertide, Iguanide and Anguide, all the Scincide, all the

thoroughly aquatic snakes, the Viperide, and here and there

some other terrestrial kinds. But to return to the Monotremes.

Can their incipient, or perhaps arrested, asymmetry be referred to

the same embryonic conditions as those which prevail in Birds ?

The bulk of the egg is formed by the yolk, the yolk-stalk might

be strong enough to cause a disturbance, the allantois protrudes

towards the right, and the left vitelline vein preponderates.

How far, and if at all, the viscera are affected by these conditions,

remains unknown. For our purpose it is significant that there

is incipient asymmetry (functional although scarcely structural),

and that this should be restricted to the only recent Mammals

AND OVIDUCTS IN THE AMNIOTA. 815

which still possess comparatively large-yolked eggs. However,

the Monotreme embryo does not turn upon its left side, it merely

sinks into the cavity of the emptied yolk-sac, forming a

proamnion ; the bird, owing to its enormous yolk, turns over

and ultimately comes to lie on its curved back; the reptilian

embryo must also sink in, but it does not turn. This turning

over, so marked a feature in the bird, may be correlated with

the conditions of incubation. Everybody knows that the

chalaze keep the blastoderm “on the top,” i.e. nearest the

source of warmth, against the hen’s body *. This does not apply

to reptiles which deposit their eggs in the ground, nor to the one

or two eggs in the moist pouch of the Monotremes; lastly, to the

embryo of viviparous and ovoviviparous creatures “ orientation ”

towards the source of heat is not only unnecessary, it would

also be impossible because the mother changes so much her own

position by moving about.

Since no trace of functional asymmetry of ovaries and ducts

appears in Meta- and EKutheria, and as that of the Monotremes

cannot well be a reptilian inheritance (because their asymmetry

is usually marked by an enlargement of the right gonads and

ducts, e.g. Snakes and Crocodiles, both sides however being

functional), the asymmetry of the Monotremes must be due to a

departure within the Prototheria, but so slight as not to have

caused any irreparable morphological reduction of either ovaries

or ducts by the time that the Prototheria entered upon the next

higher or Metatherian stage, excepting of course the Monotremes.

If Prototheria ever laid eggs much larger than they are now, the

asymmetry may have been greater and be referable to the same

primary causes as those suggested for birds, but since the recent

Monotremes seem to be actually in the process of reducing them

now, and moreover to the last possible number, the left-sidedness

seems to be a case of mere coincidence with birds.

Some simplification of the completely double female apparatus

of the Vertebrates was bound to come; it was a matter of time,

and success depended upon the grade or height of the general

organisation of those who attempted it. Any agreement

between birds and mammals versus reptiles cannot be anything

more than convergent resemblances, at best cases of Isotely.

The classes of both Birds and Mammals have gone beyond the level

of the Reptilian organisation and they represent highest termini ;

but, although the class of Birds is by far the most specialised and

in various respects has reached seemingly unsurpassable perfec-

tion, the class of Mammals is morphologically the highest, in

spite of its still comprising such lowly, undecided types as the

* The usual statement that the hen turns the eggs over from time to time in

order to ensure the equal warming of the whole egg, now the upper and then the

lower half, implies nonsense. What the sitting bird does, is to rearrange the

position of the eggs with reference to each other, to give those now lying peripherally

an equal chance of best position near the centre. With a small clutch this is not

necessary, but with a dozen eggs the frequent rearrangement is very noticeable, at

least with thoughtful sitters.

816 . DR. HANS GADOW ON OVARIES

Monotremes (cf. loss of the nucleus of red blood corpuscles,

structure of the atrio-ventricular valves, the alveolar lungs, the

abolishment of the cloaca, the cranio-dental articulation of

the underjaw, etc.).

The Elasmobranchs show various instances of precocious in-

ventions, ahead of the times, foredoomed to failure or further

improvement, because their owners are after all but low fishes.

In some the ovary is single, unpaired, but it lies in the middle.

Some Acanthopteri have succeeded in producing a median duct

out of their otherwise paired ovarial sacs. Birds, as we have

seen, have suppressed one side; a clumsy mode of procedure

because it has a lopsided result and implies the reduction, by

neglect, of one of the precious gonads. It is only the higher-

graded organisation of the Mammals which has succeeded in

simplifying the apparatus in the morphologically neatest way,

namely, by the partial fusion of the two ducts into one passage,

not only unpaired, but median, whilst the upper ends and the

ovaries remain intact and functional.

Monotremes, Marsupials, and Placentals form an unbroken,

progressive, therefore most probably monophyletic series. The

reduction in question could not be brought about until the

reptilian plan of hard-shelled eggs had given way to internal

gestation. The differences between oviparous, OVOViviparous,

implacentally and placentally viviparous, are questions of degree

only. ‘There is for instance no difference, to be expressed in a

few words, between the ovoviviparous fruit of a Viper and the

newborn feetus of a Kangaroo, except that the newborn reptile is

complete and must shift for itself. The point is that the young

bursts its egg, and other membranes, with the act of birth.

Whilst no newborn reptile requires maternal care, most birds do,

and all mammals are absolutely dependent on their mothers for

nourishment.

Within the Class of Birds every stage from almost reptilian

to practically mammalian conditions is represented. The

typical nidifugous birds are hatched with still a considerable

amount of yolk slipped into the belly, sufficient for the little ones

to hold out for days without food ‘until they are bodily and

mentally strong enough to feed themselves; in many eases they

have to be shown by the parents how to do it. Next come those

which are hatched in a more or less helpless condition and must

be fed by the parents with food either in its natural state or

already semidigested. Lastly, those which are nursed with a

milky secretion of the crop. The higher Altrices or Nidicolz are

born with but small remnants of yolk left, the digestive organs

having been hurried on at the expense of the others. | The most

significant point, however, is that through the crop-secretion

the Pigeons have established a parallel with the Mammals in so

far as the young are fed actually with parental matter, in which

proliferation and fatty degeneration of epithelial cells plays a

great part. The analogy can be carried still further, since by

AND OVIDUCTS IN THE AMNIOTA,. 817

Pigeons and some of the lower mammals this “ milk” is squeezed

or injected into the young.

The Mammalian evolution has probably gone through many

stages and vicissitudes, difficult to enumerate, because there were

many factors and not all the organs changed at the same time.

1. We start with a hypothetical stage of Sauro-mammalia.

The hard-shelled egg, when laid, contains an already far advanced

embryo, therefore ovoviviparous. This egg was not so much

incubated for warmth as covered for protection by the mother.

The young although hatched in a reptilian condition was pro-

tected by the mother.

2. Reduction of the size of the egg. Gradual preponderance

of extra-uterine nursing over uterine gestation, made possible

and introduced by the protecting insulation, in such a way that

an abdominal incubation area was developed, owing to the

reaction of the mother’s abdominal surface by the sat-upon,

adpressed, covered egg; this resulted in hypertrophic condition

of the cutaneous blood-vessels, hence of the glands, and correlated

reduction of hairs. Incidental change of absorption of moisture

through the porous shell, enhanced by loss of its calcareous

portion.— When this suppression had been well established, by

progressive inheritance, the now quite porous parchment egg had

the same chances of absorbing fluid whilst still in the oviduct.

At the same time it stands to reason that the chances of external

or brood-pouch nourishing may beconie prevalent. The shorten-

ing of the life-period within the egg implies the birth of an

unripe fetus. Fetal life must be taken as ending with the

bursting of the egg, no matter whether this act coincides with

the moment of parturition, or whether it happens some time

after the egg has been “laid.” In either case it coincides with

the cessation of any further possibility of function of yolk-sac

and allantois.

We must further assume that the actual length of time re-

quired for the production of a young animal is the same in

equal-sized creatures from the beginning of segmentation until

it is independent. Unless we assume this, the argumentation

would become too complicated.

Let us say that it took 50 days from impregnation until the

Sauro-mammalian youngster was ready to face the Permian

world. This means:

50 days of ovoviviparous, internal uterine life ............... ...(Sauro-mammal),

40 days uterine and 10 days incubation-life Mes ‘ie tide egg.........(Hypotheria).

ges a 105; a and 10 days nursing in ton

(Prototheria).

20 ” 33 5P ” ” 25 ” ” (Monotremes).

8 ” 9 0 ” ” 42, ” ” (Opossum),

The suppression of the incubation-life marks the Metatherian

stage just as sharply as the introduction of incubation of a

feetus marked the early Mammal stage. The adaptation to the

818 DR. HANS GADOW ON OVARIES

new invention of pouch-life and nursing caused the development

of an entirely new category of features—of features which were

not required either by the fetus or the adult, therefore larval,

é. g. & Suctorial apparatus with its far-reaching incidental in-

fluences upon the future adult structures.

In these respects viviparous reptiles, Hypo- and Prototheria

culminating in Monotremes, and Metatheria culminating in

Marsupials, represent a continuous progressive series, with a

logical terminus characterised by the enormous preponderance

of extra- over intra-uterine development. Compared with these

terminal Marsupials the Eutheria seem to be totally different,

provided we take as their type those which are born complete,

in this respect like the hypothetical Sauro-mammals, the whole

of the “50 days” being intra-uterine. And yet the Eutheria have

with certainty passed through the same Metatherian stage as

have the Marsupials, and this a stage comprised,

besides others, the following features*: Truly viviparous ; allan-

toic placenta ; marsupium; diphyodont teeth, the same two

middle series of a total of prelacteal, lacteal, permanent and post-

permanent sets; nipples; semi-cloaca; absence of a corpus

callosum.

To arrange any one of these features into successive morpho-

logical stages is comparatively easy, but it does not follow that

these represent exactly the phylogeny of the groups, because of

the complicated correlations with other organs which by no

means keep step with each other, neither in the same species nor

in the greater groups. Some are precocious, even hypertelic,

while others lag behind.

Just as to the large egg of the truly oviparous Sauropsids

albumen (more watery but less fatty yolk) is added, before it

receives its calcareous shell, so in the Monotremes fluid is added

to the contents of the egg, but with the remarkable difference

that fluid matter is taken into the yolk-sae itself by osmosis

from the uterine walls, after the keratine shell has already

been developed. This process is correlated with an wun-

doubted previously acquired reduction of the amount of ovarial

yolk, and is as much a secondary process as the loss of caleareous

matter in the parchment-like ‘‘keratine ” shell.

As Semon has shown, the whole shell-enclosed egg multiplies

its size during its passage through the oviduct. This mode of

growth finds a curious parallel analogy in various Lacertilia, the

parchment- -shelled eggs of which grow considerably after they

have been deposited.

Whilst the Sauropsidan allantois comes to surround the whole

yolk-sac and also nearly the whole of the albumen, so as to

spread over most of the inner surface of the egg, the Monotreme

* Cf. R. Semon: “ Monchgpaun. Marsapinlien) in Zoolog. Forschung. Australien,

ii. 1894-97; further, J.T. Wilson and J. P. Hill’s papers in Q. J. M. 8. 1897, 1898,

1900.

AND OVIDUCTS IN THE AMNIOTA. 819

allantois and yolk-sac balance each other. The shell-enclosed egg

is for a short time transferred into the marsupium (cf. Semon,

Echidna; not into a bursa as was imagined by Klaatsch, Gegen-

baur, and others), which secretes fluid (the “ nutritive sweat” in

Gegenbaur’s unfortunate diction), and this can be taken up by

the embryo through the porous shell. This may be the reason

why the shell is soon cast off.

In Marsupials the shell is at first still present, but soon absorbed

within the oviduct. The egg-membranes, etc., of the embryo

establish no structural communication with the uterus. The

ovarial yolk is much reduced. But the yolk-sac becomes en-

larged, as in the Monotremes, still occupies a great portion

of the inner egg-surface, and has established intimate contact

with the serosa. The allantois, being independent of the expansion

of the ceelom, which results in the driving away of the yolk-sac

vessels from the somatopleure, establishes a villous placenta.

Such must have been the condition of the Metatheria.

In Perameles the allantois still reaches the surface, where it

is very vascular, and fused with the serosa, a truly respiratory

arrangement. ‘The.placenta being lost in most other Marsupials,

the allantois reverts to its primary function of urinary receptacle,

although apparently late during the foetal life.

The Metatherian stage may therefore be characterised as one

in which the posterior of the two bags, the allantois, has super-

seded the previous attempts of placentation by the yolk-sac.

The new placenta was perhaps not advanced enough to prevent

the foetus from being born soon after the limited amount of yolk

was used up. Certainly it did not pass beyond the non-deciduous

stage, and it never reached the extent of even the lowest recent

Eutherian placenta. Yet one effect of this incipient organ must

have been to render the foetus less independent than that of a

viviparous reptile. It had therefore still to be transferred into

the marsupium, there to be kept moist and suckled in as

premature a condition as the Monotreme.

Two opposite tendencies are inherent to this stage. One

palingenetic, to give birth when the yolk is used up; the other

cenogenetic, to prolong the retention of the foetus because of the

compensatory, respiratory, etc. advantages incidental with a

placenta. Obviously the Metatherian stage was a half-way

house at a parting of the ways to further improvements, leading

to Marsupials* and to Placentals.

* We have here an instance of the well-known fact that Group-names based upon

single anatomical features are mostly unsuitable for classificatory purposes. ‘The

taxonomic value of these characters may be good enough, but they are not diagnostic.

If such names were used merely as labels without much intrinsic meaning, well and

good, but even the best of us cannot, on occasion, resist taking their face-value for

full value. There are Mammalia Implacentalia with a placenta, and we now know

that the young Echidna does not lie in a bursa. Odontornithes are a valueless,

heterogeneous assembly, and overcontidence in “ Ratite” was responsible for

having branded Hesperornis as a “swimming Ostrich.” “ Mammalia” is for-

tunately an excellent term, although invented before Monotremes were known.

820 DR. HANS GADOW ON OVARIES

To judge from analogy the new organ should continue straight

on, leading in this case through a diffuse and cotyledonary to a

solid, and from a non-deciduous to a deciduous placenta, ter-

minating in the birth of the young in an utterly helpless con-

dition. Asa rule the non-deciduous and diffuse placenta is found

in those mammals which at birth require least maternal care.

The latter implies greater mental capacity. But we also know

from analogy that a recent invention—although it may be

capabie of much perfection—is not always kept up if the old

string to the bow 1s still capable of further adaptation. Where

both alternatives are good, workable, and capable of being

improved, it may take a long time to settle which is after all the

better of the two.

What has induced some Metatheria to neglect the further

development of their placenta and to intensify the marsupial

alternative? The morphological and physiological momentum

was surely on the side of improving placentation. There

must have been some external, environmental influence, and

it was Dollo’s brilliant suggestion that arboreal life was the

underlying cause. His further explanation is less acceptable,

that the climbing habits have caused premature births, which

have become habitual, and that the hapless foetus had therefore

to be nursed and located in a marsupium, the bursa, or rather

multiple burs, being no longer sufficient.

I think the new conditions to be faced by intense arboreal

life have acted somewhat differently, especially if, as we assume

now, all the Metatheria were already possessed of a marsupium”™.

I venture to suggest that those Metatheria which were driven

to arboreal life, had to solve the question what to do with

their young. They had to carry them, and being already pos-

sessed of the pouch, this was not only retained but intensified

by prolonged use. That the facilities offered by such a pouch

are great, is shown by the still existing numerous and much

diversified Marsupials, and by the fact that some at least, e. g.

Opossum, have reduced the uterine gestation almost to the

conceivably lowest limit. Whilst these arborealised Metatheria

could not afford to leave their young behind, it was different

with those other Metatheria which during the Cretaceous epoch

somewhere “‘in the larger and more effective workshops of the

North” (to apply here one of Darwin’s happiest expressions) went

* Klaatsch’s view of the correlation between Bursze and Marsupium had to be

considerably modified by Semon’s discovery that the Monotreme does not lie in a

bursa but in an already typical, though transient, marsupium. Moreover, there are

unmistakable vestiges of marsupial muscles, other than compressores mammz,

in various Placentals. Lastly, each teat with its areola does represent a bursa

(Klaatsch), but if each bursa had been a brood-pouch instead of merely the result of

the attachment of young, the possession of a series of bursz from near the vulva

up to the armpit would imply an impossible condition. In fact, bursa or nipples

can be formed independent of a marsupium and in places whereto no such organ

can have extended.

AND OVIDUCTS IN THE AMNIOTA, 831

in for that further mental development which is intimately

correlated with the origin of the corpus callosum. Competition is

most effective if backed by brain-power. Where these dawning

EKutheria spread the other Metatheria had to go to the wall, or

were forced to get out of reach—into the trees. Those which,

already arborealised, found their way into countries like Australia,

where the Eutheria could not follow in time, found there

nothing to prevent them from becoming again terrestrial, as

Dollo has shown so conclusively. The chance of reviving the

degenerated placenta was, however, gone, and the Marsupials are

now, in spite of the retention of various low characters, the most

specialised subclass of Mammals.

The Edentates present a striking analogous case: being with-

out exception arboreal or fossorial, they also seem to have got

out of the way of mentally higher, dangerous Eutheria. They

have survived, whilst those that went in for heavy bulk and

armour were not a great success. At least the American Eden-

tates seem to have come into contact with the dangerous

Kutheria long after they themselves had become typical Eutheria,

so that the former possession of a marsupium could avail them

nothing.

A few words about the Eutheria. They could afford to part

with their young for a time and to suckle them intermittently,

an advantage which needs no further comment than that it

would be impossible without higher mental faculties.

Great adaptive changes have been wrought ; every chance has

been tried, now here, then there. Nearly all the Non-deciduata

are at birth precocious, able to look after themselves, no matter

whether the placenta is diffuse, cotyledonary or zonary even,

namely the aquatic Cetacea and Sirenia, and the Artio- and

Perissodactyles, the exception being the Lemurs. The Deciduata

are at birth helpless: Rodents, Insectivores, Carnivores, Chiro-

ptera, Tarsius and Simiz, except the Elephants.

The egg-laying Monotremes, the implacental Marsupials, and

the Mammals with the most advanced placenta, are born in a

condition as helpless as the highest nidicolous birds, whilst the

avowedly low non-deciduous and diffuse placenta of the Mare i

sufficient to produce a young which is as precocious and in-

dependent as a lizard.

Can these facts be brought into line? What is here primitive

and what is a roundabout return to apparent but not really

ancestral conditions 2

Proc. Zoo, Soc.—1912, No. LY. 55

829> DR. F. E. BEDDARD ON A

49, Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea. By Frank H. Bepparp, M.A.,

D.Sc., F.R.S., F.Z.S., Prosector to the Society.

[Received April 30, 1912: Read June 4, 1912. ]

(Text-figures 113-121.)

VI. On an ASEXUAL TAPEWORM FROM THE RODENT, Siber zibethicus,

showing a new form of Asexual Propagation, and on the

supposed Sexual Form.

INDEX.

Page

Str chuve cece csecsen Uke coer sasaumee seca ees mereste te § ROaO

Systematic :

Urocystidium gemmiparum, gen. et sp. n....... 840

General Résumé and Systematic Position ............ 849

Anatomical Summary ..........:0ccecereeeeeeeeeeeeeeesss 850

T received in February of this year two complete tapeworms

which were found in the hepatic duct of a Musquash (fiber

zibethicus) which died in the Society’s Gardens. The two worms

lay together in a part of the hepatic duct just before it receives

the bile-duct, which was much dilated by the parasites. Although

these two worms were very different in appearance, I believe them

to be respectively the sexual and asexual form of the same species.

T shall give reasons for this conclusion in the course of the

following description of the two individuals.

(1) Tue Asexual Form.

The general appearance of this very remarkable worm is

shown in text-fig. 113 (p. 824), which illustrates its most

remarkable peculiarity, viz., the possession of two series of

what appear to be buds at one end of the body. It will be seen

in the course of my description of this “ Cysticercoid” that it is

without any doubt to be regarded as an asexual form. It does

not, however, follow that the mature worm found with it is a

further stage in its development. On the contrary, indeed; for

as a mere matter of guesswork, the assumption would be the other

way. Still, I believe that I shall be able to prove that the two

worms are stages of the same species, in which event we have the

very remarkable fact of both the sexual and the asexual form

coexisting in the same host, and, moreover, in a situation where

one would expect to meet with sexual forms only. The Hymeno-

lepis* of the mouse is a partly, but not an entirely, parallel

instance,

At first it seems likely, from an inspection of the figure

* AH, murina.

NEW ASEXUAL TAPEWORM, 823

annexed, that the wider end of the worm is the scolex and that

the narrower end is the posterior extremity where the proglottids

are being shed. The dilated extremity suggests a scolex not

altogether unlike that of the genus Dasyurotenia, which I have

lately described to this Society *, and which is characterised by

an unusually swelled scolex. Furthermore, there is a slight

diminution in the diameter of the worm towards the narrower

end, which again conforms to this view, which, as a matter of fact,

I at first held myself. It appears, however, that the reverse in

fact is the case, and that the narrower end is the scolex end.

The opposite extremity therefore is, as I suppose, to be compared

to the persisting bladder of the bladder-worm stage of the

Cestode. To the naked eye, and even on examination with a

lens, the scolex end presents every appearance of being the proli-

ferating end of the body, for the last segments appear to be

slightly incurved, as is so commonly the case at that end. It was

not until I studied this region by means of a series of horizontal

sections that I was able to discover its true relations. And even

now certain important details are wanting.

I could find, indeed, no armed scolex, nor any trace of suckers;

if the worm is a member of the Pseudophyllidia and has there-

fore only bothria, these may easily have escaped attention in such

sections, which would not be suitable for their display. I cut the

sections, in fact, under the impression that I was dealing with the

posterior end of the body, and without making a sufficiently ex-

haustive survey of the external characters. The main arguments,

therefore, which lead me to the conclusion that this is really the

scolex end are firstly the mode of imbrication of the proglottids,

and secondly the presence of large pigment granules, a condition

which would hardly be expected at the posterior end of the body,

but which is not uncommon among tapeworms in the scolex.

As to the imbrication of the proglottids it seems to me to be

necessary to regard a segment which overlaps the next one as

being anterior to it in point of origin, and therefore lying to the

scolex side of it. Judged by this conclusion, the narrower end of

the body of this very remarkable tapeworm is the scolex end.

There is no evidence that a scolex hag been lost. On the

contrary, the body ends here in a slight median elevation, which

is quite unlike the termination of the body were this the region

of the detachment of proglottids. This little elevation, however,

bears no particular likeness to a scolex, and there are certainly no

suckers or hooks to be seen anywhere. Nor is there any neck, or

break of any kind, between this region and the first obvious pro-

glottid. In this latter, moreover, the lateral and transverse tubes

of the water vascular system are as large as in the more posterior

segments, and do not end in a coil such as is so frequent in the

anterior part of this system in other worms. It may be, of course,

that the scolex is in this genus a transitory affair, as it has been

* P,Z.S. 1912, p. 677.

La:

824 - DR. F. E, BEDDARD ON A

believed to be (though perhaps hardly now) in certain other tape-

worms. Or, on the contrary, we may have here a feebly developed

head like that of Zigula, But the great breadth of the body of

the present worm is not perhaps in favour of such a supposition.

Had there been a very narrow anterior neck, some suggestion of

this kind might have been put forward. Anyhow, the general

characters of the anterior region of the body of this Cestode are

as described above. We shall recur to their more minute anatomy

Text-fig. 113.

The upper figure represents the entire asexual form of Uroeystidinm gemmiparum

enlarged by about one-third. The lower figure is the posterior end of the same

individual more magnified. For explanation see text.

later. From this point to the extreme posterior end are very

many proglottids. They are very short and wide, and do not

appear to differ very appreciably in length in different regions of

the body. The greatest diameter is something like 4mm. This

is at and towards the posterior end of the body. Quite anteriorly,

the diameter is not more than 2°55 mm. The length of the entire

worm is about 80 mm.; the small posterior bladder is 8 mm. long

NEW ASEXUAL TAPEWORM. 825

by 2°5-3 mm. wide. The proliferating end of the worm ceases

abruptly with the commencement of the bladder. This region,

which bears a great many lateral buds, shown in the text-figure

referred to, measures 17 mm. The region of the body which

bears these presumed buds (we shall consider their nature later)

does not differ much from the preceding section. It looks,

perhaps, a little more transparent, but it is divided up exactly in

the same way into proglottids, which are of about the same

dimensions as those elsewhere in the body. The buds form a row

on each side of the body and are quite lateral in position ; they

are not at all arranged in order of growth, though representing

very many stages. That is to say, the older buds alternate with

younger and older forms. Nor does every proglottid bear a bud

or pair of buds (one on each side), Between any two buds there

was often a variable number of proglottids without any trace of

budding at all. This, indeed, is necessary; for the large size of

the buds would prevent their proper growth upon immediately

adjacent proglottids. I counted altogether 17 buds upon one

side and 18 upon the other. But I may have omitted one or two

in each case; for it is a little difficult to fix the actual first

appearance of a bud. ‘They begin, in fact, as an only just recog-

nisable rounded swelling of the edge of a proglottid. Sometimes

the swelling includes two proglottids. There is no question

whatever of the continuity of these buds (as I regard them) with

the parent stock; they are most plainly outgrowths therefrom.

In later stages the minute round bosses at the edges of the

segments swell into spherical, largely transparent, vesicles, of

which, as the text-figure (text-fig. 113) shows, the size varies—

possibly according toage. Later still, these bladders show a young

worm growing out from their distal extremity, which is usually

of considerably less diameter than the bladder which is attached

to the parent stock. It exhibits numerous wrinkles which I do

not definitely regard as the delimitations of proglottids. The

longest of these presumed young worms was about 4 mm. in

length; the bladders reached a length of 2 mm. The exact

number of these more developed buds is faithfully reproduced in

text-fig. 113.

§ Structure of the Parent Stock.

I have investigated the anatomy of the worm by transverse,

longitudinal, and sagittal sections. From an inspection of the

former (see text-fig. 114, p. 826) the depressed form of the body

. was obvious, the diameter of a section being about seven times

its depth.

At the thinner anterior end of the body, not far from the actual

extremity, transverse sections showed that the medullary region

of the proglottid was not more than two-thirds of the diameter of

the cortical region. The two were plainly marked off from each

other by delicate transverse fibres forming a very thin layer, and

826 DR. F. E. BEDDARD ON A

yet quite unmistakable. The same proportions and the same

structure were visible as plainly in the wider posterior region of

Text-fig. 114.

Part of a transverse section through a proglottid of the asexual form of

Urocystidium gemmiparum.

A, spaces in middle of proglottid with darkly staining walls referred to in text.

Um. Longitudinal muscular layer of cortex, within which is seen delicate

transverse layer of muscles. .s. Nerve-cord. v.d. Dorsal excretory tube,

v.v. Ventral excretory tube.

the body. The medullary substance is quite well stained by hema-

toxylin and has the usual homogeneous appearance. There is,

NEW ASEXUAL TAPEWORM. 827

moreover, in the present worm the usual meshwork arrangement

of fibrils round spherical masses of the homogeneous ground-

substance such as occurs in so many tapeworms, and is often a

very coarse and thus more obvious network. The homogeneous

ground-substance was, however, traversed by two sets of fibres,

which are, I presume, muscular fibres. These are absolutely at

right angles to each other and not at all closely set in either case.

The dorso-ventrally running fibres are nearly always wavy in

their course, the transverse fibres are quite straight—a matter of

different states of contraction, as I imagine.

The cortical layer is easily divisible into two regions, which ar

of about equal diameter. The innermost of these is that occupied

by the bundles of longitudinal muscular fibres. The bundles of

muscular fibres are two or sometimes three deep, and each bundle

contains a large number of fibres, which, however, are not very

closely pressed together. In addition to these bundles of rather

slender fibres there lie on either side, between the nerve-cord and

the lateral margin of the proglottid, two bundles of considerably

stouter fibres which run continuously from segment to segment

and are closely associated with a longitudinally running cavity.

These bundles appear on occasions to lie actually within the cavity.

This tubular cavity is not at all like the tubes of the water

vascular system, and it lies within a very lax tissue. Iam unable

at present to suggest its nature.

The accompanying text-figure (text-fig. 115, p. 828) illustrates a

section through the hinder part of the worm not very far in front

of the budding region. This region shows very plainly a system

of irregular spaces in various parts of the body belonging to the

water vascular complex. It may be remarked first of all that the

dorso-ventral diameter is greater here than in the more anterior

part of the body. The structure of the vertical and medullary

regions is the same, but the lower central region of the medulla

actually forms a cavity (text-fig. 114 A) which I do not think to

be artifact and which may be a forward extension of the cavity

of the terminal bladder, though I have not proved the fact, from

an unwillingness to sacrifice the specimen altogether. The in-

terior lining of the space was, however, so very strongly stained

as compared with the surrounding tissues that it appears to me to

be practically certain that the staining fluid gained access through

the two cut ends of the piece of worm, which was stained en bloc.

This, of course, argues a continuous central cavity.

It was quite plain that this cavity was quite distinct from that

of the transverse water vascular trunk. The peculiar central

cavities occupying nearly the middle of a segment here and there,

referred to later (see p. 830) in describing sections made from

earlier segments in the body, are present here and are again not

to be confused with the central lacuna. Their thick walls show

no trace of an opening into it. In addition to the main trunks

of the water vascular system, which are disposed here as they are

elsewhere in the body, there are scattered irregular spaces which

828 DR, EF. BE, BEDDARD ON A

seem to belong to the same system, These consist of larger but

very short stems with finer branches. I have not traced any con-

nection between these tubes and the main stems of the excretory

system; but I can, nevertheless, hardly doubt that it exists.

Text-fig. 115,

oY.

A move highly magnified section through excretory vessels and adjacent:

structures of the asexual form of Uroeystidiwm gemmiparun.

dv. Dorsal vessels surrounded by circular muscle-fibre. mm. Layer of transverse

muscle-fibres separating the cortex (to the right) from the medulla (to the

loft), 2». One of several tubes forming the excretory network and lying, as is

shown, in the cortex as well as in the medulla, é. ‘Transverse commissural

vossol forking to join the ventral vessel (v.v.), which is also bifurcate to receive

the branches,

These ramifications lie in the cortical layer as well as in the

medullary (text-fig. 115, 2). In one case I observed a narrow

duct leading from a wider space to the exterior—at any rate, most

ule

NEW ASEXUAL TAPEWORM, 829

undoubtedly to the cuticle, and in other cases it appeared to me

to be highly probable that the ramifications of these tubes do

open on to the exterior here and there; they at least invade the

outermost layer of the cortical tissues.

The water vascular tubes ave large in this species, and not far

from being equisized. There are, as is the rule, two upon each

side of the body. As is so often the case among the Cestodes, the

two tubes, instead of being respectively dorsal and ventral and

superposed, lie side by side, As is also quite usual among these

worms, the two tubes, dorsal and ventral, differ somewhat in

structure. | consider that the slightly smaller and more muscular

of the two tubes is the dorsal vessel, from the analogy of other

forms; in the present species, therefore, the dorsal vessel is

situated internally to the ventral. They are, however, quite in

the same straight line for the most part and also in the same

straight line with the nerve-cord, But, as a matter of fact, the

more muscular dorsal tube lies in a corkscrew fashion, which is

very plain when the proglottids are viewed in longitudinal and

sagittal sections, and thus that tube in cross-sections appears to

lie now rather dorsal of, and at times rather ventrally of, the

ventral and more external vessel. Elsewhere they are. more

directly in the same straight line. It is a remarkable fact that

both of these vessels seem to be quite as large or very nearly so

in the most anterior segments of the body, There is no great

increase in calibre posteriorly.

The general relations of these two vessels are shown in text-

fig. 114, ‘Text-tig. 115 is a more enlarged representation of the

two tubes and the adjacent parts taken from a section of the

posterior region of the body. ‘Their detailed structure is there

illustrated, In his account of the Cestoidea, in Bronn’s ‘ Thier-

reichs,’ Prof. Braun remarks—‘‘Ob in der Wand der Sammel-

rohren auch noch Muskelfaseren vorkommen, ist noch strittig.”

The figure just referred to is, as I think, quite conclusive as to the

presence of a particularly thick layer of civeular fibres constituting

the greater part of the wall of what I regard as the dorsal vessel.

This muscular layer appears to wax and wane in thickness from

place to place. But it is always thick and thus very apparent,

The actual tube is lined with a chitinous cuticle which is also often

very thick and everywhere to be seen, Outside of the layer of

circular muscles are abundant nuclei, massed round the tube, 1

did not observe any nuclei among the circular muscular fibres,

which are, indeed, very closely contiguous. There appear to be

no intrinsic longitudinal muscles to assist in the widening of the

tube again after contraction by the circular muscles; but there is

a mechanism which takes the place of such fibres to serve in their

stead.

In the neighbourhood of the dorsal excretory vessel the dorso-

ventral muscles form groups here and there of rather thicker

and more closely adpressed fibres, which, as it were, tie to or

suspend the dorsal vessel from the cortical layer ; it seems certain

830 DR. F. E. BEDDARD ON A

that the contraction of these would dilate the lumen of the dorsal

excretory tube. These fibres, which exist both dorsally and

ventrally, have a somewhat fan-like arrangement—that is to say,

they converge upon the vessel from above and from below. It is

these various muscular layers which cause so great a variation in

the lumen of the excretory tube from place to place.

The ventral vessel differs entirely from the dorsal by the entire

absence of any muscular layers and by the possession of only a

very thin lining cuticle. It follows, therefore, that the variations

in the calibre of this tube are not so great, and when they do

occur seem to be due to a squeezing by the surrounding tissues.

The ventral vessel also runs a much straighter course than the

dorsal without any marked corkscrew-like windings exhibited by

the latter. There is a transverse vessel in each segment which

has a rather peculiar relation to the ventral longitudinal tube

into which it opens on either side of the body. A little way before

reaching the dorsal tube the transverse vessel splits into two

halves, which embrace the dorsal vessel and open separately into

the ventral vessel. This is partly shown in text-fig. 115.

‘We have now to deal with a series of sac-like bodies forming

closed cavities of very problematical nature, which lie in the

middle of many segments. There is by no means one of these

sacs to each segment, and thus they are very far from consti-

tuting a tube running without intermission through the body.

I compare them, however, later in this paper to a tube found in

the young buds (see p. 833). But, as a matter of fact, one of

these sacs may actually extend through two proglottids, so that

there is a hint of a formerly continuous structure such as occurs

in the buds. Asa rule, however, they appear to be limited to a

single proglottid, of which they occupy sometimes the exact centre,

at other times pushed rather to one side. They lie close to the

transverse water vascular vessel. I cannot find that these sacs

have any connection with any other spaces. They appear to be

perfectly isolated. The first of the series occurs very early in

the body—if not in the very first segment. These sacs have a

lining cuticle surrounded by a single layer of epithelial cells of

which the borders between the individual cells are not clear.

The nuclei, however, are very obvious and large. There are no

muscle-fibres at all apparent encircling the sacs. They often

appear crumpled in transverse sections, and occasionally seem

to be Y-shaped. They are impossible to miss in a series of

sections.

The only remaining characteristic of this worm upon which I

am able to report is the structure of the ‘“‘ head” end of the body,

which has been already briefly described above. In the place of

a scolex there is a small projecting process, no larger than the

rostellar process of many unarmed tapeworms. ‘This arises from

the middle of what may be termed, for the present, the first seg-

ment of the body, which is distinguished from those that follow

by its rather less breadth and greater length. It would seem to

NEW ASEXUAL TAPEWORM. 831

be really composed of two segments, if we consider the condition

of the water vascular system therein. The two main longitudinal

vessels of this system extend well forwards into this compound

segment, and the outer ventral vessels are connected by two

transverse vessels, thus indicating, as I suggest, the double

nature of this apparently single segment. Towards the centre

of the specialised anterior segment is a large group of deeply-

pigmented oval bodies, which appear to be special pigment-cells.

They are as large, or nearly so, as the calcareous corpuscles which

are very abundant in this worm. In the middle of the concave

anterior border of these two fused segments is quite a small

conical projection, than which nothing else at all comparable to

a scolex exists. There is no sign of any breakage at the summit

of this which might suggest that a scolex had been accidentally

detached.

It is a somewhat remarkable fact that these conditions (or, at

any rate, something very near to them) occur in the genus

Dioicocestus; for, as will be pointed out later, the genus with

which I am concerned in the present paper shows some likeness

to the Acoleide and even to the genus Diotcocestus, which is of

that family. In Dioicocestus acotylus, Fuhrmann * has described

and figured the practical absence of suckers and the very rudi-

mentary condition of the rostellum—a state of affairs which is

not found in the other species of this genus. The figure referred

to also shows—as I have described in my species—the absence

of any marked diminution in calibre of the body at the head end.

But it must be remembered that the Dioicocestus was a fully-

developed sexual worm.

§ Structure of the Buds.

It now remains to consider the minute anatomy of these out-

growths and their nature. As to the latter query, there are, as

it appears to me, only two alternatives. These outgrowths must

be either “tentacles” of the nature of the processes known in

Hymenolepis villosus, or young worms budded off from the

parent stock. There is very little, if indeed anything, to be

said in favour of the former view. It is true that a large out-

growth of the body to form a tentacle might have very much the

same structure as the body, even to possessing branches of the

excretory system and lateral nerve-cords. But there would

hardly be a practical identity of structure such as I shall point

out in detail later. Furthermore, the growth of processes of the

body might be expected to be more regular than are these out-

growths (ef, text-fig. 113), and above all a gradual freeing of

themselves from firm contact with the body, as shown in the

figure, is precisely what we should expect with budding offspring,

which, as it appears to me, is the obvious and only way to in-

terpret these appendages of the fully grown worm.

* Zool. Jahrb., Abth. f. Syst. Bd. xx. 1904, p. 181.

832 DR. F. E. BEDDARD ON A

I selected for investigation, by means of transverse sections,

the most adult-looking of the young budded-off worms, in which,

moreover, segmentation was obvious. I have already referred to

this individual in my account of the budding process and need

not redescribe its external characters here. The wider end

attached to the body of the parent forms a bladder with rather

thick walls. This I take, therefore, to be the posterior end of

the young worm. The opposite end is very much thinner (as is,

of course, usual among tapeworms) and represents the scolex end

of the animal. I could observe no trace of hooks anywhere on

this scolex, nor suckers of any kind. This, however, is not sur-

prising in view of what I have already described in considering

the conditions observable in the parent stock. At about the

middle of the worm a transverse section shows the appearances

represented in text-fig. 116. The cortical layer is thick—about

the same diameter, or in places rather more than the diameter,

of the medullary region. The medullary region is plainly marked

off from the cortical by transverse muscular fibres of delicate

constitution.

There are several parallel fibres in this layer, which is altogether

of some width, the individual fibres being widely separated. The

parenchyma of the medullary region is as usual, and contains

many nucleated cells, generally triangular in form, connected

with processes at the angles. These cells get to be particularly

numerous near the water vascular tubes, which will be referred

to later. There are also heaps of nuclei visible in some sections,

which I take to be the rudiments of the sexual organs. To one

side of the body the medullary region seems at first sight to be

invaded, as it were, by the cortical region, thus producing an

asymmetry in the transverse section, which is quite apparent in

the accompanying text-figure. This asymmetry is associated with

the presence of a tubular organ (text-fig. 116, a:), which will be

described in detail on a subsequent page. This occupies a good

deal of space, and round it are large bundles of stout muscular

fibres, quite similar to those which occur in the cortical layer.

It is very easy, however, to observe that the transverse layer of

muscles bounding the medullary parenchyma is continued on

either side of this differentiated mass of tissue and ends laterally

at a point asymmetrical with its ending on the other side of the

body. I therefore regard this region as belonging to the medullary

part of the body.’ The cortical region is distinguished from the

medullary (other than that section of the medullary region just

referred to) by the bundles of stout longitudinally running fibres.

These bundles form a layer immediately outside of the transverse

fibres; but there are much smaller bundles and isolated longi-

tudinal fibres extending some way towards the exterior of the

body.

In the medullary region lie the water vascular tubes and the

nervous system. All of these lie in the same straight line with

each other and with the tubular organ already referred to, and

NEW ASEXUAL TAPEWORM. 833

this line is the longer diameter of the cross-section ; the tubes

ete, are thus laterally disposed with regard to each other. It is

necessary to emphasise what may appear to be a statement of the

obviously true, since in other regions of the body the mutual

relationships of these several organs and systems are a little

changed.

Text-fig. 116,

Transverse section through middle of strobila of oldest bud of the asexual

form of Urocystidium gemmiparum.

lv. Excretory tubes. 2. Nerve-cord. «. Tube of doubtful significance

described in text.

In all other tapeworms known to me the water vascular tubes

of the two sides of the body are symmetrical with each other in

point of position within the medulla and distance from the lateral

cortex on either side, They are also generally correspondent in

size, though not always regularly so throughout the whole body.

In the present species the water vascular tubes are asymmetrical in

both these particulars. The tube of the one side is considerably

834 DR. F. E. BEDDARD ON A

larger than that of the other and is relatively asymmetrically

placed. This latter fact is doubtless due to the bundles of longi-

tudinal fibres surrounding the large lateral tubular organ. This

structure has, as it were, pushed the nerve-cord and the water

vascular tube of its side towards the centre of the body. Indeed,

the vessel in question lies very nearly in the actual centre of the

section and quite in this position shortly before and during its

connection by the transverse vessel with the tube of the opposite

side of the body. These transverse vessels are at regular intervals

and quite easy to see as in the adult worm. I did not observe

any other branches of the water vascular tubes than these.

These sections from the middle of the body of the young worm

also show very plainly the remarkable constitution of the nervous

system in this Cestode. One of the two lateral cords (text-

fig. 116, 2) is very obvious indeed, lying between the water vascular

tube of that side and the tubular organ of doubtful meaning.

It cannot possibly be missed in these sections. But on the opposite

side of the body I could find no trace whatever of any nerve-cord

even of smaller size than the one represented in the text-figure.

T feel confident that a failure to observe a cord in this situation,

were it present, would be difficult. The medullary tissue is so

clearly differentiated by the stain (iron hematoxylin*) that

the nerve-cord, if of any size at all comparable to that of the

opposite side of the body, would stand out quite prominently.

This state of affairs is undoubtedly very anomalous and hard of

explanation. For we shall see later that the two nerve-cords are

both quite recognisable towards the end of the body.

Finally, this transverse section shows to one side a large tube

cut across, to which reference has been made incidentally once or

twice already. It is situated in the centre of a group of bundles

of longitudinal fibres, which bundles are like those in the cortical

layer of the body in every way, including their rather sparse

scattering round the centrally placed tube. The bundles in the

cortical layer are not densely pressed together as they are, for

example, in Dasyurotenia robusta t. The tube itself is of con-

siderable size and is lined by a thick layer of chitin, which is

stained precisely like the chitin which covers the body of the

worm externally. Outside of this is a single layer of stout longi-

tudinal muscle-fibres, and then a deeply staining layer of elongated

cells, interspersed among which are small bundles of stout longi-

tudinal fibres. The structure of the tube is in fact almost exactly

that of the outer layer of the body, only differing, indeed, by the

intercalation of the groups of stout longitudinal fibres among

the cells.

We shall now consider the course and structure of this tubular

organ throughout the body of the young worm. The identity of

structure with the outer layer of the body which this tube

shows in the middle region of the body, is rendered intelligible

* T am indebted to Dr. Plimmer, F.R.S., for the use of this reagent.

+ See Beddard, P. Z.S. 1912, p. 684, text-fig. 97, lm.

“4 NEW ASEXUAL TAPEWORM. 835

by the fact that in two places, one behind the other, it communi-

cates with the exterior by a narrow duct (text-fig. 117). One of

Text-fig. 117,

Two consecutive sections from another region of the same bud as that which is

represented in text-figure 116, to illustrate the opening of the tube x on tothe

exterior.

In the upper figure the orifice (o.) on to the exterior is shown and the

commencement of the invagination leading towards the tube x.

In the lower figure the outgrowth of the tube # to meet the invagination is seen.

n, Nerve-cord. JU.v. Water vascular tubes,

836 DR, F, E. BEDDARD ON A

these openings is in the bladder region and the anterior one a

very little way in front of this. Both orifices are dorsal or ventral*

in position—at any rate not lateral,—but they do not coincide

exactly. In view of the position of these orifices it is clear that

the tube of communication is of some length. In addition to

these two openings on to the external surface, the tube gives off

two short cecal processes, one of which nearly but not quite

opens into the bladder.

- Anteriorly the tube gradually comes to an end and does not

end by opening on to the exterior. It diminishes in calibre for

the space of a few sections and then simply ends. At the bladder

end of the body—-the end which is attached to the parent stock—

the tube gets to be more and more shoved to the side as the

bladder increases in size. In consequence also of this the tube

becomes flattened from side to side, at the anterior part of the

bladder. Followed backwards this tube can be recognised by its

thick chitinous lining, already referred to, and can be seen

thereby to be quite distinct from the excretory tube which has

come to lie below it in consequence also of the development of

the bladder region. The next that occurs is that the tube divides

into two coincidently with the formation of two or three septa,

partly dividing up the cavity of that part of the bladder which

lies adjacent. The two tubes lie one dorsally to the other, and

thus both of them in the same straight line with the excretory

tube. The middle tube of the two after a very short course

appears to open into one of the chambers of the bladder, and is

thus a diverticulum like the one mentioned above. Furthermore,

I am not absolutely certain that it actually opens into the

bladder ; but it ends at least in close contact.

I feel, indeed, almost inclined to assert that this tube does

open into the cavity of the bladder, since the main tube, continu-

ing a little way further back, gets very narrow and undoubtedly

ends by opening into the bladder. It is to be noted that these

orifices do not involve a mere continuity of lumen. The character

of the lining membranes changes at the point of meeting. It

is further to be noted that it is not the main cavity of the

bladder into which the tube opens, but into a portion of it,

separated off by septa. This problematical tube varies in parts

in its structure and in its relations to the nerve-cord. Anteriorly

the layer of cells surrounding it are not so conspicuous as

elsewhere, and thus the longitudinal muscular fibres come into

greater prominence; the calibre of the tube also is not the same

all through the body. It has been mentioned that in a section

from about the middle of the body, that the tube lies outside

of the nerve-cord and quite laterally to it. In other parts of the

body, particularly anteriorly, the tube lies above (or, perhaps it

may prove necessary to say, below fT) the nerve-cord.

We have, therefore, in these young worms still attached to the

* T cannot differentiate between the dorsal and ventral surfaces.

+ See footnote above.

NEW ASEXUAL 'TAPEWORM. 837

parent, though (as the narrow stalk shows) just ready for detach-

ment, a tube of the structure of the outer layer of the body

which traverses in a straight line nearly the whole of the body,

which opens into the bladder behind and has two segmentally-

arranged orifices of communication with the external world.

There is no obvious answer to the question as to its nature. In

buds within the cyst of Hchinococcus* a cavity is formed which

is not an ingrowth of the cavity of the whole cyst, but which

is alleged to be continuous up to the scolex. This, however,

has been denied and the scolex asserted to be entirely solid. But

even supposing a comparison is possible on the grounds that

the scolex of UYchinococcus is hollow, the short tubes leading to

the exterior in the worm of which the present paper contains

an account would seem to invalidate it. And, moreover, the

structure of this problematical tube is* not that of the bladder

with which it communicates. It seems, indeed, to be a special

structure and not a space connected in any way (or, rather, any

direct way) with an evagination of the scolex.

For the greater part of the body I could only find a nerve-cord

on one side, where it was quite as obvious as in the “adult” worm.

In the anterior region, however, the nerve-cord of the opposite

side was visible for a short space. It was, however, not half the

size of its fellow, though of precisely the same structure and

equally unmistakable as a nerve-cord. Both cords lie in the

medullary region just at its bordering on the cortical layer. I could

find no enlargement of the nature of a brain. Quite anteriorly, as

well as posteriorly in the bladder region, I could not recognise a

nerve-cord. It is noteworthy that the asymmetry of the two

water vascular tubes in size is exactly paralleled by that of the

two nerve-cords.

These are the facts which I have ascertained in the anatomy of

the most mature of the growing buds. It now remains to compare

the structure of the young worm with that of the parent stock of

which it is a bud,

§ Comparison of Bud with Parent Worm.

In the general form of the body and in the possession of a

relatively small bladder they agree; also the rudimentary scolex

is a marked feature of both young and old.

It is rather remarkable to find asymmetry in the young

worm, but this of course may be its normal method of growth.

Certainly I noticed no asymmetry in the parent form. What

is perhaps important as a difference is the presence of only two

water vascular tubes in the young and four of these in the

older worm. The additional water vascular tubes may, however,

make their appearance later. Furthermore, for the same

reason, too much stress cannot be laid upon the more copious

* Bronn’s Thierreichs, iv. Abth, 1B, p. 1549.

Proc. Zoou. Soc.—1912, No. LVI. 56

838 DR. F. E. BEDDARD ON A

bundles of longitudinal muscular fibres which are to be noted in

the large worm. I believe also that the continuous large tube in

the immature worm is represented by the cavities contained in

many of the segments of the adult worm, though the minute

structure, it must be confessed, differs in the two cases. Further-

more, the series of cavities in the adult worm lie very nearly in

the middle of the body, indeed quite in the middle of the body in

some segments; the tube in the young worm is, on the other hand,

as distinctly to one and the same side. But it must be borne in

mind that the position does vary in the young worm, and a further

flattening of its body might easily cause a greater similarity m

the position of the tube to that observable in the more fully

mature worm.

S$ Comparison with other Forms and Systematic

Position of the Parasite.

Apart from the question of proliferation by budding, to which

we shall return later, this Tapeworm presents a certain number

of undoubted resemblances to Cysticercus fasciolaris (of rats and

mice, ete.), which becomes Z'cenia crassicollis in the Cat. Both

these forms agree (I have compared the species which forms the

subject of the present paper with examples of the Cysticercws from

the common rat) in possessing a long segmented body and a small

bladder posteriorly. But in Cysticercws fasciolaris the bladder is

smaller and the body shorter than in my species, while the hooked

and suckered anterior end renders any confusion or detailed

comparison impossible. Nevertheless, the two forms have in

common the small bladder and long strobila. But while the one

occupies the position in the body of its host of asexual worm, ¢. e.

in a diverticulum of the alimentary tract, the other is found, as

are Oysticerci, encysted, and in the liver of its intermediate host.

Nor is it by any means certain that the species from the Mus-

quash isa member of the Teenioidea (Cyclophyllidea) at all. It

may well be a member of one of the lower groups of Cestodes.

We shall consider the arguments for and against the placing of

the worm among the 'Tenioidea.

Tt must be admitted at once that there are no absolutely con-

clusive arguments which point definitely one way or the other.

This, of course, may be explained on the assumption that we are

dealing here with quite a new type of Cestode. In the meantime,

the structure of the worm so far as it can be read does not favour

such an assumption, though it does not, for the matter of that,

appear to be necessarily contradictory of this possible view. As to

the other alternatives, the absence of a marked scolex with suckers

is greatly against the reference of the worm to the Teenias, but

the character of the terminal bladder, on the other hand, is on the

whole in favour of such a placing. This latter fact is obviously

against the supposition that the worm is a Plerocercoid of any

kind; but the lack of a definite “head” is as clearly in favour of

NEW ASEXUAL TAPEWORM, 839

this supposition. There, as it appears to me, the question must be

left—in regrettable uncertainty.

The process of asexual reproduction by budding which this

worm shows is not entirely unknown among the Cestoidea, but

there are some not unimportant differences from what has been

recorded in other forms. There are two principal comparisons to

be made.

The first of these is with Cysticercus longicollis, the bladder-

worm of Venia crassiceps. It is of further interest from the

point of view of the present comparison to note that this Cysticercus

is also found in a Rodent, Arvicola arvalis. Besides the earlier

investigators, Prof. M. Braun* has studied this form and em-

bodied his results in brief in Bronn’s ‘ Thierreichs’+. ‘The buds

from the Cysticerci do not apparently {contain a prolongation of

the bladder-cavity, ‘sondern durchweg als solide Wucherungen

der peripheren Schicht der Wand entstehen.” This is an obvious

point of similarity with my species, where, of course, the buds

cannot be continuous with the cavity of the bladder—that is to

say, in both cases the buds are solid outgrowths. Furthermore,

before being separated off from the parent Cysticercus the attach-

ment of the bud dwindles to a narrow stalk precisely as I have

described above. The buds, however, in the case of Cysticercus

longicollis seem to be limited to the bladder and always to the

hinder end of that; whereas in my species this is exactly the

region where no buds are formed.

I believe, however, that a nearer approximation to the condition

observable in the species which I describe in the present paper is

offered by a worm recently described by Ijima + with some detail.

This is a Plerocercus or Plerocercoid found parasitic in a human

being in Japan in cysts in the skin. It appears to give off actual

buds, which are, however, more or less irregularly arranged and

present nothing of the comparative symmetry manifested in the

species from Miber zibethicus. The individuals differ in the degree

of their budding, some giving off a large series of slender pro-

cesses. These buds are, as in my species, actual outgrowths of the

parent stock and not, for instance, connected with the excretory

organs or any other definite part of the body. They occur

as much at the head end as elsewhere and there differ from the

Tapeworm of Liber zibethicus. There is, however, a resemblance,

in that in both cases the more mature buds are not in front of

or behind the less mature: there is a complete irregularity in

their order of succession.

The similarity between the two cases of budding cannot, how-

ever, be carried into any detail, since it is obvious that the two

worms are not nearly allied. ‘The Plerocercoid described by

* See Centralbl. f. Bakt. u. Par. xx. 1896, p. 580; Zool. Anz. 1896, No. 514; ibid.

1897, No. 521. ‘These papers are not illustrated.

+ Bd. iv. Abth. 1B, p. 1529.

ft “On a new Cestode Larva parasitic in Man,” Journ. Coll. Sci. Japan, xx. 1905.

For a reference to this paper I am indebted to Dr. W. Nicoll.

56*

840 DR. F. E. BUDDARD ON A

Tjima is regarded by him—and, as I think, rightly—as a young

Bothriocephalid whose larval stages are also known from similar

positions in other mammals than man. ‘These larva have no

bladder, and thus the fact that in the Plerocercoid, as in my

worm, the buds are produced from the general body surface loses

its principal significance. It is, I think, more important to note

that the worm from /ber sibethicus differs from Cysticercus

longicollis, in that the budding is not upon the bladder end of the

worm, It is remarkable, however, that both in my worm and in

the Plerocercoid there is no recognisable scolex ; but at the same

time there is in the supposed young Bothriocephalid no proper

segmentation of the body as yet visible. It is thus particularly

remarkable that my species is very fully segmented throughout.

I think, therefore, that the case of budding which I bring forw ard

in the present paper may be regarded as in some respects a new

form of asexual reproduction which i is partly paralleled in a few

other forms of Tapeworms.

Inasmuch as the propagation of this worm is different in detail

from anything that is known, and as the worm itself does not

entirely conform with any known species or genus, it is desirable,

as I think, for mere convenience sake to give it a name and to

define as far as may be the characters of the Cestode, Had the

worm proved definitely a ‘Tetracotylean*, I should have con-

tented myself with veferri ing it temporarily to the genus “ Z'enia,”

a name which is generally given to forms whose generic aftinities

are uncertain. As, however, this cannot be done with any

certainty, I venture to give a new generic name which is non-

committal as to its position in the series, and I term it

accordingly

Urocystidium gemmiparum, gen, et sp. n.

Incompletely mature worm with small bladder and very numerous

proglottids, Scolew feebly developed and without hooks. Tubes of

water vascular system lateral to each other, the dorsal with thick

muscular coat and inside of ventral, the panties connected by a trans-

verse vessel which forks and surrounds dorsal vessel ; @ network pr sent

in corte as well as in medullar 7] region. Longitudinal muscle-

layer wn cortex of at least two rows of bundles with rumerous fibres

in each. Buds formed at hinder end of body on both sides, thus

forming two rows, of which the bladders are attached to the parent

stock ; the scolex is not inverted, is without hooks, and rudimentary.

Hab. Musquash (/iber zibethicus), in liver-duct.

(2) Tas Srxvan Form.

As has been already mentioned, the hepatic duct contained

besides the ‘“ cysticercoid” just described a sexual worm which I

__* I think it necessary to maintain an attitude of reserve with respect to the

identity of the asexual and sexual forms.

NEW ASEXUAL TAPEWORM., 841

believe to be the mature form of the same Tapeworm. ‘The size of

the two worms was about the same. The sexual form measured

86 mm. in length and the greatest diameter, which occurred not

Text-fig. 118.

The sexual form of Urocystidium gemmiparwm (?).

The left-hand figure represents the entire worm magnified about twice; the right-

hand figure represents the scolex with a double crown of hooks more highly

magnified,

far behind the head, was6 mm, The appearance of the worm

(text-fig. 118) was quite similar to that of the immature form, It

842 DR. F. E. BEDDARD ON A

was posteriorly of a rather delicate appearance, rather translucent,

and the proglottids were very short and enormously wide in

proportion. There was no increase in length of the proglottids

towards the end of the body, and their proportions were as in the

asexual worm. It would hardly, of course, be sufficient to insist

upon an identity upon these grounds alone; but, taken in con-

junction with other facts which will be dealt with, the similarity

in outward appearance is very striking. The “tail” end of the

body appeared to be a little excavated, as is common in tapeworms,

the penultimate segments slightly embracing the last segment,

which was not quite so wide. I can believe that the worm was

proliferating at this end.

The scolex was not quite so wide as the ensuing strobila, and no

appreciable neck separated the two. The scolex is proportionately

large, as is indicated in the accompanying text-figure (text-fig.

118), and well armed anteriorly with two rows of hooks alter-

nating in position. The anterior circle of hooks consisted of 16

separate hooks, which were about twice the size of those of the

succeeding circle, whose number I did not count, but which were

presumably the same, as they were implanted between the larger

hooks. The usual four suckers are present; they show no un-

usual features and are unarmed ; their cavity looks forwards.

I have investigated the internal structure by means of trans-

verse and sagittal sections. The cortical layer does not differ

greatly from the medullary layer in thickness, and the general

appearance of the sections is very like that of sections of the

immature worm (see text-fig. 119). The cortex, for example, is

identical or nearly so. The same bundles of longitudinal fibres

occur and are very much of the same thickness. They are also

separated from the medulla by transversely running fibres. I

have described and figured in the asexual form the bundle or

bundles of rather stouter longitudinal fibres running outside of

the nerve-cord on either side and associated with a cavity dubiously

related to the excretory system. I find the same arrangement in

the sexual worm,

The excretory system does not needa very long description,

since it agrees in its main peculiarities with that of the supposed

asexual form. There are, in fact, the same two lateral vessels on

either side lying parallel to each other. They are, moreover,

roughly equisized, and the innermost of the two has very thick

muscular walls, the fibres being circular in their disposition. I

should add that nuclei interspersed among these fibres were very

obvious. In addition to these two longitudinal trunks each pro-

glottid possesses a transverse vessel which has the same remarkable

mode of union with the ventral excretory tube that I have figured

(see text-fig. 115) and described (see p. 830) in the presumed

Cysticercoid stage, and which I need not redescribe here as the

structure seems to be identical. There is, however, one important

difference which the sexual form shows from the Cysticercoid ; and

that is the absence in the former of the peripheral water vascular

NEW ASEXUAL TAPEWORM, 843

network of the latter. I could find no trace of this in the sexual

worm, although it was easy to see in the supposed Cysticercoid of

the same. It seems to me possible (perhaps necessary) to explain

this difference by assuming in the Cysticercoid a retention and

gradual metamorphosis of the bladder into the strobila, together

with its excretory system, which latter ultimately disappears on

the assumption of sexual characters. There are, of course, other

Cestodes in which the bladder is not cast off before the acquiring

of sexual characters.

Text-fig. 119.

AZ EY, c.m.

Part of a transverse section through a proglottid of the sexual worm,

c.m. Circular muscles. d.v. Dorsal vessel of the water vascular system, N. The

three laterally running nerve-cords. ¢.v. Transverse water vascular trunk,

vv. Ventral trunk. X. Large longitudinal muscular fibres referred to in the

text as frequently running within a space.

In the neighbourhood of the water vascular trunks delicate muscular fibres are

shown running chiefly in a dorso-ventral direction, which may be associated with

the dilation and contraction of the water vascular tubes.

The nerve-cord shows no trace of the asymmetry which I have

described above in the very young worms. There is nothing un-

usual about the position or structure of the main trunks. There

844 DR. F. E. BEDDARD ON A

is, however, on either side of the body and running parallel with

the main nerve-cord a second and a third nerve-cord (see text-

fig. 119, N) which lie above and below the former and at some

distance from it. These accessory nerve-cords appear to be

exactly like the main nerve-cord in structure and are situated

atthe boundary of the medullary and cortical regions, a trifle

nearer to the middle of the segment. In sagittal sections the

supplementary nerve-cord is very obvious and is seen to be con-

nected with the main trunk by numerous (7 or 8) transverse

cords in each segment, which produce a ladder-like appearance.

There is an obvious likeness here to Bertiella, in which genus the

nerve-cord also consists of three separate strands, and to Dioico-

cestus, where the cords are wider apart. They are, however, much

closer together in Bertiella than in the present worm. I have

not observed this arrangement of three nerve-cords in the supposed

immature form already described. But it is to be observed that

the main cord is there almost as wide as the medulla, thus leaving

no room for the accessory cords which may be split off later.

§ Sexual Organs.

In the middle region of the body and possibly for some way in

front (I have not examined sections just in front of the mid-

region) the proglottids are full of ripe eggs. I have examined

these proglottids in sagittal as well as in transverse sections. The

former bring out the important fact that there is apparently no

internal boundary-line between the proglottids ; for the eggs form

continuous masses stretching without intermission from segment

to segment. In this region the most careful search failed to show

any other genital organs than these exceedingly numerous ripe

ova. It is, of course, not uncommon in fully mature segments

of tapeworms, such as these in the present form, to find nothing

but ova in the medulla; but in those forms there are at least

considerable though often altered remains of the genital ducts

and terminal apparatus. In the present species there is no

vestige of cirrus-sac or vagina and not the least trace that I

could discover of an external pore. Nor do I think that it would

have been easy to have missed these structures were they present,

particularly in a series of sagittal sections. I am, in fact, con-

vinced that they are absent.

The eggs of this tapeworm do not lie in any space or spaces

that can be strictly called a uterus. There are, however, spaces

which suggest the remains of the uterus. These are not arranged

in strict metamerism ; but in a certain number of proglottids,

and not by any means confined to a single proglottid, are large

cavities such as is represented in the accompanying text-figure

(text-fig. 120). These cavities are more or less circular and lie

in the medullary region of the body. As a matter of fact,

they are not at all full of eggs, Indeed, they are often quite

empty; only sometimes are eggs to be seen lying within them

NEW ASEXUAL TAPEWORM. 845

Text-fig. 120.

Two sections through portions of proglottids of the sexual worm.

The upper figure is a longitudinal section in the more anterior part of the body,

showing the smaller ova lying among and to the inside of the longitudinal

muscles (J.); these smaller ova lie in what is probably generative tissue, not

clearly indicated in the drawing.

4. Transverse muscle fibres. 0. Mature ova scattered through medulla and

also at the opposite side of the figure in the cortex.

The lower figure represents a transverse section through a more posteriorly situated

segment.

c. A cavity which may possibly represent a uterus; two spaces are shown in the

cortex which may or may not belong to the same category. J. Longitudinal

muscles, o. Ripe ova showing a nucleus. ¢. Transverse muscles,

846 DR. F, E, BEDDARD ON A

and adherent to their walls. I doubt, therefore, whether these

spaces are the remains of the uterus, or whether they are not

rather cavities which have arisen in the medullary region perhaps

by stretching of the walls of the body and consequent laceration.

The eggs, in fact, lie scattered through the parenchyma, some-

times singly and more often in masses of various sizes, as is

shown in the figure referred to. The eggs are not limited

to the medullary region. It is clearly to be observed that

they extend into the cortex (text-figs. 120, 121) a good way

Text-fig. 121,

Bes ntl f

Mit. le Feb

. a ae

Pe eects? 4

weet RRae i

sy *

Qe @

Note Me OSA te ae,

aa oe wy er . Sum

A ba ca e

ite ee lle Ca cS ,

a ee Fs

we WE Me gS Sa Reet iu . oe

wage’ Set Beaks Ce GN

act WAP ere GCS * . a’ PSPS RL,

fy ate “

inc Rieaan enone

Sars Rae EQ le

Part of a transverse section through a proglottid of the sexually mature worm in

which the ripe ova (0.) have been very deeply stained and are seen to be

scattered through the cortex as well as the medulla,

1, Longitudinal muscles. ¢. Transverse muscles.

towards the external layer of the body-wall. They stop short,

however, some way below it. It is quite certain that whether

or no there may be remains of the uterus, some of the eggs lie

scattered within the parenchyma, In the cortical parenchyma

they are to be seen between the muscle-bundles and closely packed

in masses. The appearance is, indeed, not at all unlike that

which I have recently described in the tapeworm Anoplotenia

dasyurt * in those parts where the cavities of the uterus were

not so conspicuous; for in the latter worm there are uterine

* P,Z.S, 1911, p. 1012, text-fig, 213,

NEW ASEXUAL TAPEWORM, 847

cavities lodging many, perhaps most, of the eggs. The present

species is, as it were, a stage beyond that exhibited by Anoplo-

tenia dasywri. As is well known, the imbedding of the eggs in

the parenchyma is not a novelty; for Oochoristica is a genus

which is largely characterised by this very feature, though in

this case the eggs are imbedded singly and there are not the

heaps of eggs seen in the present species and perhaps in Ano-

plotenia. In Anoplotania, however, the masses of eggs are

usually in cavities of the uterus. In both Oochoristica and Ano-

plotenia, however, the scattered eggs are limited to the medullary

region. But the peculiarity of the species with which I am

dealing in the present paper is also known in other forms. Clerc *

has figured eggs in the cortical region in a species of Dilepis.

I have some reason to think that the scattered eggs lie in

the interstices of the network forming the ground-tissue of the

worm; but in any case eggs were also seen lying in larger

circular cavities, I believe that these cavities (shown in text-

fig. 120) are not the remains of a uterus. ‘They are, as it

appears to me, to be directly compared with similar spaces to be

seen in mature (but less mature than in the present instance)

segments of Jnermicapsifer, where I have described and figured

them‘. In this latter case I was able to bring forward some

positive evidence to show that the spaces in question were not,

and could not be, the isolated series of chambers left by a

vanishing uterus. In the worm with which I am dealing in the

present communication I am unable to furnish any of those

proofs of the nature of the cavities in question. J can merely

point out their general similarity to those of Jnermicapsifer.

Although there were no traces of gonads like those of other

tapeworms in this region of the body, I have succeeded in dis-

covering what I regard as undoubtedly the generative tissue.

The medulla was packed with ripe ova, which, as already said,

stray into the cortical layers lying between the bundles of

muscles. But at the line of junction of the cortex and medulla

and lying in the latter is a continuous mass of tissue with

interspersed nuclei, in and just outside of which eggs in various

stages of development are to be seen. ‘This is, as I think,

undoubtedly to be regarded as a continuous layer of gonadial

tissue. I cannot see any other obvious explanation of it. It is

clearly totally unlike the ground-tissue of the tapeworm’s body

and forms a solid mass with darkly staining nuclei. Its associa-

tion with developing ova seems to be final in the matter of

argument. It is, however, possibly the case that this gonadial

tissue is wholly or in part the origin of spermatozoa also; for

testes like those of other tapeworms were not found in the

present species. I have, however, no evidence of the presence of

spermatozoa. The tissue itself is—save where developing ova

* Rev. Zool. Suisse, t. xi. 1903, pl. xi. figs. 75, 76.

+ P. Z.S. 1912, p. 588, text-fig. 77. For the opposite view, see Janicki, Denkschr.

Med.-Nat. Ges. Jena, xvi. (1911) p. 381, pl. xiv. figs. 25-27.

848 DR. F. E. BEDDARD ON A

are present—naturally indistinctive. The only way of proving

its testicular nature is the discovery of spermatozoa, An

inability to find these is not of itself conclusive proof of the

absence of testicular tissue. But an argument of weight in this

direction is to be derived from a study of the ripe eggs. These

- are rather small as compared with some species and possess a

very thick hyaline shell, which is with difficulty penetrated by

staining reagents. Thus in many of my sections the eggs seem

to be simply oval structureless bodies. In cases, however, where

the staining has been more successful the true ovum within the

shell becomes obvious. It consists of a single cell with a large

nucleus. I say a single cell because there are not vitelline cells

enclosed within the shell, anything in the nature of a vitelline

body being completely absent. The absence of a vitelline gland

is rare among tapeworms, but is known in the genus called

on that very account Avitellina*. In no case did I find this

ovum in course of division—and I have examined a large

number of individual ova in many segments. I infer, therefore,

that no spermatozoa are formed in this individual worm and

furthermore that there is no entry of spermatozoa from another

individual—a fact which is also supported by the absence of a

vagina. But it must be remembered that the host may not

have contained in the bile-ducts another sperm-producing indi-

vidual. It may be that fertilisation occurs outside the body.

But this is clearly a mere suggestion for the present, though

not repugnant to such evidence as there is.

There being no trace of the actual uterus in the middle

region of the body, I naturally sought for these bodies elsewhere

and made a series of sections from the anterior region of the

body. I selected the rather thicker region which immediately

follows the head and examined a piece cut from about 10 mm.

behind the anterior extremity. Here, as I imagined, the im-

mature uterus might be found or at any rate some trace of its

existence. I could, however, find in this part no essential

differences from the posterior region of the body which has just

been described. The body was in the same way packed in places

with the ripe ova, which were present everywhere in a less crowded

state; indeed, they seemed to me to occur nearer to the external

surface here than posteriorly. I found the eggs only just beneath

the external cellular layer of the body-wall. The only conclusion

that I can come to is that this worm possesses no uterus, or

that it exists for a very transitory period only, and also that very

possibly the sexes are separated as in the genus Dvozcocestus.

The specimen described here being a female, this conclusion

is obviously based upon negative evidence only and is thus less

valuable. It is, however, quite clear to my mind that the genera-

tive products develop simultaneously in the proglottids and that

* See Gough, Quart. Journ. Micr. Sci. vol. lvi. pt. 2, 1911, for an account of this

genus.

NEW ASEXUAL TAPEWORM. 849

therefore there is no growth of the worm when it has arrived

at sexual maturity. This conclusion, which fits the facts that I

am able to set forth, agrees also, it may be pointed out, with the

very worm-like cysticercoid stage. There is, so to speak, not

much necessity for this immature worm to grow in length before

assuming the sexual condition. Far otherwise is it with the

typical Cysticercit, which are provided with but short strobila

as compared with the mature forms of their species. There is

at the very least a relation between the two series of facts which

is worth noting. It seems to me to be furthermore likely that

at no period are there sexual ducts developed—at any rate, no

female ducts. This latter circumstance, if true, is not new; for

the absence of a female orifice has been asserted in more than

one genus of worms belonging to more than one family.

Aporima among the Anoplocephalide and many genera among

the Acoleide are instances to the point. It is mainly, indeed,

the plain absence of a cirrus-sac which leads me to believe that

this worm possesses a dicecious habit so rare in this group.

§ General Résumé and Systematic Position.

It is, I think, obvious from the. foregoing account of the

sexual form of this worm that it is the representative of a new

genus which differs in a good many particulars from any known

form. It will be convenient to give a short résumé of the

essential characters of this worm, for which the name already

given to what I regard as the asexual form may be retained.

My reasons for retaining the name of the asexual form are

firstly that I have been able to give a more complete account

of it than of the presumed sexual form of the same Cestode,

and secondly that I cannot fully define the sexual form, concerning

whose identity with the asexual form, moreover, some doubt may

be considered to remain.

In any case the following are the principal characters of the

sexual worm :—Length 86 mm., greatest breadth 6 mm. Head

with two rows of hooks 16 in each row, the hooks of the anterior

row twice the size of those of the second row. Suckers normal

and unarmed. No neck; strobila consisting of numerous pro-

glottids very short and not appreciably longer at posterior end of

worm.

Body flat, thicker anteriorly. Cortex about the same diameter

as medulla. Two layers of bundles of longitudinal muscular

fibres in cortex. Water vascular tubes two on each side lying

side by side; dorsal (?) tube with thick muscular walls. A trans-

verse vessel in each proglottid forking round dorsal vessel to join

ventral at two points. No water vascular network present.

Nervous system consisting of a larger lateral trunk and two

smaller trunks, one dorsal and one ventral, connected by many

cords in each proglottid to main trunk. The sexes are apparently

separate or the worm is protogynous or protandrous throughout.

850 ON A NEW ASEXUAL TAPEWORM,

Female organs consist of a layer of gonadial tissue lying in the

medulla at its junction with the cortical layer from which ova are

shed into the body, which they permeate even to the outer layers of

the cortex. There is no trace of segmentally arranged gonads or of

a uterus, nor are the eggs surrounded by any kind of * capsule.”

There is no vagina or female passage of any kind; the vitelline

glands are totally absent. The ripe eggs are surrounded by a

thick hyaline shell and none were observed to be dividing and

no embryos were discovered in them. Thus it is possible that

fertilisation oceuwrs outside of the body,

These characteristics do not entirely fall in with those of any

other family of tapeworms. They obviously point to an aftinity

with the Acoleide, but do not definitely necessitate the inclusion

of this remarkable worm within that family.

§ Anatomical Summary.

Tt may be convenient to extract from the foregoing account

of this new tapeworm the more remarkable anatomical facts

which I have been able to make out.

(1) The absence of a marked scolex in the Cysticercus of a tape-

worm which cannot be placed among the Bothriocephalids

and of which therefore it would be expected that the scolex

would be very prominent. And correlated with this the

necessary assumption that, as in Bothriocephalids, the

scolex only develops part passe with the growth in maturity

of the worm,

2) The enormous size of the strobila as compared with the

bladder—a rare condition among the Tetracotylea, but

paralleled in Cysticercus fasciolaris.

(8) The very thick layer of muscle surrounding the dorsal

vessel of the excretory system and the bifureation of the

transverse vessels round the dorsal vessel to open into the

ventral water vascular tube.

(4) The total absence of generative ducts or (presuming that

the species is diccious) of the female tubes, which is, so

far as I am aware, a unique anatomical character.

(5) The total absence of a uterus or of any trace thereof.

(6) The diffuse and non-metameric character of the ovaries,

which are not sharply differentiated into relatively small

bodies of a definite shape.

(7) The enormous quantity of eggs produced and their exist-

ence in quite anterior as well as posterior segments; the

eggs are, moreover, found quite as abundantly in the

cortical layer as in the medulla. The conditions observable

in this part of the generative system are simply an

c<aggeration of what is to be met with in other genera,

where the eggs come to be ultimately scattered through

the medullary parenchyma. .

ON TWO NEW TREMATODES. B51

50. On Two new Trematode Parasites from the Indian Cobra.

By Wiii1am Nicoxt, M.A., D.Sc., M.D., F.Z.S., Lister

Institute of Preventive Medicine, London.

[ Received May 28, 1912: Read June 4, 1912.]

(Text-figure 122.)

INDEX.

Page

Ethology: Two new Trematode parasites in the gall-bladder

and ureters of the Indian Cobra (Naja tripudians) ...... 851

Geographical Zoology : India; Cobra, two new flukes from

the gall-bladder and ureters ...............-00ss0crecesseovevecs S01

Systematic :

Xenopharyns (gen. u., Fam. Dicroceliide) solus, sp. n.,

from gall-bladder of Indian Cobra wi... 00.00. ecceeceeeeeeee B51

Styphlodora naje@, sp. u., from ureters of Indian Cobra,

closely resembles S. horrida Leidy ..........ssccccceeeeceeeee 854

In an Indian Cobra (Naja tripudians) which died in the

Zoological Society’s Gardens on 11th October, 1911, a few speci-

mens of two interesting new Trematode parasites were found.

Of the first of these a single specimen was found in the gall-

bladder; of the second, four somewhat macerated specimens were

met with in the ureters. Four different species of Nematode

parasites were also present in the same animal, so that altogether

it was infected with six different species of parasitic worms.

The first species belongs to the family Dicroceliide and is of

particular interest from the fact that members of this family

are not commonly found in Reptiles. Hitherto the only typical

representative in Reptiles is that described by de Faria (1910).

The present species shows most of the characteristic features of

the family, but at the same time it exhibits several divergences of

such importance as to warrant its being regarded as the type of a

new genus. For that genus I propose the name Xenopharynz.

XENOPHARYNX SOLUS, gen. et sp. n. (Text-fig. 122, B.)

Only a single specimen was found in the gall-bladder. It

measures 4°55 mm. in length and 1°68 mm. in greatest breadth,

which is about the middle of the body. The outline is almost

elliptical and the body is fairly flat. There are no cuticular

spines.

Pthe oral sucker has a diameter of -42 mm., but its length is

only 30 mm. The ventral sucker is circular, with a diameter of

‘41mm. It is situated 1:13 mm. from the anterior end. The

neck is therefore almost exactly one-fourth of the body-length.

The pharynx is contiguous with the oral sucker and measures

‘'22x°24 mm. It possesses the curious shape shown in text-fig.

122, B. At first sight this shape was thought to be the result of

852 DR. WILLIAM NICOLL ON

unequal contraction of the pharyn geal walls or perhaps due to the

drawing of the esophagus into the lumen of the pharynx, but

neither of these possibilities seemed to accord with experience.

The condition appears to be brought about by the thinning of the

Text-fig. 122.

M. Rhodes del.

A. Styphlodora naje, sp.n. Ventral view. x 40. C., cirrus;

Ov., ovary; R.S., receptaculum seminis.

B. Xenopharyne solus, gen. et sp.n, Ventral view. xX 25.

Ov., ovary.

posterior part of the pharyngeal wall, which anteriorly is of the

usual thickness. In section the appearance is somewhat horse-

shoe-shaped. It is unfortunate that no second specimen was

TWO NEW TREMATODES. 853

; available for comparison. Following the pharynx, and about as

long as it, is a wide cesophagus, which bifureates about midway

‘between the pharynx and the ventral sucker, The diverticula are

fairly uniform, but become somewhat sinuous towards their ter-

mination, which is -45 mm, from the end of the body.

The excretory vesicle is Y-shaped, with a long median stem

reaching the middle of the body and two short limbs, the left of

which extends as far forward as the left testis. Owing to the

fact that sections could not be made, it was impossible to determine

whether Odhner’s view with regard to the shape of the excretory

vesicle in the Dicroceeliidee holds good in this case. To all

appearance, however, the paired limbs are part of the vesicle,

Their walls stand ont as distinctly, and are about the same

thickness, as those of the main stem.

The testes lie not far behind the ventral sucker. They are two

transversely oval bodies, separated from each other by a distance

equal to their diameter, They are situated obliquely, the right

being about half a diameter in advance of the left, and being

separated from the ventral sucker by a similar distance. They

measure *31 mm, in transverse diameter. The cirrus-pouch is of

small size and les entirely in front of the ventral sucker. It

measures ‘29 x'13 mm., and contains a small convoluted vesicula,

seminalis, a comparatively long pars prostatica, and a short narrow

ductus ejaculatorius. The genital aperture is median, just over

the intestinal bifurcation,

The ovary lies a considerable clistance behind the testes, about

midway between them and the end of the body, It is on the

right side overlying the right intestinal diverticulum, Its outline

is transversely oval and it is much Jarger than the testes,

measuring 31 x°47 mm. There appears to be no receptaculum

seminis, or if present it is obscured by the uterus, The yolk-

glands are situated chiefly in the neck, the sides of which they

almost completely fill. They extend on each side from the oral

sucker to well behind the ventral sucker, but they are more ex-

tensive on the left than on the right, On the right they stop at

the level of the bifureation of the excretory vesicle, although

there are a coupie of small folligles near the ovary. On the lett

they extend slightly beyond the level of the ovary. In the neck

the follicles spread well in towards the mid-line of the body, but

behind the ventral sucker they lie entirely to the outer side of the

jntestinal diverticula. ‘The transverse yolk-ducts cross the body

about the level of the testes, and the shell-gland lies just behind

the right testis,

The uterus is moderately yoluminous and is thrown into

irregular narrow convolutions. It does not extend much behind

the ovary, but in front it oyerlaps the intestinal diverticula, and

winding between the testes it passes forward on the left side of

the ventral sucker and makes a few short turns in front of the

sucker before passing into the short vagina. The eggs are small

and numerous, measuring ‘036-039 x (018-019 mm.

Proc. Zoou, Soc.—1912, No, LVI. 57

854 DR. WILLIAM NICOLL ON

The new species differs from all other Dicrocceliide in the dis- |

tribution of the yolk-glands and in the shape of the pharynx.

Another important feature is the distance separating the ovary

from the shell-gland complex. From the genus Dicroceliwm it is

further distinguished by the position of the testes and the ovary

and by the extent of the uterus. With Platynosomwm it is more

closely allied, yet the differences between it and this genus are

greater than those separating Dicrocelium from Platynosomum or

Eurytrema. Yt is rather curious that this new species does not

show any particularly close relationship to “ Dicrocelium ” infidum

de Faria from the snake, Hunectes marina. The latter is closely

allied to the avian genus Platynosomwm and should be included

in it, unless de Faria’s doubtful observation in regard to the

excretory vesicle proves to be correct.

STrYPHLODORA Nas#, sp.n. (Text-fig. 122, A.)

Four somewhat macerated specimens of this species were found

in the ureters of the Cobra. The species is a typical member of

the genus Styphlodora and presents a very great resemblance to

S. serrata Looss and to S. horrida Leidy. The features separating

it from these two species are so slight that I have some hesitation

in regarding it as a distinct species. They are, however, quite as

distinctive as those separating the above two species from each

other. The difficulty is rather increased by the fact that the

present specimens, although mature, are possibly not fully grown.

The length is 2°0-2°4 mm., the greatest breadth -42—55 mm.,

which occurs a little behind the ventral sucker. The breadth of

the whole postacetabular region is fairly uniform, and there is

only a very slight attenuation in the neck. There isa considerable

amount of dorso-ventral flattening. Im each of the specimens

cuticular spines were entirely absent, but it is practically certain

that they have fallen off, and it would be unreasonable to suppose

that this is an unarmed form. On that account I have had spines

depicted in the drawing.

The oral sucker is subterminal and hasa diameter of -22--25 mm.

Tt is rounded and rather shallow. The ventral sucker is slightly

transverse and measures -24x:25 mm. It is only very little

larger than the ora sucker, and it is situated at a distance of

-67—84 mm. from the anterior end, 7. e. about one-third of the

body-length. There is a distinct prepharynx followed by a large

pharynx measuring about -13x-14 mm. The cesophagus is about

the same length, and the bifurcation takes place midway between

the suckers. The intestinal diverticula are fairly straight and of

considerable width. They are longer than in the other species of

Styphlodora, reaching to within ‘18-29 mm. of the posterior end.

They are very slightly unequal in length.

The main excretory vesicle was entirely invisible, but it was

apparent that it gave off numerous lateral branches, which,

anastomosing freely, gave the body a honeycomb-like appearance.

The testes lie obliquely behind one another, the left being in

TWO NEW TREMATODES. 855

front. They are separated by the uterus, which passes between

them, but does “not overlap them to any great extent. Their

outline is irregular, but they appear to be roughly triangular ar

trilobate. The anterior testis lies almost exactly midway between

the two ends of the body, but its position in relation to the ovary

varies somewhat, and it may be slightly nearer than is shown in

the figure. The posterior testis is about -27 mm. behind the

anterior, They touch the intestinal diverticula or overlap them

to a very small extent. Their dimensions are, on an average,

*21 x17 mm. and '22x°19 mm, The cirrus-pouch is of moderate

length and uniform width, measuring °31x-11 mm. It extends

to about the posterior border of the ventral sucker, and contains

a convoluted vesicula, a short pars prostatica, and a fairly long

ductus ejaculatorius. In each of the specimens the cirrus was

exserted and was longer than the pouch itself. The genital

aperture is situated in the middle line, almost immediately in

front of the ventral sucker.

The small round ovary lies just behind the ventral sucker, a

little to the right side, and measures -13 mm, in diameter,

Immediately behind it les a somewhat smaller receptaculum

seminis. ‘The yolk-glands lie on each side at the level of the

ovary, They are, for the most part, external to the intestinal

diverticula, but they also overlapthem dorsally. They consist on

each side of about a dozen fairly large follicles, which extend from

the posterior border of the ventral sucker to the anterior border

of the left testis, The transverse yolk-ducts cross the posterior

border of the ovary.

The uterus is poorly developed, It passes back between the

testes to near the posterior end of the body and returns along

much the same path, terminating in a short vagina. It is only

slightly convoluted and contains a comparatively small number

of eggs, It is confined within the space bounded by the intestinal

diverticula, and it does not form any convolutions behind the ends

of the intestine, as is the case in the other species of Styphlodora.

The ova are rather elongated, oval, possessing a large operculum

and, in a few cases, a small knob at the anopereular pole. They

measure °038—'048 mm. x ‘019-024 mm., the average size being

"043 x 021 mm.

The features which appear to distinguish this form as a definite

species are the length of the intestinal diverticula and the

restricted extent of the uterus. It is not impossible that both

these features may be less marked in a fully-grown specimen,

In addition the yolk-glands are slightly less extensive than they

are in the other species of the genus. From S. serrata it is

further distinguished by the relatively larger size of the suckers,

Both S. condita and S. horrida also possess relatively smaller

suckers, and in these species the distance between the suckers is

considerably less. The only other species of Styphlodora, namely

S. similis Sonsino and S. bascaniensis Goldberger, present marked

features of difference from the present species.

57*

856 DR. WILLIAM NICOLL ON

REFERENCES.

1. De Faria, Gomez. 1910.—Contribution towards the Classifi-

cation of Brazilian Entozoa. I. Dicrocelium infidum,

n. sp., Parasite of the Gall-bladder of Hunectes marina L.

Memorias do Instituto Oswaldo Cruz, 1i. (1) pp. 22-28, pl. 2.

2. De Farra, Gomez. 1911.—Beitrige zur Systematik der bra-

allianischen Helminthen. IV. Styphlodora condita, n. sp.

Memorias do Instituto Oswaldo Cruz, iii. (1) pp. 40-45, pl. 1.

3. GoLDBERGER, J. 1911.—A New Trematode (Styphlodora

bascaniensis) with a blind Laurer’s Canal. Proc. U.S. Nat.

Mus. xl. pp. 233-239.

4, Lerpy,J. 1850.—Descriptions of Two Species of Distoma, with

the partial History of one of them. Journ. Acad. Nat. Se.

Philadelphia, ser. 2, vol. 1. pp. 301-310, pl. 43.

5. Looss, A. 1899.—Weitere Beitrige zur Kenntniss der

Trematoden-Fauna Aegyptens. Zoolog. Jahrbiicher, Abt. f.

Syst, x1. pp. 707-708, pl. 26, fig. 28.

6. Opuner, T. 1910.—Nordostafrikanische Trematoden. Results

Swedish Zool. Exped. to Egypt and White Nile, 1901, p. 53,

note 2, fig, v.

D1. Statistical Note on the Worm Parasites collected from

the Animals dying in the Zoological Gardens, from

December 1910 till April 1912. By Witi1am Nico.t,

M.A., D.Se., M.D., F.Z.S., Lister Institute of Preventive

Medicine, London.

[ Received May 30, 1912: Read June 4, 1912. |

INDEX.

Page

Ethology: Animals dying in the Zoological

Society’s Gardens, numbers infected with

PATASIUIC! WOLMISIse.c..0.;hee cReeeaeerce ete ee OO

Geographical Zoology : Importance of ascertaining

distribution of parasitic worms .................. 857

At the scientific meeting of the Society held on May 21st last,

{ referred to the excellent work which is being done by the

Prosectorial department in acquiring information concerning the

parasites which infect the animals living in the Gardens. This

is particularly noteworthy in regard to the worm parasites, about

which many valuable facts have been obtained, Following my

remarks at the meeting, Dr. Beddard suggested that I might be

able to supply some general information regarding the animals

which had been sent to me for examination, and acting on this

suggestion I venture to offer the following communication.

The scheme, which owes its initiation to the Secretary of the

WORM PARASITES. 857

Society, was put into action in N ovember 1910, and the practical

working has been evolved as follows : A general examination of the

viscera of all the animals killed or dying in the Gardens is made

by the Pathologist, after which certain selected examples are

forwarded to me for further examination. The desirability of this

latter procedure is shown by the fact that a very large proportion of

Entozoa are too small to be detected in the course of an ordinary

routine examination; and although the detection of such forms

does not involve any very special skill, yet it 1s a tedious process

and one which necessitates some experience. In justification of

this extra trouble it may be remarked that the comparatively

few animals which have been submitted to this additional

examination have yielded almost as large a number of parasitic

worms as all the other animals put together. This fact, I

venture to believe, is a plea for a still more extended and

exhaustive system of examination, dealing with all the animals

which die in the Gardens. This, naturally, would involve some

additional expenditure, which the Society is probably not at

present in a position to incur, but it would certainly yield a very

rich return.

In addition to supplying information with regard to the

diseases and habits of the animals living in the Gardens, as was

the original intention of the scheme, it also provides oppor-

tunities of studying the parasitic fauna of the various countries

from which the animals come. This is a very important con-

sideration, for it is an unfortunate fact that in most of the large

faunistic expeditions which have hitherto been undertaken

practically no attempt has been made to deal with the internal

parasites. As important exceptions to this may be mentioned

the German Expedition to Spitzbergen in 1898, and the Swedish

Expedition to Egypt and the White Nile in 1901. By both of

these expeditions a large number of parasitic forms has been

collected, and the results have been published in a series of very

valuable monographs. Dr. Leiper has, at these meetings, on

more than one oceasion called attention to this regrettable

neglect of the parasitic fauna, and has urged on collectors the

great service they would render by making even a cursory

attempt to collect parasitic worms.

During the past seventeen months a total of 198 animals have

been submitted to me for examination, the great majority of

which have been birdsand reptiles. Of these, 87 were found to

be infected with worm parasites of one kind or another, 2.¢.,

about 44 per cent. In most cases the infection was single, only

one species of parasite occurring ; but in an Indian Cobra as many

as six different species were found. As is generally the case,

Nematode infections were much more numerous than any others,

there being 60 cases. Trematodes were found in 28 cases, and

Cestodes in 15. The Trematodes belong to about 20 distinct

species, a large proportion of which are new, and several of

which are new generic types.

858 ON WORM PARASITES.

From the reports of Dr. Beddard and Dr. Leiper I gather that

during the same period 41 additional cases of infection with tape-

worms, and 53 of infection with Nematodes have been observed.

To these must be added a very considerable number of infections

with larval Filariz observed by Mr. Plimmer. These figures,

excluding the larval Filarie, give a total of about 180, of which

the infection with Trematodes, Cestodes, and Nematodes is

respectively 28, 56, and 113. This appears to show that

Trematode parasites are much rarer than other forms, but that

is certainly not the case. It must be remembered that only a

comparatively small number of the animals passing through the

Prosectorium were adequately searched for Trematodes. The

extent to which Trematodes and the smaller intestinal Nematodes

escape notice except when thoroughly searched for may be

gathered from the following table.

Trematodes. Cestodes. Nematodes.

Total number of cases of infection 28 56 Ike

Number of cases found only on

further examination ........ esp 20 15 60

MASI {SIA EEX) a Mig A ee al Mean cE ORE SF (pil 26 53

It may be added that the bulk of the Cestoda which were

not noted on first examination were small immature forms or

fragments of no value. It may therefore be safely affirmed that

only a negligible fraction of the mature tapeworms actually

escape detection in the Prosectorium. The matter, however, is

different with Trematodes ; and in view of the fact that only about

one-tenth of the total number of animals dying in the Gardens

have been submitted to a thorough examination, it must appear

that the number of Trematodes actually occurring is considerably

greater. Even allowing that they only occur in 5 per cent. of

cases, which is taking an extremely low estimate, we ought to

have had instead of 28 records a total of well over 100. In the

same way the number of small intestinal Nematodes ought to

be much increased.

These remarks, however, are not intended as a criticism of the

efforts of the prosectorial staff. Parasitic worms are, after all,

only a very small part of the work of this department of the

Society, and as I have mentioned in the beginning of this com-

munication, considerable praise is due for the results actually

obtained. My chief intention is to show that parasitic worms

are undoubtedly an important concern and that their importance

justifies the attention which is being paid to them. Their claim

to notice had until recently been somewhat neglected by the

Society, and it seems necessary that someone should offer a plea

for their vindication. There can be no doubt that the attention

which the Society, thanks to the efforts of Dr. Chalmers Mitchell,

is devoting to these forms will do much to benefit Zoology as a

whole.

B.Brosm del,

SOUTH AFRICAN FOSSIL je SAID Sy,

Huth, Lith? London.

iB apo bs ia dai oI,

Huth,Litht London.

SOUTH APRICAN FOSSIL REPTILES.

R.Broom del.

PZ. S192 PL xem,

Huth, Lith? London.

R.Broom del

SOUTH APRICAN HOSSi. Be Sitka.

Py sige SL <eoun

BR.Broom del, . Hoth,Lith? London.

SOUTH AFRICAN FOSSIL REPTILES.

ON NEW FOSSIL REPTILES. 859

52. On some new Fossil Reptiles from the Permian and

Triassic Beds of South Africa. By R. Broom, D:Se.,

C.M.Z.S.

[Received May 13, 1912: Read June 4, 1912. |

(Plates XC.—XCIII.*)

INDEX.

Systematic : Page

Taurops macrodon, gen. et Sp. . ......-seee eee eee ees 859

Galeops whaitsi, gen. et Sp. De ....-. 2.4.22 see eeeeee eee 860

Scymnognathus whaitsi, gen. et sp. Me .....---.-++ ++ 861

Flurosaurus striatidens, sp. n. ......cecceeeeeee eee es 868

Pristerognathus platyrhinus, sp. U........--...-esree 863

Alopecorhinus parvidens, gen. et Sp. Ne .....-...++- ++ 864

Tctidognathus hemburyi, sp. WM. ...-....-...s00eeeere eee 865

Hindothiodon whaitsi, sp. ms ..ic..-:c-se-ceeeeeeeene ess 866

B. platyceps, Sp. W..........ceesce eee ceeeee cee rte ereetecee ees 867

Prodicynodon beaufortensis, sp. Ne ......-..+16eee sree 867

Dicynodon to include Oudenodon ..........-..-2+ 868

Pristerodon to include Opisthoctenodon............... 868

Dicynodon laticeps, sp. 0...........0seeee testers cree 868

D. psititacops, SP. W. ....-...eeeeseerenenene teers ceneee ces 869

D. lutriceps, Sp. We... 2.0... see ceeceeeee eee eee seetteces sees 870

Emydops minor, gen. et sp. 0. -.e-ee-eeeeseeereerrees 871

Ictidopsis elegans, gen. et SP. De ......+eeeeeeee serene 872

Nythosaurus browni, sp. 0. 2.0.0.0... terres 874

Endothiodon subdivided: see Addendum ............ 875

Emydochampsa, gen. D........-..20eeeeeeeeeterees teeeetens 875

The following series of new reptiles forms an important addition

to our knowledge of the Karroo faunas. The majority have

been found by the Rev. J. H. Whaits, of Beaufort West. Of

the others one was found by Mr. Alfred Brown, of Aliwal North,

one by Mr. H. J. Hembury, and a few by myself.

Suborder DINOCEPHALIA.

TAUROPS MACRODON, gen. et sp.n. (Pl. XC. fig. il)

The snout of this large Dinocephalian was obtained at Bos-

manshoek at the foot of the Komsberg. Though so little has

been obtained we have enough to give the distin guishing characters

of this new type. In size it almost equals Z'apinocephalus

atherstonei, but the snout though narrower is considerably deeper,

and it differs in the great degree of development of the teeth,

especially of the incisors. Where the snout is broken across,

about the transverse plane of the back of the anterior nares, the

width is 190 mm. and the height about 100 mm. The anterior

nares are unusually small and situated about 90 mm. from the front

of the snout. The distance between the two nostrils is about

45 mm. The condition of the bone renders it almost impossible

to make out the sutures, the only one that is distinct being that

between the two premaxillaries.

%* For explanation of the Plates see p. 875.

860 DR. R. BROOM ON

The teeth are for the most part badly preserved, but as they

are broken off at different levels and as a large number of

replacing teeth are present, the structure can be made out without

much difficulty. There are 14 teeth preserved, of which the

first five are large and long. From the 6th backwards the teeth

steadily decrease in size and the crowns become quite short.

Possibly there are two or three teeth lost behind the 14th, but

it seems improbable that there are many. The anterior teeth

are of the usual Dinocephalian type. They are, except where the

root is being absorbed by a replacing tooth, of great length, the

whole root and crown being about 90 mm. The anterior cusp is

long and narrow, and on Ecrtion is semicircular. At its base it

measures about 9 mm. across and gradually narrows towards the

point. None of the anterior upper teeth shows the whole anterior

cusp, but it probably measures about 20 mm. in length if not

more. The posterior basal cusp is about 15 mm, in width and

has a slightly concave surface. The posterior teeth have short

cusps. The 12th tooth has a crown only 12 mm. in height.

The teeth in the lower jaw are apparently closely similar in

type to those in the upper. Most of the functional teeth are

badly weathered away, but the weathering shows that at the base

of each is a well-developed replacing tooth similar in type to the

functional one. It seems probable that the long anterior cusps

of the front teeth interdigitated and that the gr inding took place

by the meeting of broad posterior cusps. The edges ice the long

anterior cusps would thus form an admirable cutting appar atus,

and the internal cusps would take the part of molars, In the

Dinocephalians almost the whole dental appar atus 1s placed in the

front of the snout, and the absence of grinding molars such as are

seen in ibe contemporaneous or slightly, earlier Ameri ican herbi-

vores, e.g. Diadectes, is explained by the crushing or grinding

function iar been taken up by the peculiarly specialised

incisors.

Suborder DROMASAURIA.

GALEOPS witArtst, gen. et sp.n. (Pl. XCT. fig. 6.)

This new genus is founded on the anterior half of the skeleton

of a Dromasaurian. The skull is crushed but fairly well preserved,

and the shoulder- -girdle and front limb are in good condition,

while remains of about 18 vertebre are seen. Unfortunately the

matrix is very hard and little can be done in the way of develop-

ment. The specimen was found by Mr. Whaits on the farm

La-de-da, about 20 miles to the west of Beaufort West, and

probably belongs to the upper part of the Pareiasaurus- Zone.

The skull is in many respects very remarkable. The orbit is

exceedingly large and the temporal fossa deep and very narrow,

and the squamosal has a long descending process which brings

the quadrate below the back of the orbit. ihe face is ver 4 short,

and there appear to be no teeth in either upper or lower jaw.

The bones of the skull are not in good condition for showing

NEW FOSSIL REPTILES, 861

sutures. The snout is missing in front of the nostril, but from

the shape of the lower jaw its length can be fairly well assumed.

The nostril is large and the distance between it and the orbit very

short. It is probable that the septomaxillary meets the lachrymal.

The suborbital and postorbital arches are slender. The frontal

and parietal regions are both fairly wide and there is a large oval

parietal foramen. The squamosal is somewhat like that of the

Dinocephalia, but is slender and the descending process very long.

There appears to be a distinct quadrato-jugal. The palate, so far

as preserved, agrees with that of the Therocephalia. There is a

pair of long slender prevomers, and the palatines and back of the

pterygoids are not unlike those of Scylacosaurus, but whether in

other respects the palate is Therocephalian or Dinocephalian the

evidence does not show. Certainly the palate is not the least

like that of the Anomodont. The lower jaw is short and tooth-

less. It agrees fairly closely with that of the Anomodont, but

there is a rudimentary coronoid process.

The shoulder-girdie has a large distinct precoracoid and coracoid

loosely articulated to each other. The scapula is somewhat like

that of the Dinocephalia and also a little like that of the Thero-

cephalian Jetidosuchus, but unlike that of the Anomodont. The

limb bones are long and slender.

Galeops is an entirely new type of Dromasaurian and represents

a new family, the Galeopide, characterised by the absence of

teeth and the presence of a small coronoid process. The other

family, which may be called the Galechiride, includes Galechirus

and Galepus, both with teeth and without a coronoid process.

Suborder THEROCEPHALIA.

SCYMNOGNATHUS WHAITSI, gen. et sp.n. (PI. XC. figs. 4, 5.)

This interesting type was found by Mr. Whaits near Beaufort

West. Within quite a small area remains of four or five

animals were obtained. Unfortunately, most of the bones are

extremely weathered, and so infiltrated with lime as to be

practically limestone nodules so hard that any development is

almost impossible. Two skulls are sufticiently well preserved to

show the general characters of the genus, though it is impossible

to he sure of most of the sutures.

The new type is a very near ally of Gorgonops torvus Owen,

and it is only after considerable deliberation that I have decided

to place it in a new genus. With perfect certainty it can be

placed in the Family Gorgonopide. When Owen first described

Gorgonops in 1876 he unfortunately came to the conclusion that

the temporal fossa was roofed over as in Labyrinthodonts or

Pareiasaurus, and this mistake was also made by Lydekker in

1890, and by Seeley in 1895. On examining the type and, at

that time, only known specimen when in London three years ago,

I discovered that the temporal fossa is not roofed over, though

the parietal region is broad, and noted the observation in a paper

862 DR. R. BROOM ON

published in 1910*. Within the last few months Mr. Whaits

has fortunately discovered a good skull of Gorgonops which shows

that it has quite a large temporal opening, very similar to that

in the present genus.

In general appearance Seymnognathus differs from the more

typical Therocephalians mainly in having the intertemporal region

about as wide as the interorbital. The snout is long, the orbits

small, the temporal region fairly wide, and the squamosals more

powerfully developed than in most Therocephalians.

The total length of the skull is, in the type, 305mm. In a

second but much crushed specimen the length is probably about

325 mm. From the front of the snout to the front of the orbit

the measurement in the type is 165 mm., and the length of the

orbit about 40mm, The interorbital width in the type is 70 mm. :

in another very imperfect specimen it measures 78 mm., and na

third about 76 mm. The narrowest part of the parietal region

measures in one of the larger specimens 78 mm.

The parietal foramen is situated well back and is 6 mm. in

diameter.

Though the limits of many of the bones cannot clearly be made

out, much of the cranial structure can be seen. The bones of the

snout seem to agree with those of the better known Thero-

cephalians, though the septomaxillary is relatively larger. The

jugal is unusually well developed and the postorbital is very large.

The postfrontal also seems to be much larger than in any other

type hitherto examined. The squamosal is more massive than in

most Therocephalians, and the quadrate is relatively small and

largely hidden by the squamosal. The occipital condyle is single.

The lower jaw is fairly similar to that previously described

and figured in Lycosuchus and Aloposaurus. The front of the

dentary is very deep and has a well-marked mental process.

Posteriorly the dentary has a short coronoid process and for a

considerable distance lies above the angular. The surangular is

much smaller than in other known Therocephalians.

There are five upper incisors, one large canine and three or four

molars. The incisors are long pointed teeth, oval in section, and

apparently without serrations. The space occupied by the five

incisors in three specimens is 37 mm., 39 mm., and 40mm. The

space between the last incisor and the canine in four specimens is

18 mm., 22 mm., 25 mm.,and 26mm. The length of the canine

in seven specimens is 16 mm., 18 mm., 20 mm., 20 mm., 20 mm.,

20 mm.,and 21 mm. Behind the canine is a diastema varying from

7 mm. to 16 mm., followed by either three or four molars. The

number of molars apparently depends on age. When three molars

are present the distance occupied by them is 20 or 21 mm.: when

four are present they occupy 25 mm. to 29 mm.

SS) 1 3 or 4

»1. 4, ¢. 5m

The dental formula is i ™ sor4°

* “Observations on some specimens of South African Fossil Reptiles preserved in

the British Museum.” ‘Tr. Roy. Soc. 8. Afr. vol. 11, pt. 1, 1910, p. 20.

NEW FOSSIL REPTILES. 863

Scymnognathus may prove to be related to Seymnosaurus.

The dental formule are practically the same in the two genera,

but Scymnosaurus seems to be a higher type of Therocephalian,

and the shape of the snout and lower jaw differs sufticiently

markedly to justify one in keeping the genera distinct.

AELUROSAURUS STRIATIDENS, sp.n. (Pl. XCI. fig. 7.)

The imperfect snout which forms the type of this new species

was discovered by me at Kuilspoort, near Beaufort West, at a

horizon which is probably 500 feet above the town.

It is a smaller species than J. felinus, and in addition differs

from this and the other known species in the relatively small size

of the teeth, in the feeble development of the serrations, and in the

incisors and canine having feeble vertical ridges on the enamel.

So far as preserved there is a pretty close agreement in the

structure of the bones of the snout with those of previously

described species. The septomaxillary is larger than usual. The

mandible has a broad angular chin.

The incisors are small, rounded, pointed teeth, placed so near

each other as to be almost touching. Those whose crowns are

preserved (3rd, 4th, 5th) have the enamel folded into about half

a dozen vertical corrugations. The 5th tooth has fine serrations

on its posterior edge. The space occupied by the five incisors is

16mm. The diastema between the 5th incisor and the canine is

7 mm.

The canine is long and slender. The base measures antero-

posteriorly 7 mm. and the height of the crown as preserved is

14mm. It originally probably measured about 20 mm. The

molars are small with only faint indications of posterior serrations.

Two are well preserved and there are remains of the other two.

Probably 5 is the complete number, which would make the dental

formula agree with the other species of @luroswurus. The four

preserved molars measure 10 mm., and the front one is only 9 mm.

behind the canine.

PRISTEROGNATHUS PLATYRHINUS, sp. n. (Pl. XCI. fig. 8.)

The specimen which I take as the type of this new species is an

imperfect snout found by Mr. Whaits at Grootfontein, about 12

miles to the west of Beaufort West, and probably from the upper

part of the Pareiasauwrus-Zone. The specimen consists of the

front half of the skull. It is broken into three pieces and the

upper nasal region is missing. The matrix is extremely hard and

difficult to clear off, but most of the characters can be satisfactorily

made out.

In the large majority of Therocephalians the snout is deeper

than broad. In this specimen the snout is broad and flat, and the

lower jaw comparatively straight and with very little of the usual

upeurving in the canine and incisor region. The widest part of

the snout is immediately above the canine, where it measures

55 mm, and on the same plane the height of the snout is 40 mm.

864 DR. R. BROOM ON

There is very little crushing. Nearer the front of the nose the

height is only about 25 mm. and the width 45 mm. The

maxillary bone is rather deeply pitted. The palate is broad

and flat behind and in front slopes upwards to the opening of the

internal nares. The prevomers are narrow slender bones which

in section are seen to have thin vertical plates. The palatines

are apparently as in Seylacosaurus, but appear to approach each

other more closely in the middle line. The pterygoids have each

a very thin vertical plate which is closely placed against its

neighbour.

There are apparently 6 incisors, though only evidence of the

last five are preserved. The anterior ones are fairly round on

section, but the last three are more flattened and havea posterior

edge which is not improbably serrated. The teeth decrease in

size as we pass backwards, the 6th being only about half the size

of the 4th. The last four teeth occupy a space of 20°5 mm. and

the whole six probably 30 mm. The diastema between the 6th

incisor and the canine is 9mm. ‘'Thecanine at its base measures

10 mm. by 7mm. Behind the canine is a diastema of 7 mm.

followed by 8 pointed molars. Each of the anterior molars has

an antero-posterior diameter of nearly 3 mm., and the whole series

occupies a space of 29 mm.

The dentition of the lower jaw is mainly concealed in front.

Probably there are 3 incisors and 1 canine. The molars are well

shown on the right side and are 8 in number, and occupy a space

of 32mm. There is no evidence of any serrations, and were they

present the specimen would be expected to show some of them.

WSS eh

There is some doubt whether this specimen belongs to the same

genus as Pristerognathus polyodon Seeley. The type is only the

front of a snout, and the number of molars is unknown, and only

a figure of the underside is given by Seeley. However, the two

specimens are probably from near the same horizon, and the

arrangement of the teeth, so far as known, is sufficiently close to

render it advisable to place this new species provisionally in

Seeley’s genus Pristerognathus.

The dental formula is probably i. - c.

ALOPECORHINUS PARVIDENS, gen. et sp.n. (PI. XCI. fig. 9.)

This new genus is founded on animperfect snout discovered by

Mr. Whaits at Beaufort West. It consists of the greater part of

the left maxillary and dentary and much of the right maxillary

and dentary,

It resembles Pristerognathus platyrhinus in the broad short

nose and approximates in dental formula, but differs in having

a much more slender jaw and in the relatively smaller size of the

teeth, especially the molars, and in having a much shorter

precanine portion of the snout.

The anterior upper incisors are lost but there is evidence of the

last three. The whole series probably measured 19mm. The

NEW FOSSIL REPTILES. 865

diastema between the last incisor and the canine is only 2 mm.

The canine measures at its base 6 mm.by 5mm. The molars

are at least 7 in number, and are small, pointed but apparently

unserrated teeth. The seven occupy 17°5 mm. Between the

canine and the first preserved molar is a diastema of 9 mm.

The lower incisors are preserved in section and are 4 in number,

the 4th being inside the line of the others. The canine is far

forward and small and rounded, the section measuring 3:5 mm.

in diameter.

IcTIDOGNATHUS HEMBURYI, sp.n. (Pl. XCI. figs. 10, 11.)

This new species is founded on four imperfect snouts found by

Mr. H. J. Hembury at Beaufort West. There are minor

differences between the snouts probably due to crushing, to age,

and possibly to sex, but I believe they all belong to one species.

To avoid any possibility of confusion I shall take the skull

(fig. 10) as the type. It is the best preserved specimen but does

not show the molars satisfactorily.

As in Alopecorhinus the snout is broader than deep. This

is probably also the case in /ctidognathus parvidens, but the only

known specimen of this species is considerably crushed. From

the front of the snout to the front of the orbit the measurement

is 42mm. The antero-posterior diameter of the orbit is 18 mm.

The interorbital region is 19 mm. across. The measurement

across the snout at the canine region is about 27 mm.

The premaxillary is a very small bone forming the anterior and

lower margins cf the nostril. It carries six small pointed, rounded,

smooth incisors. I fail to detect serrations on any of them, The

S1x Incisors occupy a space of 10 mm.

The septomaxillary is unusually large and formsas large a part

of the facial surfaces as does the premaxillary. There isthe same

foramen between it and the maxillary seen in typical Thero-

cephalians, but it is relatively smaller than in other forms.

The nasal bone is well developed and is interesting from the

fact that it is only very little broader behind than in front.

The maxillary is typically developed. As in so many Thero-

cephalians, the centre part of the bone is markedly pitted as if for

the accommodation of glands or sense organs. In the same region

there are numerous foramina passing into the bone, and some of

these foramina lead back into a canal in the bone. As there is

no large foramen in the maxillary bone which might be regarded

as the foramen for the maxillary branch of the Vth nerve, I

think it probable that this nerve subdivides in the maxillary

bone and comes to the surface by a number of small foramina,

and that the pits were for sense organs which were supplied by

this nerve. The tactile vibrissee of mammals, or the remarkable

sense organs of the beak of Ornithorhynchus may be the modified

homologues of these supposed ancestral organs,

866 _ DR. R. BROOM ON

There are two canines—a very small anterior and a fairly large

long slender posterior one. The small canine has a diameter of

-5} mm.: the large canine measures 3°5 mm. by 4mm. The two

are a little more than 2 mm. apart. The exact number of

molars is a little doubtful but appears to be 8, and they occupy

16 mm. in one of the specimens. The molars appear to be

all short, smooth, pointed teeth without serrations.

In the lower jaw there are four incisors, a large canine, and eight

molars. The dental formula would thus be i. : ¢. = Mm. 3. In

Ictidognathus parvidens there are certainly nine upper molars and

possibly ten, but as this species is certainly allied to 7. parvidens

it seems better to keep both in the same genus.

Suborder ANOMODONTIA.

* ENDOTHIODON WHAITSI, sp.n. (Pl. XCIII. fig. 18.)

This new species is only known by the skull, a few vertebree

and ribs, and a couple of limb-bones. Fortunately the skull is in

beautiful condition. The specimen was discovered by the Rev. J.

H. Whaits at Beaufort West. It is a near ally to Hndothiodon

uniseries Owen, and has the molars in a single row, but it

differs from H, uniseries in a considerable number of points

besides in being nearly twice as large. In EHndothiodon wniseries

the greatest length of the skull is 360 mm.: in Hndothiodon

whaitsi the skull measures in length 570mm. With the ex-

ception of Oudenodon magnus it is the largest known Anomodont.

As I hope shortly to publish a full account of the genus Endo-

thiodon, I shall heve merely give a preliminary description of this

interesting species.

The skull is narrow and deep. The greatest width across the

squamosals is probably about 360 mm., while the interorbital

region is only 140 mm. and the greatest width of the palate

150mm. ‘There is a slight degree of crushing, but not such as to

make these measurements far wrong. ‘The notch in the pre-

maxilla for the point of the lower jaw is very deep and narrow.

The nostrils are large, measuring 65 mm. by 45mm. The nasals

do not overhang them as in #, wniseries, the whole width of the

nasals being only 85 mm. The orbit is situated 180 mm. behind

the front of the snout and measures from 60 to 65 mm. in

diameter.

The parietal crest forms a huge arch over the top of the head

about 300 mm. in length. Itis very narrow for its depth, and

is mainly formed by the postorbitals and the squamosals, the

parietals being relatively small. Near the union of the anterior

and middle third the two postorbitals are pushed apart by an

enormous development of the preparietal. This forms a pro-

minent boss 90 mm. in length and 50 mm. in width, and rising

20 mm. above the edge of the postorbitals. In it is situated the

* See also Addendum, p. 875.

NEW FOSSIL REPTILES. 867

parietal foramen, which is only about 10 mm. in diameter. This

is the more remarkable in that in H. wniseries the foramen

measures nearly 20 mm. in diameter.

The zygomatic arch is very massive. The malar process of the

Jugal is less marked than in the smaller species. The squamosal

extends forwards to below the postorbital arch. Anteriorly the

zygomatic arch measures 65 mm. in depth and posteriorly

80 mm.

The lower jaw is very powerful and measures 400 mm. in

greatest length.

* ENDOTHIODON PLATYCEPS, sp.n. (P]. XCIII. fig. 19.)

This new species was also discovered by Mr. Whaits, near

Beaufort West. It is founded on a fairly complete but slightly

crushed and probably barely mature skull. The lower jaw is

complete, and the only important part missing from the skull is

the whole of the nasal region and the anterior part of the

frontal.

The greatest length of the skull is 275 mm., and the greatest

width about 215 mm.

The frontal region is flat, and the parietal crest instead of

rising up from this is continued straight back. There does not

appear to be a distinct postfrontal bone. The preparietal is large

and passes forwards for 35 mm. in front of the pineal foramen.

The foramen is 14 mm. wide and 8 mm. in antero-posterior

diameter. Behind it is a large boss formed by the preparietal.

There is nothing particularly noteworthy about the squamosals,

jugals, or occiput.

The dentition is better seen in the lower than in the upper

jaw. It is made up of a series of 7 teeth in a fairly regular row.

Behind this the same row has 9 more teeth, but median to these

there appear others, so that the front half of the dentition is an

irregular single row, the back half an irregular double row.

Parts of the crowns of two teeth are preserved and there appear

to be no serrations on them. If this prove to be the case, this

species will require to be placed in a new genus, as Hndothiodon

bathystoma has the teeth markedly serrated in front and behind.

The lower jaw is pointed in front, but at the lower part of the

symphysis it is very broad. The back part of the dentary is

unusually slender. Hndothiodon platyceps differs from E. uniseries

and #. whaitsi in the double row of teeth behind and also in the

flat broad head; while from Z. bathystoma it differs even more

markedly. ‘he teeth are smaller than in the other species.

PRODICYNODON BEAUFORTENSIS, sp.n. (Pl. XCIII. fig. 21.)

This small species is founded on an imperfect skull obtained by

me at Kuilspoort at about the same horizon at which Taognathus

megalodon was found. The specimen consists of the crushed ant-

orbital portion of a small skull with the front two-thirds of the

* See also Addendum, p. 876.

868 DR. R. BROOM ON

lower jaw in position. At first sight the skull might readily be

taken for that of a small Oudenodon, but there are two marked

differences. There are a number of small maxillary teeth, and

the lower jaw ends in front in a pointed beak which fits into a

deep depression in the premaxillary exactly as in Hndothiodon.,

There is no tusk.

The nostril is 8 mm. in length by 7 mm. in depth. The

orbit is 12 mm. behind the nostril. The inter-orbital width is

probably 14mm, Close to the outer edge of the maxillary bone

are at least two small smooth pointed molars, and other molars

are arranged in a row further in.

The lower jaw is almost typically Endothiodont.

The only form nearly related to the present one is that de-

scribed by me eight years ago as Prodicynodon pearstonensis. The

two agree in the arrangement and structure of the molars, but

differ markedly in the proportions of the head. P. pearstonensis

has a much broader snout and an enormous premaxillary. It

also has the orbit much further back. As neither type is in

good condition it is impossible to affirm with certainty that the

two species belong to the same genus, but at present it will be

convenient to keep them together,

When Prodicynodon was first described, and until recently, the

presence or absence of a tusk was believed to be a generic dis-

tinetion. Dicynedon and Oudenodon were believed to be distinct

genera, For the last five years the evidence has been steadily

accumulating in favour of the tusk being merely a sexual character,

and now the evidence seems to be conclusive. Mr. Whaits has

collected a large number of the common little Beaufort West

Endethiodont Dielurodon whaitsi, and tusked and tuskless

specimens seem to be about equally common, while there are no

other characters te separate the specimens. Mr, D, M.S. Watson

has succeeded in obtaining two specimens of Oudenodon bolorhinus

recently described by me from Kuilspoort, at the same locality

as afforded the type, but one specimen is tusked and the other

tuskless,

This conelusion will necessitate the giving up of the genera

Oudenodon and Opisthoctenodon and placing the species of these

old genera under Dicynodon and Pristerodon, Fortunately very

little confusion will result,as I have for years assumed the

possibility of the tusk being merely sexual,

DicyNopon LATICES, sp. n. (PL XCII. figs, 12, 13.)

This new species of Dicynodon is founded on a beautiful skull

obtained by Mr. J. H. Whaits on the Nieuwveld. With the

exception of the lower jaw being missing and the tips of the

maxillaries with most of the tusks being broken off, the skull may

be regarded as perfect. It belongs to the very unusual broad-

headed variety of which only a few specimensare known. When

viewed from above the resemblance is so close to Dicynodon

figriceps Owen as to suggest that it might be a young specimen,

NEW FOSSIL REPTILES. 869

but the palatal view shows that the tusk is differently placed and.

relatively very much larger.

The greatest length of the skull measured obliquely from the

snout to the back of the squamosal is 270 mm., and the greatest

width across the squamosal is also 270 mm.

The snout is very short and the nostrils completely roofed over

by the projecting nasals. The breadth across the nasals is 83 mm.

The premaxillary is broad and shallow. The maxilla is short and

also shallow. The tusk is large and situated right beneath the

orbit, and directed downwards. In diameter it measures 25mm.

by 28 mm. at its base. The frontal region is broad and flat, the

narrowest part between the orbits being 63 mm. If there is &

distinct postfrontal it is very small. The parietal foramen is

large, measuring 15 mm. by 10 mm. The preparietal lies mainly

in front of it. The parietals are unusually large, and the posterior

branch of the postorbitals more slender than in most species.

The postorbital forms the front half of the long slender post-

orbital arch. The squamosal extends forwards below the post-

orbital arch and has a large articulation with the maxilla. The

pterygoids where they meet are broad and, except for the median

ridge, flat. Considering the great width of the skull the quadrates

are not far apart.

The general structure of the skull will be better understood

from the illustrations given.

DicyNopon psirracors, sp.n. (PI. XCII. fig. 17.)

In working at the troublesome genus Dicynodon we have con-

stantly been in doubt as to whether the small Dicynodon specimens

are distinct species or only younganimals. Frequently they agree

sufficiently in general shape and structure as to suggest the

probability of their being young specimens of Dicynodon

leoniceps or some other large species. But this is certainly not

always the case. Many specimens of Dicynodon jouberti are

known from the Pareiasaurus horizon and all small, while no

large Dicynodon is known to occur in the same zone. At Beaufort

West a considerable number of specimens of a small Dicynodon

also occurs, but there is no evidence of any large Dicynodon

having lived at the period. The large Anomodonts are the

Endothiodons. There is thus satisfactory evidence that the

common Dicynodon of the Beaufort West commonage is an adult

animal, and apparently a new species. The best specimens are a

good skull with much of the skeleton and a fairly good skull with

nearly the complete skeleton. I take the latter as the type.

Both specimens were obtained by Mr. J. H. Whaits.

The greatest length of the type skull is 102 mm., and of the

second specimen 112 mm. The width of the type across the

squamosals is about 66 mm. Across the maxilla the maximum

width is 42 mm. The interorbital width is 20 mm., and the

intertemporal 18 mm.

The following are the most noteworthy characteristics of the

Proc. Zoon, Soc.—1912, No. LVIIT. 58

870 DR. R. BROOM ON

species. The nasals are so narrow that the nostrils look almost

directly upwards. The upper part of the nasals is thickened,

and the prefrontal region of the orbital margin is also elevated.

The frontal region is broad and flat. The parietal foramen is

situated in an elevated preparietal. The postorbitals approach

each other behind the foramen and nearly touch, forming a

parietal ridge. The tusk is small and directed forwards and

downwards. The lower jaw has the front portion unusually

broad and deep. The foramen behind the dentary is very

small.

The whole skeleton from the snout to the end of the tail

probably measures 500 mm.; the humerus measures 50 mm. and

the femur 58 mm.

DICYNODON LUTRICEPS, sp. n. (Pl. XCII. figs. 14-16.)

The type of this new species is an imperfect skull found by me

at Kuilspoort, Beaufort West district. The skull has lost the tip

of the beak, the postorbital and zygomatic arches, and there is

about 20 mm. missing from the postpterygoid and from the

parietal region so that the contact between the occipital and

anterior portions of the skull is lost, but otherwise the skull is

complete.

The most noteworthy characters of the type are the relative

shortness of the beak, the broad concave frontal region, and the

broad flattened intertemporal region, the upper surface of which

is almost entirely formed by the postorbitals. In a maxillary

from the same locality, and believed to be of the same species, the

tusk is feeble and directed downwards and slightly forwards.

The greatest length of the skull from the snout to the back of

the squamosa] is about 195 mm., and the greatest width across the

squamosals is 145 mm. From the front of the beak to the orbit

is probably about 48 mm., and the antero-posterior diameter of

the orbit is about 43 mm. The interorbital width is 35 mm.,and

the intertemporal region 29 mm.

The type specimen is tuskless. The caniniform process is

directed downwards and forwards and has a marked low outer

ridge which passes upwards towards the jugal arch. Below the

nostril is an anterior ridge parallel to the other and forming a

well-marked valley between the two. The nostril is fairly large

and the nasal is considerably thickened above it, forming an

overhanging supranasal ridge.

The prefrontals are small, but the frontals are well developed.

Posteriorly they enclose between them the small preparietal and

meet the anterior ends of the parietals. The postfrontals are

long and narrow. ‘The preparietal is small and seems to lie

entirely in front of the pineal foramen. ‘The postorbitals are

very large where they overlap the parietals, but the postorbital

arch is unusually feeble.

The parietals are large and powerful, but are almost completely

NEW FOSSIL REPTILES. 871

hidden by the postorbitals. There has been a little doubt as to

whether the pair of bones usually supposed (Seeley, Broom, etc.)

to be parietals are really the parietals, or whether the median

bone in which lies the pineal foramen, and usually called the

preparietal (Seeley, Broom) or interparietal (Newton), may perhaps

be the true parietal. This latter view has recently received the

support of Jaekel. The median bone, called for convenience

preparietal, is met with in most Anomodonts. In some (Zndo-

thiodon) it is very large; in others (Cistecephalus) it is quite

absent. The size of the paired bones depends to a considerable

extent on the development of the preparietal. In Cistecephalus

there can hardly be any doubt that the large pair of bones behind

the frontals are the parietals. They have the same relations to

the squamosals, interparietal, frontals, and postorbitals as the

parietals have in most reptiles, and there can be, I think, no

reasonable doubt but that these bones are homologous with the

parietals of the mammals. When the preparietal appears and

the intertemporal region becomes narrowed the parietals are

much reduced in front, but posteriorly the relations to the

squamosals, interparietal, and postorbitals remain constant. In

Endothiodon the preparietal is so large that the parietal seems to

be completely separated from it by the frontal. In Dicynodon,

as exemplified by this skull, the parietals still meet the frontals.

What the preparietal is, is not clear. It certainly is not the

interparietal. J am inclined to look on it as a neomorph

developed in connection with the pineal eye. There is no trace

of it known in Dinocephalians, Dromasaurians, Pelycosaurs,

Therocephalians, or Cynodonts, though in some of these the

pineal eye was probably as large as even in Endothiodon.

There is little of special note in the palate or occiput.

The horizon from which the specimen was obtained is probably

about 300 feet above that of Beaufort West.

Emypbops MINoR, gen. et sp.n. (Pl. XCIII. fig. 20.)

When Owen, in 1876, described the specimens of Cistecephalus

in the British Museum, he named one species Kistecephalus

arctatus and referred two specimens to it. The type differs very

considerably from Cistecephalus microrhinus, the type species of

the genus, and Lydekker in his Catalogue places C. arctatus

doubtfully under Cistecephalus. On more than one occasion I

have also expressed the opinion that C. arctatus does not belong

to Cistecephalus.

Recently, I discovered at Kuilspoort a small imperfect skull

which apparently belongs to the same genus as Owen’s C. arctatus,

though a distinct species. Pretty certainly the genus is not

Cistecephalus, and the question arises, is it Owdenodon, or rather

Dicynodon? The only specimens known are tuskless, and there

are apparently no molar teeth. In the imperfect state of the

specimens it is impossible to clearly differentiate the genus from

58*

872 DR. R. BROOM ON

Dicynodon at present, but there seems little doubt they represent,

if not a distinct genus, at least a subgenus. The more note-

worthy characters are the wide parietal region with large

parietals, slender postorbital arch, and feeble beak.

The length of the skull is probably about 45 mm., and the

greatest breadth about 30 mm. The orbit measures 12 mm. in

diameter. The intertemporal region is 14 mm. across.

The beak is in very imperfect condition, and little can be made

out with certainty as to its structure. It may be stated with

confidence that it was short.

The frontals are large, and form an interorbital region 10 mm.

wide. From near the supraorbital margin to the anterior end of

the parietal there runs backwards and slightly inwards a shallow

groove. There is a moderate sized triangular postfrontal and

a large median preparietal. ‘This latter seems to le entirely in

front of the pineal foramen. The parietals are large and form

the greater part of the broad intertemporal region. The post-

orbital is long and slender. It forms a feeble postorbital arch

and the inner margin of the temporal fossa. The squamosal is of

the typical Anomodont type. Its zygomatic position extends

forwards to below the orbit. The articular region is badly

preserved.

The lower jaw is very like that of Oudenodon, but the beak

portion is small, and probably little more than the symphyseal

region was covered with horn.

Suborder CYNODONTIA.

TcTIDOPSIS ELEGANS, gen. et sp.n. (Pl. XCITI. fig. 22.)

This new genus and species is founded on a nearly perfect little

skull obtained at Harrismith, Orange River Colony. It is a very

near ally of Vythosaurus larvatus Owen, but is much smaller, and

differs in the number of molars and in other cranial characters.

In general shape the skull agrees fairly well with Vythoswurus.

The orbit is near the middle of the skull and relatively larger than

in the better known genus, while the jugal arch is more slender.

In Jctidopsis the snout is shorter, and the molars are 6 in number

instead of 7.

The greatest length of the skull is probably 63 mm. and the

width is 42 mm. The interorbital width is 12°5 mm.

The premaxillary bone is badly preserved, but it is manifest that

there are four incisors. The first three incisors are moderately

round, but the last is more flattened. There do not appear to

be any posterior serrations.

The maxillary is relatively shorter than in Vythosaurus, and

deeper. Above the canine, and in. front of the lachrymal are

little elevations of the bone. On the canine elevated area are

three small foramina and near the root of the 3rd molar two other

NEW FOSSIL REPTILES. 873

foramina, while two more are near the anterior end of the bone.

As there is no single large supra-maxillary foramen, it is probable

that all these small foramina are for branches of the maxillary

nerve. The canine is long and slender. It is ridged somewhat

after the manner of the canine of the cat. Behind it are 6 molars.

The Ist is small, pointed, and without any cusps. The 2nd, 3rd,

4th, and 5th all closely resemble one another. There is a large

pointed median cusp and a small anterior and posterior cusp.

While essentially similar in type to the molars of Vythosaurus

they differ in that the anterior and posterior cusps are relatively

smaller. The 6th molar is a small tooth, and unfortunately the

crown has been lost from both sides of the skull.

The incisors measure about 6°5 mm. Behind the last incisor

is a diastema of 4mm. The canine measures antero-posteriorly

about 2°5 and its height is 7 mm. Ata distance of 1-5 mm.

behind the canine is the lst molar, and the whole series of six

occupies the space of 13 mm.

Only a small fragment of the septo-maxillary is preserved, but

it manifestly forms part of the face, and was probably as in

Nythosaurus.

The nasal is moderately wide in front, but narrows on passing

backwards, and then near its middle it becomes about twice as

wide as in front. Round the bone near where it meets the

maxilla is a series of three or four foramina.

The lachrymal forms the front of the orbit and, as in Vytho-

saurus, it is larger than the prefrontal.

As in NVythosaurus and most Cynodonts the prefrontal meets

the postorbital, shutting out the frontal from the orbital margin.

The postorbital forms about half of the postorbital arch and

overlaps a small part of the parietal.

The parietal is large, and there is an obvious pineal foramen.

The squamosal is like that of Wythosaurus, except that in

Ictidopsis there is a much more prominent auditory groove.

The fractured edge of the occipital crest shows the interparietal

distinct from the parietals, and the lateral bone, which I believe

to be the opisthotic, distinct from the parietal, the interparietal,

and from the squamosal.

The occiput and palate have not been cleared.

The dentaries are in position, but the posterior bones of the

jaw have been detached and displaced, probably by insects before

the skull was fossilized.

Thety pe of Ictidopsis elegans wasfoundat Harrismith in association

with Lystrosaurus. Unfortunately the geology of Harrismith is

unknown. Some of the first specimens of South-African Dinosaurs

were got there by Mr. J. M. Orpen in 1853, and there can be no

doubt that these are from the Red Beds of the Stormberg Series,

It is not a little startling to find that the same commonage

yields fossils which in Cape Colony belong to a horizon about

3000-4000 feet lower than the Red Beds. Probably the Molteno

874 DR. R. BROOM ON

Beds and the Burghersdorp Beds are greatly thinned out, or

possibly there is an unconformity.

Some years ago I divided the Upper Beaufort, or Triassic Beds,

into three zones: (1) the Lystrosawrus-Zone, (2) the Procolophon-

Zone, and (3) the Cynognathus-Zone, While these zones in the

main hold good, the limits of them are still unknown. In the

extensive Lystroswurus beds of Colesburg, Middelburg, and

Cradock no Procolophons or Cynodonts are known; but Mr. D.

M. 8S. Watson has recently found Lystrosaurus associated with

Cynodonts to the west of Burghersdorp, and a similar association

we now know occurs at Harrismith. Again, while no Lystro-

saurus or Cynodont remains are certainly known from the

Procolophon beds, the Procolophon-like genus Thelognathus occurs

at Aliwal North with Cynodonts, and possibly Procolophon itself.

It seems not improbable that later work will further subdivide

the Triassic Beds, each zone counting from the point where a

new type begins, but before we can make any further advance

with confidence we require to have a much fuller knowledge of

the distribution of the fossils of the Upper and Middle Trias.

In the meantime I think we are safe in stating that Vytho-

saurus, Ictidopsis, and probably Galesaurus come from an sities

zone than the Cynognathus beds. None of the known specimens

have been found near Burghersdorp, nor have any traces of

Lystrosaurus ever been found, from which we may conclude that

Lystrosaurus and probably these small Cynodonts became extinct

before Cynognathus appeared.

NyTHOSAURUS BROWNI, sp.n. (PI. XCIII. fig. 23.)

This new species is founded on an imperfect lower jaw obtained

by Mr. Alfred Brown at Aliwal North. A large part of both

dentaries is present, but most of the symphyseal portion is missing

with the canines and incisors. Five molars are well preserved on

the left side and three on the right.

While in a number of respects the specimen differs from

Nythosaurus larvatus Owen, it seems probable from the position

of the symphysis that there were seven molars, and as in general

structure the molars agree with those of Vythosaurus larvatus, we

may consider it as probable that the species belongs to this genus.

The dentary differs from that of Vythosaurus larvatus in haying

a more slender horizontal ramus, in having a more marked angle,

and in the ascending ramus passing more upwards.

The molars preserved are probably the 3rd to the 7th. All

have three cusps. What is apparently the 3rd molar has the

median cusp short, and the others only feebly developed. In the

Ath and the other later molars the middle cusp is about twice as

long as the anterior and posterior cusps. In the second last

molar there is an additional small anterior cusp which gives it

four cusps, but there appear to be only three cusps in the last

molar. The five molars occupy 13°5 mm.

NEW FOSSIL REPTILES. 875

Though the crowns of the molars resemble considerably those

of the Triconodont mammals there is the marked difference that

in NVythosaurus there is no trace of a cingulum, and there is

apparently only a simple root to each tooth.

Apprnpum (29th July, 1912).—Since the above paper was

read evidence has been obtained which shows that the genus

Endothiodon must be subdivided. Seeley twenty years ago

placed Hndothiodon uniseries in a distinct genus, Esoterodon, and

until quite recently I have followed him in this. A fragmentary

maxilla recently found by Mr. Whaits shows that Seeley was

correct. In Endothiodon bathystoma the teeth have long pointed

crowns with coarse serrations both in front and behind. In

Endothiodon wniseries the teeth have flattened crowns serrated

only behind, and in Zndothiodon platyceps there are no serrations

on either side. This latter point has been confirmed in a second

specimen. Until the crowns of the teeth of Endothiodon whaitst

are known we may provisionally place it with Hndothiodon wii-

series. he group would thus be formed of

Endothiodon bathystoma Owen.

'Hsoterodon uniseries Owen.

Esoterodon whaitst Broom.

Emydochampsa platyceps Broom, gen. nov.

EXPLANATION OF THE PLATES.

PuatE XC.

Pig. 1. Side view of snout of Tawrops macrodon Broom. Nearly } nat. size.

Fig. 2. Side view of tooth of probably Eecasaurus priscus Broom. Nat. size.

This tooth, though not associated with the type of Hecasaurus priscus,

is trom the same horizon, and as it is the tooth of a Dinocephalian of

the sizeof Hecasaurus it very probably belongs to this genus and species.

Fig. 8. Upper view of tooth of probably Wecasaurus priscus Broom. Nat. size.

Fig. 4. Side view of skull of Scymnognathus whaitsi Broom. A little over $ nat.

size. The skull is much flattened and distorted by crushing.

Fig. 5. Lower jaw of Scymnognathus whaitsi Broom. % nat. size. Though this

jaw is rather smaller than that of the type specimen it almost certainly

belongs te this species. It may have belonged to a young animal or to a

female. ‘The specimen is considerably crushed.

PuatEe XCI.

Fig. 6. Side view of the skull of Galeops whattsi Broom. Nat. size. The

specimen is slightly crushed and considerably weathered. The orbital

margins and the preorbital portion of the skull and the lower jaw are the

outer surface of the bones viewed from within. ‘The back portion of the

skull shows the quadrate and parts of the squamosal and opisthotic. A

sclerotic plate is seen in the orbit.

Fig. 7. Side view of snout of A2lurcsaurus striatidens Broom. Nat. size. The

specimen is considerably crushed.

Fig. 8. Side view of snout of Pristerognathus platyrhinus Broom, 5%; nat. size.

Fig. 9. Under view of snout and lower jaw of Alopecorhinus parvidens Broom.

42 nat. sine.

Fig. 10. Upper view of imperfect skull of Ictidognathus hemburyi Broom.

Nat. size.

Fig. 11. Under view of snout of Ictidognathus hemburyt Broom. Nat. size.

876 PROF, S. J. HICKSON ON THE

Puate XCII,

Fig. 12. Upper view of skull of Dicynodon laticeps Broom. 5 nat. size.

Fie. 13. Under view of snout of Dicynodon laticeps Broom. -; nat. size.

Fig. 14. Side view of skull of Dicynodon lutriceps Broom. %s nat. size. The

: contact between the front part of the skull and the occiput is missing,

but the relation of the parts must be almost as restored.

Fig. 15. Upper view of skull of Dicynodon lutriceps Broom. About 7% 15 nat. size.

Fig. 16. Palatal view of skull of Dieynodon lutriceps Broom. About 5% 16 nat. size.

Fig. 17. Upper view of skull of Dicynodon psittacops Broom. About 4 nat. size.

Prater XCIII.

Fig. 18. Side view of skull of Hndothiodon whaitsit Broom. About 2 nat. size.

Fig. 19. Upper view of lower jaw of Hndothiodon platyceps Broom. About 35

nat. size.

Fie. 20. Upper view of skull of Hmydops minor Broom. 2 nat. size.

Fig. 21. Side view of skull of Prodicynodon beaufortensis Broom. About 7% nat.

size.

Fig. 22. Upper view of skull of Ictidopsis elegans Broom. 2 nat. size.

Vig. 23. Side view of left dentary of Nythosaurus browni Broom. % nat. size.

53. On the Hydrocoralline Genus, Errina. By Professor

S. J. Hicxson, F.R.S., F.Z8., The University of

Manchester.

[ Received April 29, 1912: Read June 4, 1912.]

(Plates XCIV.-XCVL)

INDEX.

Page

Revision of the genera Hrrina, Labiopora,

Spinipora ......... eau cael SeRGEn ea eRe aU,

Description of new species . sais . 882

Notes on the structure of rn NOVE 1, eelaniiens 884

Note on geographical distribution ..................... 886

Description of old species ..........:eeseeeeereeeeeesees 887

General remarks on structure ................:000000002. 891

List of Known Species ..........ccceseeeeeeeeneeeeeeessene 893

Additional motes Ness. jecssasene eee eeceee eee eeeee nes cee OO

Literature ...... SARE ERRORS 94

Explanation aE he Plates SBS GRBI an apnacocasenmuseauss pete

Four years ago Professor Benham sent to me some specimens

of Stylasterina from New Zealand, with a request that I would

name them for him. At the same time he forwarded some notes

and drawings which have proved to be of considerable assistance

to me in working out their details. JI am also indebted to

Professor Dendy for the loan of another specimen also from New

Zealand; to the late Mr. Morgan, of Worthing, for the loan of a

specimen from an unknown locality; and to Mr. Gilchrist for

a specimen from the Cape of Good Hope.

All these specimens clearly belong to one of three genera,

Errina, Labiopora or Spinipora, as they exhibit the characters

that these genera exhibit in common and by which they can be

separated from other Stylasterima. These characters are: (1) a

P.Z.8.1912, Pl. XCIV.

Huth, coll.

EY OD) is Orn ei Ge NUS) ie REINA

Piao. ole, Ply xev.

we 8.

Huth coll.

BYDHOIDS OF THE GENUS ERRBRINA.

i al}

oo fe

ee beans Ga

itaa) | e

of 2 4 3g] |e @

Hoaa ee e ie)

a2ean Wl [GPK ec

Bex e eo §

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VF eo Va 0 2

5 4 og Pee

> hl 5 @aas

i Ze 2 & %

3 i Pete.

Wel 7 9@e a

if ae ga

Et (9 Oe @e-

Ph 11.09 2-0e@

po ao Vf

Ni 3° af

Gye n @P J

mn ( [a Gse

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Nn ~ePea

hi eo °@.@

14.

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a we =~ F-@ oe oes

Warn:

BMD OIDS. Of THE GENUS” MRRINA.

P Z.S,1912. Pl. XCVI.

tie

HYDROCORALLINE GENUS ERRINA, 877

style in the Gasteropores ; (2) no style in the Dactylopores; and

(3) the Dactylopores, or some of the Dactylopores, guarded by a

grooved lip or spine—called by Moseley the “ nariform process.”

To any one acquainted with Moseley’s classical memoir of the

Stylasterina (8), and with the subsequent literature, which is not

very extensive, it might seem a simple matter to determine

whether the specimens about to be described belong to any one

of the three genera -or not; because, according to Moseley’s

descriptions, the following characters were diagnostic :—

Errina, with only one kind of Dactylopore.

Lahbiopora, with two kinds of Dactylopores. The larger kind

of dactylopore with a nariform process * and arranged in

rows. The smaller kind of dactylopore, without a nari-

form process, arranged between the rows.

Spinipora, also with two kinds of Dactylopores. The larger

kind of dactylopore with long grooved spines, not arranged

in rows. The smaller kind of dactylopore at the base of

the larger ones.

Moseley’s memoir was published in 1881, and since that date

only four new species of Hrrina and one new species of Labiopora

(LZ. moseleyi) have been described; but owing to the rarity

of these Hydrocorallines in the seas that have been recently

investigated, very little progress has been made in our knowledge

of them.

The specimens from New Zealand and elsewhere that I have

examined convinced me that a thorough revision of the genera

was necessary, and consequently the task of naming Professor

Benham’s specimens has taken me much longer than I anticipated.

The general results of my investigations have been to show that

the limits or frontiers between the three genera are ill-defined

and that it is necessary to accept von Marenzeller’s (6) proposal

to unite Labiopora with Hrrina.

In the first place, I have found, as von Marenzeller (6) has

done, that the presence of two kinds of dactylopores is not a very

reliable character; because in some forms that are otherwise

closely related the dactylopores of the smaller kind that have no

grooved spines may be numerous, scarce, or altogether absent

(ce. g., Errina nove zelandiw). A genus such as Labiopora

cannot therefore be absolutely separated from Hrrina by the

character of the dimorphism of the dactylozooids.

Moreover, the arrangement of the grooved spines in definite

rows is another character that is subject to considerable variations

and cannot be relied upon for diagnostic purposes. The arrange-

ment of these processes seems to be correlated in some way with

the method of growth of the hydrophytum as a whole, and that

is again, I believe, dependent upon the conditions of the en-

vironment.

* The term “nariform process” introduced by Moseley is not very convenient, and

T have consequently used the the expression “grooved spine” for the coenosteal

processes that shelter the dactylozooids.

878 PROF, S. J. HICKSON ON THE

The characters that appear to me of more importance and to

signify a more profound differentiation are based upon :—

1. The texture of the surface of the corallum.

2. The aspect of the grooves in the nariform processes.

Moseley described the surface of Hrrinw as composed of a

compact, hard, glistening, white, calcareous tissue, and he adds

that the canals generally are in this genus larger in proportion to

the size of the zooids than in most other forms, and the mesh-

works formed by them are comparatively wide open. Associated

with these two characters we usually find that the surface is

marked by shallow longitudinal grooves perforated by a series

of small apertures through which the vertical canals pass. These

ccenosteal pores are well defined in the species of this group that

I have examined, and in the type specimens of &. labiata and

LH. ramosa they are about ‘05 mm. in diameter. The surface of

Spinipora is hard and compact as in Hrrina, and it is also

perforated by well defined ccenosteal pores.

In the eriginal description of the genus Labiopora, Moseley

describes the ccenesteum as being ‘‘ minutely reticulate in tex-

ture,” and in the description of Labiepora moseleyi Ridley (10)

also describes the surface as being “ minutely reticulate.” In all

the specimens I have examined there is a very marked contrast

between the surface of the specimens now included in the Zabio-

pora group of species and of those I propose to retain in the

Hrrina group.

I should prefer to describe the surface of the former group of

species as being “ granular” rather than “reticulate.” In Zrrina

(Labiopora) capensis it is coarsely granular and in all the other

species of the group it is minutely granular (cf Pl. XCVI.

figs. 14 & 15). Below the surface, the ccenosteum is minutely

reticulate, being perforated by a network of small-meshed canal

passages (fig. 14), in contrast with the wide-meshed canal

passages of the Zrrina group.

As regards the grooved spines. In the following species that

I have examined the groove in the spine that protects a dactylo-

pore is turned towards the apex of the branch on which it is

situated: H. labiata, H. horrida, and #.ramosa. The groove has

the same aspect in #. glabra and £. carinata, if we may judge

from the figures given by Pourtales (9).

In the only two specimens of the genus Spinipora that are

known the grooves also turn towards the apex of the branch

(Pl. XCV. fig. 8). In the Labiopora group of species there is

considerable variation in the direction of the grooves. In some

of them, all or nearly all the grooves are turned away from the

apex (Pl. XCVI. fig. 11), but in others the grooves are turned

in all directions, the grooved spines forming irregular clusters on

the surface (Pl. XCVI. figs. 12 & 13). The same arrangement of

the grooves occurs in Lrrina gracilis and in Errina macrogastra,

according to von Marenzeller, who writes ‘“‘Ihre Oeffnung ist

nach hinten gerichtet, selten seitlich” (5), and also in Errine

Jissurata of Gray (2).

HYDROCORALLINE GENUS ERRINA. 879

Taking these characters as guides it seemed to me that the

diagnostic characters of the three genera might be stated as

follows :—

(a) Coenosteum hard and compact, perforated by well-

defined ccenosteal pores. Grooved spines turned

towards the apex of the branch.

1. With short grooved spines and only one kind of

daetylopope tetas Me. focal dasis ate thens Errina.

2. With long grooved spines and two kinds of

dactyloponeset seis AP ie J ae oe Spinipora,

(6) Ceenosteum granular and reticulate, without well-

defined ccenosteal pores. Grooved spines turned

away from the apex of the branches or irregularly

place Uhiiie.. sais cea eas eae Labiopora.

If the three genera be joined together to form a subfamily, the

Errinina, this subfamily might be defined as follows :—

Hydrophytum arborescent and irregularly flabelliform, gastero-

pores and dactylopores not arranged in cyclosystems. Gastero-

pores with a large brush-like style. Dactylopores without a

style. Some of the dactylopores protected by a grooved spine

(narial process) on the surface of the ccenosteum.

This arrangement of the genera, however, breaks down on

further analysis, and I see no cther course than to arrange all the

species in three groups under the one generic name Lrrina.

The genus Hrrina was founded by Gray in 1835 for a species

of coral found in the Mediterranean Sea and formerly called

Millepora aspera by Linneus.

As von Marenzeller has pointed out, Gray’s description of the

Spines in this species as “‘Superne longitudinaliter fissee”’ is not

consistent with the description of the species known to Linneus

and Esper.

I have examined the type specimens in the British Museum,

and have found that Gray’s description is not correct. The

spines in these specimens are irregular in arrangement, but where

they are isolated and not in clusters the groove is directed away

from the apex. Moreover, the character of the surface of the

ceenosteum, the presence of a few small dactylopores without

grooved spines, and other features prove that this species is more

closely related to the type species of Labiopora than it is to any

of the other species of Hrrina.

According to the system I had, at first, proposed the type

species of Hrrina would thus be a species of Labiopora and

Moseley’s Hrrina ramosa would become the type species of the

genus.

Such a proposal, therefore, would not only be contrary to the

rule of zoological nomenclature, but it would also be extremely

inconvenient. Moreover, one species at least (7. macrogastra)

would oceupy an intermediate position, having a surface similar

880 PROF. S. J. HICKSON ON THE

to that of Hrrina and grooved spines similar to those of

Labdiopora.

A plea might still be made to keep the genus Spinipora distinct,

but I am convinced that intermediate forms will be found

between the deep-sea species of Hrrina and the only known

species of Spinipora, and that sooner or later it will be found

impossible to keep it apart from the others.

In the following pages, therefore, I have regarded all the

species that have been attributed to the three genera as belonging

to the one genus, Hrrina of Gray, but for convenience of reference

I have added after the generic name (Labiopora) or (L.) in the

case of those species that were formerly described as belonging to

the genus Labiopora and to others that belong to that group of

species, and (Spinipora) or (S.) in the case of Hrring echinata, the

only known species of the Spinipora group.

Genus Errina Gray.

With the characters of the subfamily Errinina (p. 879).

The “ Hrrina” group of species.

Ceenosteum hard and compact, perforated at the surface by

well-defined ccenosteal pores usually arranged im rows in shallow

longitudinal surface-grooves. Gasteropores with or without a

scale. All the dactylopores protected by short grooved spines

(narial processes) with the grooves turned towards the apex of

the branch.

The only species of this group that have been sufficiently well

described to make identification possible without reference to the

type specimens, are Hrrina labiata Moseley and ZH. ramosa

Hickson & England and 2. horrida H. & EH. Other species

are rrina carinata Pourtales and #. pourtalesit Dall.

Pourtales in 1871 described three species, which at first he placed

in the genus Hrrina but subsequently transferred them to a new

genus, Lepidopora. These species were referred back again to

Ervina by Moseley. Their names are #. glabra, H. cochleata, and

E. dabneyi. These three species were distinguished from Hrrina

by the presence of a lip or lid-like process similar to that of

Cryptohelia hanging over the gasteropores. Hrrina fissurata of

Gray may have been a specimen of Labiopora, but as the original

specimen has been lost it is useless to speculate on its supposed

affinities.

The species may be arranged as follows :—

* q@, Gasteropores with a definite lip or scale :—

E. glabra, E. cochleata, LH. dabneyi, and HL. ramosa.

b. Gasteropores in the angles formed by the branches :—

LE. horrida.

c. Gasteropores without scales and distributed on the surface

of the ccenosteum :—

E. labiata, E. carinata, EF. pourtalesii (?).

* See Note p. 894.

HYDROCORALLINE G2NUS ERRINA, 881

As regards the distribution it may be said that all the species

are inhabitants of deep water (i. e. 50-600 fathoms). Being

deep-sea species they are probably widely distributed, but at

present 2’. pourtalesii, H. ramosa, and HL. horrida have only been

found in the Pacific Ocean and Malay Archipelago, and the

remaining species in the Atlantic Ocean.

Spinipora group of species.

Ceenosteum hard and compact, perforated at the surface by

well-defined ccenosteal pores usually arranged in shallow branching

grooves irregularly arranged on the surface. Gasteropores

without a scale. Dactylopores of two kinds. The larger kind

being guarded by long grooved spines (narial processes) crowded

and overlapping on the terminal branches, often worn down

short and separated by considerable intervals on the stems and

older branches (Pl. XCV. fig. 8). All the grooves of these

spines turned towards the apex of the branch. Smaller dactylo-

pores not protected by grooved spines, but scattered between and

on the projections that guard the larger dactylopores.

The Spinipora group is represented by only one species,

Errina (S.) echinata Moseley, and this species has been found off

Rio de la Plata in 600 fathoms by the ‘ Challenger ’ Expedition,

and off Providence Island in the Indian Ocean in 75 fathoms by

Professor Stanley Gardiner (4).

Labiopora group of species.

Ceenosteum granular and minutely reticulate, without clearly

defined ccenosteal pores. Gasteropores without scales but some-

times protected by short grooved projections or “lips.” Dactylo-

pores protected by grooved spines of variable lengths, the grooves

turned away from the apex of the branch or irregularly in all

directions. Some of the dactylopores usually without grooved.

spines.

The species of this group that have been described already

are Errina aspera Gray, Errina (Labiopora) antarctica Gray,

Errina (L.) moseleyi Ridley, and Lrrima gracilis von Maren-

zeller.

Errina aspera was the name given by Gray to the Linnean

species Millepora aspera from the Mediteranean Sea. Hrrina (L.)

antarctica was originally described by Gray as a Polyzoon

(Porella antarctica), but was subsequently redescribed and figured

by Moseley as the type species of Labiopora. The type specimen

was found off the East coast of Tierra del Fuego, but a second

specimen was discovered by the ‘ Alert’ Expedition in 30 fathoms

off S.W. Chili and described by Ridley (10). Hrina (L.) mose-

leyi was found by the ‘ Alert’ Expedition off the same cosat in

2-10 fathoms. Lrrina gracilis was found off the pack ice in the

Antarctic Ocean in deep water (von Mareuzeller) (6).

I have had an opportunity of carefully examining a piece of

882 ' PROF. S. J. HICKSON ON THE

the type specimen of Hrrina (L.) antarctica and comparing it

with the specimens from various localities mentioned at the

beginning of this paper. I have found, as I expected, the same

difficulties in the determination of species that are met with in

the systematic zoology of other zoophytes. My impression is

that all the specimens from New Zealand belong to one distinct

species, that the specimen from the Cape of Good Hope belongs

to another distinct species, and that both these species are distinct

from the four species that have already been described. But

there is so much variation in the specimens from New Zealand

that it is clearly desirable to have a careful description of each.

I have also added for convenience’ sake a new deser spHOR YO of the

two earlier species for comparison.

Errina (LABIOPpoRA) NOV ZELANDIAZ. (Facies Ramosa.) (PI.

XCIV. fig. 3 & Pl. XCVI. fig. 9.)

This specimen was obtained from Preservation Inlet, W. coast of

South Island of New Zealand, in about 3 fms. of water, and was

lent by the Canterbury Museum to Prof. Benham. The colony

is flabelliform, with profuse ramification but without anastomoses.

The terminal branches are usually delicate. This may be ex-

pressed in figures by saying that at a distance of 3 mm. from the

extremity of a terminal branch the diameter may be not more

than 1 mm. The larger branches are slightly compressed in the

plane of the flabellum ; the others circular in section.

Colour : salmon-pink*.

Surface minutely granular, substance of the ccnosteum

minutely reticulate.

Grooved spines (narial processes) numerous, arranged in rows,

rarely in clusters. The groove, in nearly all cases, turned directly

away from the apex of the branch.

Gasteropores more numerous on one side of the flabellum than

on the other, sometimes provided with a lip. Diameter of gas-

teropores 0°27 mm.

Large dactylopores 0:06 x 0:16 mm. (The large kind of dactylo-

pores of the genus are protected by the grooved spines, and the

measurements given indicate approximately the width x depth of

the groove at its deepest part.)

Small dactylopores rare or absent. The small dactylopores are

often difficult to determine until the coral is thoroughly cleaned

by boiling in eau de javelle. I have examined and re-examined

a small branch thus cleaned and can find no small dactylopores,

but as I have only a small amount of material at my disposal,

and as Benham states in his MS. notes that the small dactylo-

pores are “rare,” I cannot deny their existence.

* There are so many shades of red to be found in corals that I have used the

technical term which expresses the shade of red that comes nearest to that shown

by this coral.

HYDROCORALLINE GENUS ERRINA. 883

| Erriva (Laprorpora) Nov# ZELANDI&. (Facies Benhami.)

(Pl XCIV. figs: 1&2); Pl. MOVE. fe! Dap)

This specimen was also found in Preservation Inlet and lent to

Prof. Benham by the Colonial Museum. One branch of this

Specimen was well preserved in spirit. The colony is flabelliform,

with profuse ramification and abundant anastomoses. The

terminal branches are thick, the diameter of such a branch at a

distance of 3 mm. from the extremity being about 3mm. All

the branches are approximately circular in section.

Colour : salmon-pink.

Surface minutely granular and substance minutely reticulate.

Grooved spines numerous, arranged roughly in rows. These

spines are more numerous and longer than in the facies ‘‘ Ramosa.”

They are, moreover, frequently arranged in clusters, so that they

have the appearance of “branched spines” (fig. 13). The

grooves are in general turned away from the apex, but when

the projections are clustered they turn in all directions.

Gasteropores equally numerous on the two sides of the flabellum.

Without a lip. Diameter varying considerably from 0°13 mm.

to 0°17 mm.

Large dactylopores 0:06 x 0°16 mm.

Small dactylopores not infrequent, 0:05 mm. in diameter.

ErRinA (LABIOPORA) NOV ZELANDI#. (Facies Dendyi.)

(Pl. XCIV. fig. 4.)

This specimen was obtained in Milford Sound, W. coast of

South Island, and was lent to me by Professor Dendy.

It is not very profusely branched, but probably has a flabelli-

form mode of growth. ‘The terminal branches are delicate and

of approximately the same diameter as those of facies “‘ Ramosa.”

There are no anastomoses in the specimen.

Colour: salmon-pink.

The surface is minutely granular and the substance minutely

reticular.

Grooved spines not very crowded but quite irregularly disposed,

not in rows, never in clusters. The groove in all cases turned

away from the apex of the branches.

Gasteropores on both sides of the branches but rather more

numerous on one side than the other. Usually guarded by a

small lip. Diameter 0°22 mm.

Large dactylopores 0:06 x 0-11 mm.

Small dactylopores 0°05 mm. in diameter, usually guarded by a

shallow collar or lip.

A feature of this specimen that should be mentioned is the

suppression of the spines on the larger branches. Only the

terminal branches are echinate.

884 PROF. §. J. HICKSON ON THE

Errina (LABiIoporaA) NoV# ZELANDI@. (Facies Cooki.)

(Pl. XCV. fig. 5; Pl. XCVI. figs. 10, 11, 12.)

This specimen was obtained from the cable in Cook Straits

between the two islands of New Zealand, and was lent to

Professor Benham by the Colonial Museum. According to

Professor Benham’s notes, two specimens were obtained, one

being 50 mm. in height x 70 mm. across, and has six main

branches; the other is smaller, 30 mm. in height x60 mm.

across. The general form of the cenosteum is “flabellate, the

main axis flattened but the branches circular in section. The

branches do not anastomose in the specimen examined. Only a

small piece of one of these colonies was sent to me, and from

that I have drawn up the following notes.

Colour: pure white.

- Surface and substance as in the other facies.

Grooved spines not very crowded and not arranged in definite

rows, frequently in clusters with the grooves pointing in all

directions (fig. 12).

Gasteropores evenly distributed on both sides of the flabellum,

without any lip or collar, 0°22 mm. in diameter.

Large dactylopores 0:06 x 0°08 mm.

Small dactylopores rare or very rare, 0°09 mm. in diameter.

In comparing these four facies of the species, several points of

interest may be observed.

They all agree in the general texture of the ccenosteum, and

they all have a more or less flabellate form of growth.

As regards the size of the hydrophytum as a whole, it is

impossible, owing to the broken condition of all the specimens

examined, to give exact measurements. A specimen of uncertain

facies in the Colonial Museum is 90 mm. x 70 mm. (according to

the MS. notes of Professor Benham), the specimen of the facies

“*Cooki” was 50 mm. in height x 70 mm. in expanse. Judging

from these figures and from the size of the branches of the other

specimens, it seems probable that the normal size of a full-grown

specimen of the species is not more than 100 mm. x 100 mm., or

that, in words, it is a coral that does not normally attain a very

large size.

Of the other characters, perhaps the most important one to

consider is the dimorphism of the dactylopores, because this cha-

racter has been used as a diagnostic character for the separation

of the genera Labiopora and Hrrina. In the facies “‘ Benhami”

and ‘ Dendyi” there are clearly many small dactylopores lymg

on the general surface of the coenosteum between the grooved

spines and distinct from the larger dactylopores. In the facies

‘* Ramosa” no such dactylopores could be found in the specimen

I examined (although Professor Benham says they are rare), and

in the facies ‘‘Cooki” they are certainly very rare. With the

many points of resemblance in form, colour, size of pores, ete.

between the two facies “ Ramosa ” and ‘ Benhami,” it would be

HYDROCORALLINE GENUS ERRINA. 885

very rash to propose that they should be separated into distinct

species on account of this one character,

It seems to me, therefore, that the presence of small dactylo-

pores in addition to the dactylopores of the ordinary type, in

other words, the character of dimorphism in the dactylopores, is

not a character that should be regarded as absolutely diagnostic

either of the genus or of any one of its species.

As regards the gasteropores there seem to be some variations.

In “ Benhami” the gasteropores vary considerably in diameter

from 0-13-0:17 mm., but in ‘“* Ramosa,” ‘‘ Cooki,” and ‘“* Dendyi”

they are more constant in diameter, being 0°27 mm. in the first

named, and 0°22 mm. in the latter. It is probable that these

figures are not of much value for systematic purposes. There

are many technical dificulties in the way of making accurate

measurements of the mouths of a large number of gasteropores

on any single specimen, and unless the average diameter of a

large number of gasteropores of one specimen can be compared

with similar averages from other specimens of the same facies or

species, the figures given simply represent a statement of fact

concerning a given specimen, The real value of the figures I

have given is that they prove that the diameter of the gastero-

pores 1s a variable quantity and cannot be used, except in a very

general way, as a guide to the determination of species. One

oint of rather special interest is that in the facies ‘‘ Ramosa ”’

with slender terminal branches the gasteropores appear to be

actually larger than they are in ‘“ Benhami” with thicker terminal

branches. This seems to indicate that there is no relation

between the thickness of the branches and the size of the gastero-

pores, since the expectation would be that the stouter terminal

branches would bear the larger gasteropores.

The presence of a raised margin on one side of the gasteropores

in some of the specimens of this species is a feature of some

general interest. In some of the gasteropores of the facies

‘‘Ramosa” these processes are of considerable size, and bending

over the pore have an appearance very similar to the lid of a

Cryptohelia. In ‘‘Dendyi” they are rudimentary, but in the

other specimens they are absent.

The presence of a definite lip or scale on the edge of the gastero-

pore, it must be remembered, was the principal character relied

upon by Pourtales for the separation of the genus Lepidopora

from the genus Hrrina, but if we accept Moseley’s view that the

species of Lepidopora should be incorporated with Hrrina, then

we have a parallel series of variations as regards this character in

the Hrrina group, to that in the Labiopora group.

At one time I thought that the difference in the length of the

grooved spines (nariform processes) might be a useful character

for the separation of the species in this genus. In all the speci-

mens, however, I found that the projections on the young actively

growing terminal branches are longer than they are on the older

branches, and consequently there is a difficulty in fixing a standard

Proc. Zoou. Soc.—1912, No. LIX, 59

886 PROF. 8. J. HICKSON ON THE

of measurement for comparison. Taking the measurement of a

few of the projections at a distance of about 3 mm. from the apex

of the branches, I have found that the average is in ‘‘ Benhami”

0:7 mm., and in “ Dendyi” 0:4 mm., the projections in “ Ramosa ”

and ‘Cooki” being intermediate in size between these two

measurements. The differences between these averages are so

small, and the difficulty of avoiding a relatively large error in the

calculation is so great, that the measurements are of no more

scientific value than to express roughly the general impression of

observation that the projections are longest in ‘‘ Benhami,”

shortest in ‘‘ Dendyi,” and of medium length in other specimens.

The longest projections in “ Benhami” are, however, not simple

nariform processes as they are usually in the other facies, but

groups of two, three or four of these processes clustered together

(fig. 13). The clustering together of the grooved projections

may be seen in some of the other facies, such as ‘“‘ Ramosa” and

“‘ Cooki,’ but it 1s never such a pronounced feature as it is in

“ Benhami.”

Lastly, a word about geographical distribution. All the

specimens were dredged off the coast of New Zealand, and two of

these four were found in the same bay (Preservation Inlet).

There is every reason to believe, moreover, that they were all

found in shallow water. In my opinion these facts have some

weight in determining the question whether the specimens

should be placed in one species or in several species. In a rare

genus such as Hrrina, species found at widely separated localities

will in all probability be affected by their isolation and show

differences that entitle them to rank as distinet species, but

there is much less probability that the genus would be able to

develop or to maintain specific differences in the same waters.

Unless, therefore, a very clear case is made out that the dif-

ferences between the specimens of the genus Hrrina from New

Zealand waters are constant or of fundamental importance, the

most convenient as well as most scientific course to pursue is to

place them together in the same species.

Errina (LABIorora) CAPENSIS, Sprdmeigh (Pl) XO Ve tigle7

Pl. XCVI. fig. 15.)

This species is represented in my colléction by three broken

terminal branches. The largest piece is 36 mm. in length, the

diameter at the base is 7 mm., and the diameter of the branch

3 mm. from the apex is 455 mm. In the largest specimen of

the New Zealand species (‘‘Dendyi”) the diameter at the base

(evidently the base of attachment) is 6 mm., and the diameter of

a branch 3 mm. from the apex is only 2 mm.

- From these facts it seems probable that in this species the

hydrophytum reaches to much greater dimensions than does that

of the New Zealand species.

The branches terminate in blunt, slightly flattened and ex-

panded extremities.

HYDROCORALLINE GENUS ERRINA. 887

The hydropbytum is probably flabellate in growth, the ramifi-

cation not very profuse and anastomoses rare.

Colour : salmon-pink,

Surface coarsely granular and substance coarsely reticulate.

As this species seems to approach the #rrina group in some

respects, attention may be called to the striking difference there

is between the coarsely granular character of its surface and the

fine smooth porcellanous character of the surface of the Hrrina

group.

Grooved spines very short, numerous, quite irregular in arrange-

ment and never in clusters. The form of the spine is that of a

shallow semicircular ridge open on the side turned away from the

apex.

Gasteropores equally distributed on both sides of the branches,

never provided with a lip. Diameter 0:3 mm.

Large dactylopores about 0°25 mm. in diameter.

Small dactylopores:—It may be open to discussion whether

there is or is not any true dimorphism of the dactylopores in this

species. It has been shown that in one of the specimens from

New Zealand (‘“‘ Dendyi”) the small dactylopores are provided

with shallow collars. In the Cape specimen some of the dactylo-

pores are considerably less in diameter than the majority, but

they are provided with exactly the same kind of semicircular

ridge as the larger ones. The question of dimorphism, therefore,

resolves itself in this case into a question whether the essential

feature of the dimorphism of the dactylopores consists in their

size or in the presence of a grooved spine. Ridley states that in

irrina (Labiopora) moseleyi the dactylopores are of the usual

uniform size, and the two kinds can only be distinguished by the

presence or absence of nariform processes. These facts seem to

emphasize the conclusion that the so-called dimorphism of the

dactylopores is not really a feature of very great importance, and

to suggest as a probability that the small dactylopores are the

shelters for young dactylozooids which in their later stages of

growth increase in size and become protected by a grooved spine.

The specimen described and figured by Gray (1872) as Hrrina

jissurata, from the Antarctic seas, was apparently very closely

related to this species. Unfortunately the specimen has been

mislaid (jide Moseley) and cannot therefore be re-examined, but

the figures show a similar robust habit of growth and short semi-

circular grooved spines with the grooves all turned away from the

apex of the branch.

‘Errina (LABIOPORA) ANTARCTICA Gray.

Porella antarctica Gray.

Labiopora antarctica Moseley.

Labiopora antarctica Ridley.

The type- specimen was found off the Falkland lieu

54° 27'8., 59° 40’ W., in 45 fathoms.

A second specimen, attributed to this species by Ridley, was

found in Trinidad Channel, 5.W. Chili, in 30 fathoms.

5o*

888 PROF. S. J. HICKSON ON THE

Hydrophytum flabellate in growth. There is no statement to

the effect that the branches anastomose. No record of size

beyond the statement that it is smaller than L. moseleyi.

Colour: bright crimson, with the compressed forked tips paler.

Surface minutely reticulate. Grooved spines arranged in rows,

but not in clusters, on both sides of the branches, all turned away

from the apex of the branch.

Gasteropores without a lip, 0°22 mm. in diameter (in the Chili

specimen).

Large dactylopores 0-09 x 0:2 mm.

Small dactylopores 0°08 mm., without any lip or collar.

Errina (LApropora) MosELEYI Ridley.

The single specimen of this species was found at Port Rosario,

S.W. Chili, 2-10 fathoms.

Hydrophytum flabellate in growth. Anastomoses frequent.

95 mm. in height x 185 mm. in width.

Colour: “ vermilion.”

‘¢ An anterior clearly distinguishable from a posterior surface,

by the development on it of numerous tubercles, chiefly in the

terminal branches, which are very slightly indicated in the latter.”

Surface minutely reticulate. Grooved spines not arranged in

definite rows nor in clusters.

Gasteropores without any lip, 0°32 to 0°35 mm. in diameter.

Large dactylopores 071 to 0-14 mm. in longitudinal diameter.

Small dactylopores without lips or tubercles, of about the same

size as the large dactylopores.

Errina (LApropora) ASPERA Linn. (Pl. XCV. fig. 6.)

IT have examined the type-specimen of this species in the

British Museum on which Gray (1) founded the genus Hrrina.

There can be no doubt that the affinities of the species with the

genotype of Labiopora are closer than they are with Moseley’s

Errina labiata. It belongs to the ZLabiopora, and not to the

Errina group of species.

One of the colonies in the British Museum is 80 mm. in height

by 85 mm. in width. It forms a flabellum with a clear difference

between the anterior and posterior surfaces. The branches

terminate in fine points and do not anastomose. The branches

are about 3 mm. in diameter at a distance of 3 mm. from the

apices. The surface is minutely granular. The grooved spines

sometimes occur in clusters, but when solitary the grooves are

turned away from the apex of the branch. There are a few small

dactylopores without spines and some with small or rudimentary

spines. This specimen was dredged off the coast of Sicily.

A little while ago a specimen of a Stylasterine coral was sent

to me by the late Mr. John Morgan of Worthing. It was

purchased in a sale and there was no record of its locality.

At first I thought it should be placed with the other specimens

HYDROCORALLINE GENUS ERRINA. 889

in my collection in the species Lrrina (L.) nove zelandie, but,

on comparing it with the specimens in the British Museum, I

came to the conclusion that it is probably related more closely to

Errina (L.) aspera. Itis moderately branched and roughly flabelli-

form in growth. The terminal branches are fairly thick, being

about 2-2°25 mm. at a distance of 3 mm. from the apex. ‘The

branches do not anastomose.

The surface and substance of the ccnosteum are minutely

granular.

The grooved spines are very crowded and clustered on the

terminal branches, and not arranged in rows (fig. 6). The

grooves are turned in all directions, but the majority of them

away from the apex of the branch.

The gasteropores are equally distributed on both sides of the

flabellum, and are without any lip or collar. Size 0°13-0°17 mm.

in diameter.

The large dactylopores are very variable in size, ‘09 x ‘09 mm.

to ‘06 x 0-11 mm.

The small dactylopores are very numerous, 0°06 mm. in

diameter, and provided with small curved lips.

The specimen is like the type in being white in colour.

Mr. Morgan’s specimen differs from the type in having rather

more slender branches, in having the pores equally distributed

on the two surfaces of the flabellum, and in the presence of

numerous small dactylopores.

Errina (LABIopoRA) GRACILIS von Marenzeller.

Several specimens of this species were found by the ‘ Belgica’

Expedition attached to the swabs when dredging off the pack ice

in the region of 71°S. and 88°W., i.e. about 20 degrees west

and 15 degrees south of the Straits of Magellan. The depth

is not recorded by von Marenzeller, but it is probably between

500 and 600 metres.

The hydrophytum is flabellate in growth, with well-marked

anterior and posterior surfaces. One of the specimens, which

proved to bea female, was 25 mm. in height and 30 mm. in width.

Another, which proved to be a male, was 100 mm. by 140 mm.

The surface of the ccenosteum is finely wrinkled, and marked

with transverse and longitudinal ridges (Kammschen). There are

apparently no well-marked ccenosteal pores.

Grooved spines not arranged in definite rows, but in irregular

clusters or singly. The grooves of the grooved spines turned

away from the apex of the branches or sideways.

Gasteropores with a lip, ‘015 mm. in width.

Large dactylopores sheltered by spinous projections provided

with a deep groove.

Small dactylopores with or without a lip.

Colour: white or brownish.

There can be little doubt that this species belongs to the

890 PROF. S. J. HICKSON ON THE

Labiopora group. It is true that the description given by von

Marenzeller of the surface of the ccenosteum does not agree with

that of the cther species of the group, but it does not agree either

with the description given of the surface of the ceenosteum of the

Errina group. The absence of well- defined ccenosteal pores, the

grouping of some of the grooved spines in clusters, and the direc-

tion of their grooves—all point to the affinities of the species with

the Labiopora group.

It may be remarked that this is the only species of the group

that occurs in deep water.

Review of the Labiopora Group of Species.

The careful examination of the specimens belonging to the

Labiopora group leads me to the conclusion that there are very

few characters that can be used with much confidence for the

separation of species *. However, it may be convenient for the

present to recognise six species :—

Errina (Labiopora) aspera Linn. Mediterranean Sea.

Lirring (Labiopora) antarctica Gray. Chili and Falkland Islands.

30 to 45 fathoms.

Lrrina (Labiopora) moseleyi Ridley. Chili. Shallow water.

Hrrina (Labiopora) nove zelandie Hickson. New Zealand.

Shallow water.

Errina (Labiopora) capensis Hickson. Cape of Good Hope.

30 fathoms.

Lrrina (Labiopora) gracilis von Marenzeller. Antarctic Sea.

Deep water.

Of these six species Hrrina (Labiopora) capensis appears to be

the most sharply defined. It probably attains to a much larger

size, has more robust branches terminating in blunt and some-

what flattened extremities. Its substance is coarsely reticular

and its surface coarsely granuiar. The grooved spines are short

and semicircular in shape. All the dactylopores are guarded by

these spines.

The other five species are very closely related. Hrrina (L.) ant-

arctica appears to be distinguished from the others by the grooved

spines being arranged in definite rows, and Lrrina (L.) moseleyi

by the differentiation of an anterior from a posterior surface

of the flabellum. Of Hrrina (L.) nove zelandie all that can

be said is that it appears to be a very variable species which

does not exhibit any one particular distinguishing feature.

Hrrina (£.) gracilis is distinguished from the others by the

texture of the surface of the ccenosteum, and the colour is not

red but white or brownish. rrina(L.) aspera has a close resem-

blance to some of the facies of Hrrina (L.) nove zealandie, but

it is always white in colour, in this respect resembling the facies

‘** Cooki.” '

* See Note, p. 894.

2 HYDROCORALLINE GENUS ERRINA. 891

General Remarks on Structure.

Gasteropores.—Before concluding this account of these species

I may refer briefly to one or two results of a negative character

that [I have obtained. It occurred to me that if the speci-

mens I examined belonged to different species, there might be

some measurable difference to be observed in the styles of the

gasteropores. The style is in all cases like an elongated and

sharply pointed brush, but careful and constantly repeated com-

parisons of the styles of the gasteropores of different specimens

revealed no characters by which they could be distinguished.

The type of style is the same in all the specimens, and although

the brush is more slender in some cases than in others, there is a

greater range of variation in this respect in the gasteropores of

a single specimen than there is in the gasteropores of different

specimens taken at random (Pl. XCVI. figs. 14 & 15).

The way in which it is possible to study the shape of the

gasteropore styles is to make a vertical fracture in a plane parallel

with the long axis of a branch. Ina large percentage of such

fractures the whole length of at least one gasteropore with its

style will be exposed.

In such fractures it may be observed that some of the gastero-

pores extend as far down as the axis of the branch, but in others

they extend only a short distance below the surface. In the long

gasteropores the style can be traced right down to the base of the

pore, but it is usually supported by one or, in some rare cases,

more than one, tabula (fig. 14) similar to the tabule described by

Miss England (5) in the gasteropores of Spinipora. In the short

gasteropores the style ends abruptly at the base of the pore and

is not supported by a tabula (fig. 15).

In order to make any scientific use of this character in the

classification of the species it would be necessary to make a large

number of fractures, and for that purpose I have not sufficient

material.

The results I have obtained on the few fractures I have made

are as follows :—

In Errina (L.) nove zelandie facies “‘ Ramosa” and facies

“‘Cooki” the gasteropores (I have observed) are long and exhibit

atabula. In the same species facies ‘‘ Benhami” the gasteropores

are short, and the same is the case in the facies “ Dendyi.” In

Mr. Morgan’s specimen of Hrrina (L.) aspera the gasteropore 1s

also short. In Hrrina (L.) capensis the gasteropores are short,

extending only a little way beneath the surface. As to this

character in the gasteropores of the other species I have no

evidence to bring forward.

My conclusions, from this evidence, are that the length of the

gasteropores and the presence or absence of a tabula are not, at

present, characters of any value for purposes of classification.

(Yonophores.—All the specimens I have examined show ampulle.

They can usually be seen on the terminal branches as shallow

892 PROF. 8. J. HICKSON ON THE

conyexities on the surface, sometimes between the spines, some-

times carrying the spines with them. They appear to be always

absent in the older branches and stems. Our knowledge is still

so very imperfect concerning the growth and relations of the

gonophores in the Stylasterina that it is not reasonable to use any

differences that may be observed in the characters of the ampulle

for purposes of classification.

The questions to be answered are the following :—

1. Are there any constant differences between the male and the

female ampullee ?

2. Is there any correlation between the sex of the colonies and

characters of the general structure of the ccenosteum ?

These two questions could be answered without much difficulty

by the examination of a number of preserved specimens of the

‘same species from the same locality.

In Professor Benham’s notes on the structure of a specimen

which probably corresponds with Hrrina (Labiopora) nove

zelandice facies “‘ Ramosa,” I find the following remark :—‘ Each

ampulla seems to have a small pore (?dactylopore) on its surface.”

In the specimen I have examined of this species I cannot find

these small pores on the ampulle. Von Marenzeller also remarks

that the male ampulle exhibit small dactylopores “in niedrigen

Spitzchen.”

In a previous paper (3) I have shown that, in the ripe male

gonophores of Distichopora and Allopora, the spermatozoa are

discharged by a spout-like seminal duct. In the case of the female

gonophore, however, the only way of escape of the embryo is by the

rupture of the whole surface of the ampulla. It seems possible,

therefore, that the pore on the ampulla mentioned by Professor

Benham may be for the opening of the seminal duct and that the _

specimen may be a male. On the other hand, it must not be

assumed that when the surface of a specimen is marked by large

shallow depressions having the size and appearance of ruptured

ampulle, the specimen is necessarily a female colony, because in

the specimen of Zrrina (Labiopora) capensis in which these

shallow depressions are very well-marked I found to my surprise

that it was a male colony.

As regards the structure of the gonophores there is very little

to be said at present. The male gonophores of Hrrina (Labiopora)

capensis exhibit a well-developed spadix (manubrium), and in this

respect the genus seems to resemble Distichopora and to differ

from Allopora. The only other spirit specimen I have received

was a small branch of Errina (Labiopora) nove zelandice facies

‘* Benhami,” and this proved to be a female. The specimen was

not, however, in sufficiently good condition to enable me to study

the structure of the gonophore.

Cenenchymal Canals—From the two spirit specimens I have

ascertained certain isolated facts which may be of some use when

our knowledge of the genus is extended.

HYDROCORALLINE GENUS ERRINA. 893

According to Ridley the cenenchymal canals of Z. moseleyi and

L. antarctica are 0°035 to 0:07 mm. in diameter, the meshes

between the canals are about the same diameter in LZ. moseleyi,

but in ZL. antarctica they vary from the same diameter up to

0:14 mm. Ridley does not state how his measurements were

obtained, consequently the results I have obtained cannot be

compared with his results with any degree of precision.

I have measured the diameter of a number of canals that have

been cut transversely in my sections at a level corresponding

with the base of the dactylozooids, and I find that the average

diameter of these canals is in Lrrina (Labiopora) nove zelandie

0-03 mm. and in Hrrina (L.) capensis 0°05 mm. As the specimens

must have undergone very considerable contraction during de-

calcification and imbedding, I have regarded the measurements of

the meshes as worthless. All that can be said is that in both

species the meshwork of canals is close and elaborate.

In the ‘ Challenger’ monograph Moseley gives as a character

of the genus Hrrina that the number of tentacles on the gastero-

zooids is four. In the Siboga species of Hrrina the number of

tentacles varied from four to five.

In Errina (Labiopora) nove zelandie I have found some diffi-

culty in counting the number of tentacles, but judging from a

single series of sections, there are not more than four. In the

specimen of Hrrina (Labiopora) capensis, however, a very large

number of gasterozooids can be clearly seen without decalcification,

and in these the number varies from four to six. From this fact

it is obvious that the number of tentacles on the gasterozooids

cannot be relied upon as a generic or even as a specific character.

List of the Species of the Genus Errina.

A. Coenosteum hard and compact, perforated by well-defined

ccenosteal pores. Grooved spines turned towards the apex

of the branch.

1. With short grooved spines and only one kind of dactylopore.

The Lrrina group.

Errina labiata Moseley.

Errina ramosa Hickson & England.

Errina horrida Hickson & England.

Errina carinata Pourtales 2

Errina pourtalesii Dall ?

*Errina glabra (= Lepidopora glabra Pourtales).

*Hrrina cochleata (= Lepidopora cochleata Pourtales).

*Krrina dabneyt (= Lepidopora dabneyi Pourtales).

2. With long grooved spines and two kinds of dactylopores.

Errina echinata (= Spinipora echinata Moseley).

* See Note, p. 894.

894 PROF. 8. J. HICKSON ON THE

B. Cenosteum granular and reticulate, without well-defined

coeenosteal pores. Grooved spines turned away from the

apex of the branches or clustered and irregularly placed.

The Labiopora group.

Errina aspera Linn.

Lrrina fissurata Gray.

Errina antarctica (= Labiopora antarctica Gray).

Errina moseleyt (= Labiopora moseleyi Ridley).

Lrrina nove zelandic Hickson.

Hrrina capensis Hickson.

Lrrina gracilis von Marenzeller.

The following species has the grooves turned away from the

apex as in the Labiopora group, but has well defined coenosteal

pores :—

Errina macrogastra yon Marenzeller (7).

(Additional notes on Professor Hickson’s paper on the

genus Hrrina, received August 14th, 1912.

1. The adjective ‘‘ Hydrocoralline” is used in the title of the

paper at the suggestion of the Secretary of the Society for the

convenience of Zoologists who do not possess a special knowledge

of these Coelenterata. I have in several previous papers declared

my opinion that to unite Millepora with the Stylasterina in one

Order, the Hydrocorallina, is unsound. I have found no reason

to change my mind in that respect. The word “ hydrocoralline ”

may still be used, however, to signify the corals that are Hydrozoa

in very much the same way as the words “corals,” “ zoophytes,”

“worms,” are used, without any strict systematic significance.

2. Since the paper was read I have been able to examine a

number of specimens in the possession of the Muséum d’ Histoire

Naturelle, Paris, for which I am indebted to the kindness of

Prof. Joubin.

In this collection there are some specimens which I have

identified as Hrrina dabneyi Pourt. from the Azores. An

examination of these specimens convinced me that they belong to

the Labiopora group of species, as they exhibit all the charac-

teristic features of that group. From this it seems probable, that

the other two species, L. glabra and £. cochleata, that were in-

cluded by Pourtales in the genus ZLepidopora should also be

transferred from the Lrrina to the Labiopora group. |

Literature.

(1) J. E. Gray.—(No title.) Proce. Zool. Soc. 1835, p. 85.

(2) J. E. Gray.—Notes on Corals from the South and Antarctic

Seas. Proc. Zool. Soc. 1872, p. 744.

(3) 8. J. Hickson.—The Gonophores of Allopora and Disticho-

_porad. Quart. Journ. Microscop. Sci. vol. xxxu. 1891,

p. 382.

HYDROCURALLINE GENUS ERRINA, 895

(4) 8. J. Hickson & Heten M. Eneuann.—The Stylasterina of

the Indian Ocean. Trans. Linn. Soc. xii., 1909.

(5) S. J. Hickson & Heten M. Eneranp.—The Stylasterina of

the Siboga Expedition. 1905.

(6) E. von Marenzecter.—Madreporaria and Hydrocorallia.

Résultats du Voyage du 8.Y. ‘ Belgica.’ 1903.

(7) E. von Marenzecier.—Stein- und Hydro-Korallen. Bull.

Mus. Comp. Zool. xliii. 2. 1904.

(8) H. N. Mosetny.—Zoological Collections of H.M.S. ‘ Chal-

lenger,’ vol. ii. 1881, p. 50 seg.

(9) L. F. pe Pourrates.—Deep Sea Corals. Illustrated Cata-

logue Museum Comp. Zoology, Harvard, 1871.

(10) S. O. Rrptey.—Zoological Collections of H.M.S. ‘ Alert.’

Proc. Zool. Soc. 1881, p. 105.

EXPLANATION OF THE PLATES.

Prate XCIV.

Fig. 1. Errina (Labiopora) nove zelandie (facies Benhami) ; a portion of a branch

showing the grooved spines arranged in clusters or with the grooves

turned away from the apex. X 4 diam.

Fig. 2. Another photograph of the same facies (Benhami) showing a number of

closely packed anastomosing branches. In this view it will be seen that

although the greater number of the grooves are turned away from the

apex of the branches (the top of the photograph), in places where they are

clustered they are turned in all directions. > 4°75 diam.

Fig. 3. The same species (facies Ramosa). he grooved spines are not so crowded

and rarely form clusters. Nearly all the grooves are turned away from

the apex. X 9°3 diam.

Fig. 4. The same species (facies Dendyi). The grooved spines are still more scattered

and nearly all the grooves are turned away from the apex. in this

photograph several gasteropores can be seen. > 9°3 diam.

PLATE XCV.

Fig. 5. Errina (Labiopora) nove zelandie (facies Cooki.) In this facies many of

the spines are in clusters, consequently the grooves appear to be turned

in all directions. X 45 diam.

Fig. 6. Hrrina (Labiopora) aspera. In this specimen the spines are rather

water-worn, but frequently occur in clusters and the grooves point in all

directions. > 43% diam.

Fig. 7. Errina (Labiopora) capensis. The spines are very short and the grooves

all point away from the apex. In this photograph the style can be seen

in many of the gasteropores. XX 45 diam.

Fig. 8. Errina (Spinipora) echinata. In this species, as in the Hrrina group of

species, the grooves are turned towards the apex. x 44 diam.

Prate XCVI.

Fig. 9. Errina (Labiopora) nove zelandie (facies Ramosa). Slightly enlarged,

showing the profuse ramification without anastomoses, and branches

terminating in delicate points. Drawn by Prot. Benham.

Fig.10. Hrrina (Labiopora) nove zelandie (facies Cooki). Slightly enlarged,

showing the method of ramification and branches terminating in blunt

points. Drawn by Prof. Benham.

Fig. 11. A terminal branch of the same species (facies Cooki), showing the dactylo-

pores (D) protected by the grooved spines and the gasteropores (G).

X 6 diam. Drawn by Prof. Benham.

Fig. 12. A portion of a larger branch of the same species (facies Cooki), showing the

grooved spines in clusters. XX ca. 6diam. Drawn by Prof. Benham.

896 MR. E, DUKINFIELD JONES ON

Fig. 13. Evrina (Labiopora) nove zelandie (facies Benhami). One of the clusters

of four grooved spines as seen from above. X ca. 12 diam.

Figs. 14 & 15. These two drawings are intended to show the difference in texture

between Hrrina (L.) nove zelandie (fig. 14) and Hrrina (L.) capensis

(fig. 15), as seen in vertical section of the ccenosteum. Another point

of difference between the two specimens is that in the former (fig. 14)

the gasteropore penetrates into the depths of the branch and shows a

tabula (¢) in its course; whereas in the latter (fig. 15) the gastero-

pore ends abruptly a little way below the surface. This difference,

however, cannot be regarded asa specific distinction. G, gasteropore ;

D, dactylopore; St., style. Both figures X 35 diam.

54, Descriptions of new Butterflies of the Genus Thecla

from 8.E. Brazil. By E. Duxtyrietp Jonus, F.Z.8.,

FES.

[Received May 13, 1912: Read October 29, 1912.]

(Plate XCVII*)

InvEx.

Systematic : Pages

Thecla, 13 new species Of ...ec..c.eeceeecseeeeeeesee 896-902

The species described in the present paper were captured by

myself and the types are in my own collection.

THECLA HAMILA, sp.n. (Pl. XCVII. fig. 1.)

Allied to 7. gispa Hew.

Male. Upper side rich purplish blue; costa and termen of fore

wings narrowly black, wider at apex; base of costa greenish ;

a large black silky spot with blue reflection beyond cell; cilia

black. Hind wings: costa and termen narrowly black; inner

margin greyish black, rather broad at tornus; tails black with

white tips; a few white scales on inner side of lobe; some

greyish green at tornus above the black margin; submedian hairs

greenish blue; head black; thorax greenish blue; abdomen

purplish blue. Under side dull stone-green suffused with black,

a pale green reflection over all. Fore wings: a medial row of

diffused light stone-green spots from costa to vein 2, the spot

above vein 3 distal; an indistinct subterminal diffused pale green

band; termen black; cilia black with pale green band at apex.

Hind wings darker; a broad diffused black medial shade, very

dark below costa; a narrow dark postmedial shade and broad

subterminal shade; a medial series of green spots between the

veins from inner margin to vein 5, those above veins 2 and 3

distal ; termen and cilia black.

Expanse 34 mm.

Hab. Castro, Parana, Brazil.

* For explanation of the Plate see p. 902.

PAS, Jol2, Pee Vil,

Horace Knight delet lith. West, Newman chr.

NEW SPECIES OF THECIA FROM 5S. E. BRAZIL.

“NEW BRAZILIAN BUTTERFLIES. 897

THECLA FANCIA, sp.n. (Pl. XCVII. fig. 2.)

Allied to 7’. punctwm H.-S.

Male. Upper side bright metallic blue with a greenish reflection

on basal half of wings; costa of fore wings narrowly, apex and

termen broadly black; a brown circular brand on upper angle of

cell; hind wings outwardly broadly black; tails tipped with

white. Under side yellowish grey: fore wings, a postmedial

band of indistinct brown lunular spots outwardly bordered with

white from costa to vein 2; hind wings: postmedial band inwardly

dark brown, outwardly white, angled on veins | and 2, inwardly

oblique from 3 to 4, placed more distally from 4 to 6 and above 7 ;

a small orange spot at tornus anda larger one between veins 2

and 3.

Expanse 30 mm.

Hab. Castro, Parana, Brazil.

THECLA SICRANA, sp. n. (Pl. XCVILI. fig. 3.)

Male. Upper side dull metallic blue; costa and termen of fore

wings narrowly black ; submedian hairs on hind wings greyish ;

tornus dull orange; tails and cilia black tipped with white.

Under side bright green; inner margin of fore wings bluish

grey ; hind wings: an indistinct postmedial series of lunular spots

inwardly black, outwardly white; dark red-brown subterminal

spots above veins 1, 2, and 3, the one above 2 larger; tornus dark

red-brown; cilia black tipped with white. Palpi white with

some lateral tawny scales, 3rd joint black; frons brown, with a

few metallic green scales; a white point between antenne ;

abdomen bluish grey above, white beneath.

Expanse 30 mm.

Hab. Castro, Parana, Brazil.

THECLA BERTHA, sp.n. (Pl. XCVILI. fig. 4.)

Female. Upper side smoky brown, inwardly suffused with

bluish grey ; fore wings irrorated from median nervure to inner

margin with light blue; cilia smoky brown interrupted by white

between the veins; in the hind wings the irroration covers the

cell also; tornus tawny, with a dark spot on lobe; tails black

with white tip; cilia outwardly whitish. Under side bright

green ; basal half of submedian area of fore wings brownish grey,

outer half light grey; a rather broad clear white postmedial

band with indistinct dark central line from costa to vein 2; cilia

reddish brown, tipped with white between the veins. Hind

wings: a nearly straight white medial line defined by maroon on

both sides from vein 8 to vein 1, angled inwards at origin of

veins 3 and 4, broken at vein 2, being placed more distally below

the vein, angled inwardly on submedian fold ; a large white spot

slightly irrorated with maroon above upper angle of cell; three

spots in cell; one spot above vein 2 and one above vein 1;

a terminal series of lunular maroon spots irrorated with white,

fusing at the veins, the one above vein 2 being produced inwardly,

898 MR. E. DUKINFIELD JONES ON

forming a large maroon spot without irroration; cilia reddish

brown, tipped with white between the veins.

Expanse 30 mm.

Hab. Castro, Parana, Brazil.

THECLA SCHAUSA, Sp.n. (PI. XCVII. fig. 5.)

Male. Upper side: fore wings black; cell and submedian area

to near tornus metallic purplish blue ; ; extreme edge of costa

tawny; a small dark grey brand beyond cell; cilia grey ; hind

wings metallic purplish blue; costa and apex black; termen

narrowly black; tornus tawny red; cilia grey. Under side:

fore wings brown ; a purplish reflection on medial area below

costa; a “postmedial wavy dark red line from costa to vein 2;

a terminal band of dark red irrorations wide at apex and taperi ing

down to vein 2; termen tawny red: bind wings purplish brown ;

an inner and a medial line of red irrorations ; ‘outer half of wing

somewhat greyish, very thickly irrorated with red; a diffused red

subterminal band; termen tawny red. Frons tawny red ; palpi

white mixed with br own; pectus white.

Female. Very similar to the male; the blue of the wings is

duller; cilia almost white ; termen of hind wings more broadly

black.

Expanse, male 17 mm., female 17 mm.

Hab. Castro, Parana; Sao Paulo, S.E. Brazil.

THECLA JAPOLA, sp.n. (Pl. XOVIL. fig. 6.)

Allied to 7. gaina Hew.

Female. Upper side dull steel-blue ; costa, apex, and terminal

area of fore wings broadly greyish black ; a dark diffused spot

beyond cell; basal half of extreme costa tawny ; cilia outwardly

brown. Hind wings: costal and terminal areas greyish black,

lighter than fore wings; an indistinct subterminal row of tere

lunular spots, conspicuous above veins 1 and 2; a red spot on

lobe ; tails black, tipped with white; termen black, preceded by

whitish line; cilia black at base, white at centre, brown at tips.

Under side brownish grey; fore wings: an inwardly diffused

reddish-brown postmedial band, outwardly white, straight from

costa to vein 2; terminal area pale grey with two diffused brown

bands, the inner broad, the outer narrow; termen dark brown;

cilia light brown ; hind wings: an inwardly reddish-brown, out-

wardly white medial band straight from costa to vein 2, angled

on veins | and 2 forming a very acute W; terminal area whitish

grey with an inner brown band, broad and diffused at costa,

narrow and distinct at tornus, bent strongly towards base on

inner margin, followed at tornus by orange-red to a black spot on

lobe, some white at base of lobe; an orange-red lunular spot with

black point above vein 2; termen dark brown; cilia lighter with

pale band.

Expanse 28 mm.

Hab. Castro, Parana, Brazil.

NEW BRAZILIAN BUTTERFLIES. 899

THECLA FERNANDA, sp.n. (PI. XCVII. fig. 7.)

Allied to 7. nubilum H. H. Druce.

Male. Upper side dark blackish brown; a red spot on lobe and

a few white scales at tornus; tails black tipped with white.

Under side yellowish grey-brown ; fore wings: a dark postmedial

band outwardly bordered with white, nearly straight from costa

to vein 2; hind wings: postmedial band inwardly shaded with

red and outwardly with white, angled on veins 1] and 2, proximate

between veins 3 and 4, straight from 4 to 7, distal above 7

forming conspicuous spot at costa. Differs from 7’. nubilwm in

the band on fore wing being straighter, that on the hind wing

less acutely angled on veins 1 and 2 and in the distal spot at costa,

also in absence of discocellular bar.

Expanse 29 mm.

Hab. Fernandes Pinheiro, Parana, Brazil.

THECLA NORA, sp. n. (Pl. XOVII. fig. 8.)

Male. Upper side dark metallic lilacine blue; costa and termen

of fore wings narrowly black, broader at apex; a large black

brand beyond celi; cilia black at base, whitish metallic blue at

tips; hind wings: costal area greyish brown; a fine black

terminal line ; cilia black with bluish-white band. Under side

brownish grey: fore wings suffused with steel-blue on submedian

area; postmedial band inwardly dark brown, outwardly white

from costa to vein 3, below vein 3 the brown changes to blue ;

a very indistinct subterminal line, inwardly white, outwardly

brown; cilia brown; hind wings: postmedial band dark brown,

shaded inwardly with orange and outwardly with white, strongly

angled on veins 1 and 2, outwardly displaced between 4 and 5;

a series of subterminal lunular spots shaded inwardly with white,

followed by orange spot at tornus, a dark spot irrorated with

white below vein 2 and an orange spot containing black point

below 3; a narrow white terminal line; cilia dark brown with

bluish-white tips.

Expanse 24 mm.

Hab. Castro, Parana, Brazil.

THECLA MOLENA, sp.n. (Pl. XCVII. fig. 9.)

Closely allied to 7. lemona Hew.

Male. Upper side dull dark metallic blue; fore wings: outer

half of costa, apex and termen black; a large oval black brand,

crossed by blue discocellular bar ; extreme costa tawny at base ;

hind wings: termen narrowly black; tails black tipped with

white; a red spot on lobe. Under side brown with a lilacine

reflection on terminal area of fore wings and the whole of the

hind wings. The bands and lines are much the same as in

T. lemona, but they are much more diffused and subdued; this

difference and the liacine reflection readily distinguish the

species. . 7’. lemona is also a larger insect than the present.

900 MR. BE. DUKINFIELD JONES ON

Female. Upper side brown, suffused with lilacine on basal area.

Under side as in male, but the lilacine reflection is much less

pronounced,

Expanse, male 24 mm., female 24 mm.

Hab. Castro, Parana, Brazil.

THECLA ESMERALDA, sp.n. (PI. XCVII. figs. 10, 11.)

Close to 7. janias Cr., but differs in the very much wider

black margins of the upper side and in the postmedial line, and

in the orange spot in place of the black spot on the under side in

janias.

Male. Palpi and head vivid metallic green; legs green; tarsi

inwardly whitish and banded with black; thorax, abdomen, and

wings rich purplish blue. Fore wings broadly bordered with

black on costa and termen, very broad at apex; a small circular

dark grey brand. Hind wings broadly black on costal and apical

areas, narrowly from vein 4 to tornus; a green spot on the black

at tornus; tails black tipped with white. Under side: fore

wings vivid metallic green, inner margin grey; cilia brown:

hind wings vivid metallic green; a fine black wavy postmedial

line, most distinct at inner margin; some lunular subterminal

spots near tornus, that above vein 2 being bright orange; some

white scales at tornus.

Female. Thorax green ; wings and abdomen brown, with slight

lilacine grey tint on hind wings and basal half of fore wings.

Under side as in male, but the postmedial line clearer and the

orange spot above vein 2 much larger, there is also some orange

at tornus; cilia inwardly black, outwardly whitish.

Expanse, male 27 mm., female 25 mm.

Hab. Castro, Parana, Brazil.

THECLA CASTRENA, sp.n. (Pl. XCVII. figs. 12, 13.)

Allied to 7’. phrosine H. H. Druce.

Male. Upper side dark blackish brown ; a few lilacine scales

at base and along inner margin of fore wings; a slight lilacine

reflection on hind wings; termen dark and cilia white in both

wings. Under side: fore wings pale lilacine blue; apex and

inner margin whitish; a lilacine reflection over the whole ;

a medial V-shaped mark on submedian fold; a slightly wavy

red-brown postmedial band from costa to vein 2, followed by

white spot above vein 6; a fine black terminal line, becoming

brown at apex ; cilia white at tornus, shading to brownish at apex.

Hind wings white with lilacine reflection ; a bright orange-tawny

irregular postmedial band of elongated spots, placed distally below

veins 1, 3, 5, 6 and 8, and proximally below 2, 4 and 7; a sub-

terminal row of minute tawny and black spots between the

veins; a fine black terminal line, shading to brown at tornus ;

cilia white.

Female. Upper side the same as the male, except that the cilia

NEW BRAZILIAN BUTTERFLIES, 901

are brown. Under side yellowish grey, the hind wings lighter

than fore wings; bands as in male, but the postmedial is out-

wardly shaded with white and there is a subterminal series of red

spots from vein 2 to 6; termen brown ; cilia brown : on the hind

wing the subterminal spots are much larger than in the male,

termen brown, inwardly shaded with white, cilia greyish

white.

Expanse, male 25 mm., female 24 mm.

Hab. Castro, Parand, Brazil.

THECLA CAMPA, sp.n. (Pl. XCVII. fig. 14.)

Near 7’. biblia Hew.

Male. Upper side dull blackish blue, shading to black on outer

half of fore wings; cilia brown; hind wings; termen narrowly

black ; fringe of abdominal fold white; cilia black, with white

band at tornus, outwardly brown at apex. Under side light

green with a light lilacine-blue reflection; cilia of fore wings

light brown ; hind wings: an obscure greyish-white discocellular

bar; an obscure sinuous greyish-white postmedial line, more

clearly defined from inner margin to vein 3; a black spot at

tornus with some white scales above and below it; a minute

black point below vein 2 and a small orange-red spot ringed with

black below vein 3; termen near tornus bluish white. Palpi

white with green reflection, third joint black above, white

beneath ; frons metallic green; back of head and tegule dark

brown ; eyes surrounded with white; body blue-black above,

creamy white beneath.

Expanse 22 mm.

Hab. Castro, Parani, Brazil.

THECLA DATITIA, sp.n. (Pl. XCVII. fig. 15.)

Very near 7’. tadita Hewitson, but differing from it in the

possession of tails, the narrow black margins on upper side of

the hind wings, and in the terminal rows of lunular spots and the

black spots on the under side of the hind wings.

Male. Upper side: fore wings smoky black; a very dark

medial suffusion from costa to vein 2; outer two-thirds of cell

filled with raised brown scales; a submedian patch of lilacine

blue from base to near tornus; termen dark; cilia greyish;

hind wings lilacine blue, broadly black on costa and apex;

a series of diffused black subterminal spots, confluent below apex ;

termen dark ; cilia greyish ; tails dark with white tips. Under

side grey with light brown bands. Fore wings: a light brown

discocellular bar; a well-defined light brown postmedial band,

diffused on inner side; a narrower and wavy subterminal band

followed by a row of diffused spots; termen light brown; cilia

grey at base, brown at tips. Hind wings: base light brown

irrorated with grey; a light brown discocellular bar; a dark

narrow postmedial band, diffused on inner side and followed by

Proc. Zoot. Soc.—1912, No. LX. 60

902 MR. R. LYDEKKER ON

narrow grey line and broad light brown suffusion ; a medial spot

ringed with grey at costa; a terminal series of diffused lunular

spots, containing a black spot above vein 2.

Expanse 23 mm.

Hab. Fernandes Pinheiro, Parana, Brazil.

EXPLANATION OF PLATE XCVII.

Fig. 1. Thecla hamila 3. Fig. 9. Thecla molena @.

2. » jancia 6. 10. » esmeralda @.

3. » sterana 6. init . mh ;

4. » bertha 2. 12. » ¢castrena 3.

5. » schausa 3. 13. + ss ‘

6. » japola &. 14. » canpa o.

‘ihe » fernanda g. 15. » adatitia 3.

8. » nora 6.

55. The Bornean Bantin.

By R. Lypexxer, F.R.S., F.Z.8.*

[Received October 1, 1912: Read October 29, 1912.]

(Text-figures 123-125.)

That the Bantin of Borneo should be subspecifically distinct

from the typical Bos sondaicus of Java is what might naturally

be expected, and the only wonder is that its distinction has

not long since been recognized and a name assigned to the

local race. This, however, is not improbably due to the fact

that, so far at least as this country is concerned, the Javan

Bantin is an exceedingly rare animal in museums, the only com-

plete adult example in the British Museum (Nat. Hist.) being

the mounted skin of a bull obtained by exchange with the Leiden

Museum in 1846, and now, as might have been expected after

long and unprotected exposure in the galleries at Bloomsbury, in

an exceedingly dilapidated condition. The Museum also possesses

a fine mounted head of a bull, presented in 1904 by Baron L. T.

H. van Heckeren-tot-Walien; and also the skeletons of a bull

and cow, with the horns, obtained from Leiden at the same time

as the mounted skin, the skull of the former being exhibited in

the Pavilion at the further end of the Lower Mammal Gallery.

Tn addition to these is the mounted skin of an immature male,

and also a young skull, with horns—from the Lidth de Jeude

collection—both of which present all the characteristics of the

Javan race.

On the other hand, the Museum has a considerable series

of skulls and horns of the Bornean Bantin presented by Mr.

H. B. Low in 1880 and 1887, as well as two presented by

Mr. W. B. Pryer in 1886, all these being from North Borneo.

Sir Edmund Loder also possesses at least two frontlets and

* By permission of the Trustees of the British Museum.

THE BORNEAN BANTIN, 903

horns of subadult Bornean Bantins in his private museum at

Leonardslee.

As regards the colour of the Bornean Bantin I know nothing

definite ; but, in the absence of any statement to the contrary,

I presume it is approximately similar to that of the typical

Javan race; that is to say, adult bulls are blackish brown or

black with a large white rump-patch and white “ stockings,” while

the cows and young bulls have the dark area rufous or chestnut.

In all the above-mentioned Javan Specimens in the Museum

the horns are of the type of those shown in text-fig. 123, which

Text-fig. 123,

Skull and horns of male Javan Bantin (Bos sondaicus) in the

British Museum.

represents the skull of a bull obtained from Leiden in 1846. At

their origin the horns are directed mainly outwards, but at rather

more than half their length they curve suddenly inwards with

a somewhat forward and finally a backward inclination. This

60*

904 MR, R. LYDEKKER ON

causes them to have a relatively large maximum span and a, cor-

respondingly small tip-to-tip interval, as is indicated in the table

of measurements given below. In consequence, I take it, of the

outward and somewhat backward direction of the basal portion

of the horns, the frontal region of the skull is somewhat convex,

while the intercornual ridge has a distinct prominence in the

middle line.

Text-fig, 124.

Skull and horns of male Bornean Bantin (B. sondaicus lowi), from the

Rejang Valley, in the British Museum.

In the Bornean Bantin, as typified by the largest of the skulls

presented by Mr. Low to the British Museum (text-fig. 124), the

horns, which are relatively stouter, are less curved and have a

THE BORNEAN BANTIN. 905

more upright direction, arising at an angle of about 45° from the

ends of the frontal ridge of the skull, and finally curving inwards

to a less extent than those of the typical race, the terminal

portion thus being shorter and the maximum span less. Similar

characters are shown in the two frontlets with horns in the

Text-fig. 125.

A & B.—Frontlets and horns of two Bornean Rantins (B. sondaicus lowi)

in the collection of Sir E. G. Loder.

collection of Sir E. G. Loder, represented in text-fig. 125, these

also exhibiting the relatively smaii size of the horns which

appears to characterise many Bornean Bantin heads. The skull

of the Bornean Bantin is characterised by the flatness of the

forehead and the straight intercornual ridge.

906 MR. E. G. BOULENGER ON THE

The dimensions, in inches, of the specimens represented in

text-figs. 123 & 124 are as follows :—

B. sondaicus. B. s. lowi.

Length of horn on outer curve......... 25 21%

Basal ourth con idibuowieies. 604. te es. oe ie 123

Maximum Spam of Worms p20... --cene 28 235

Tip-to-tip interval of ditto ............ 16 13

Length from intercornual ridge of

siull toshipsotmasals ee. tees cece ene 16 133

Minimum width of skull between. base

of horn-core and orbit ............... 9 le

These differences fully justify the right of the Bornean Bantin

to racial distinction, and as none of the comparatively few

synonyms of Bos sondaicus appears referable to that race, I

propose that it should be known as Bb. sondaicus lowi; the skull

and horns represented in text-fig. 124 (B. M. No. 87.2.10.4)

being the type.

It may be added that both the Burmese Bantin or Tsaine, for

which I proposed the name of B. s. birmanicus in the Society’s

‘Proceedings’ for 1898, p. 277, and the Siamese Bantin, for

which I have suggested the name of B. s. porteri, Ibid. 1909,

p- 669, have horns of the general type of those of the typical

race, although those of the Siamese race are more heavily wrinkled

at the base than any Javan horns in the British Museum.

Information is still required as to whether the Bantin occurs

in Sumatra.

56. Notes on the Breeding of the “ Millions” Fish ( Girar-

dinus peciloides). By Epwarpv G. BouLencer, F.Z.8.,

Curator of Reptiles.

[Received October 18, 1912: Read October 29, 1912. |

The little fish Girardinus paciloides, popularly known as

‘“‘ Millions” in Barbados on account of the enormous numbers

frequenting all the shallow pools in that island, is said to be

of great practical value on account of its devouring mosquitoes ;

the absence of malaria, which is so prevalent in the neighbouring

islands, is believed to be due, as first pointed out by Mr. C.

Kenrick Gibbons, to the presence of this fish, for the insect con-

veying the malarial disease breeds only in shallow pools, where

these fish abound and, it is well to add, have no competitors,

G. peciloides being the only freshwater fish inhabiting Barbados.

For this reason, the Society arranged with the Colonial Office to

keep a large stock and to supply specimens to various tropical

stations in the British Empire. The fishes have bred on numerous

occasions in small aquariums in the Reptile and Tortoise Houses,

BREEDING OF GIRARDINUS PCCILOIDES. 907

and I have thus been able to observe certain facts in relation

to their breeding habits, to one of which in particular I wish to

draw attention, namely that in some cases the male breeds before

the assumption of its secondary sexual characters. Before dealing

with this matter it is necessary to mention briefly the sexual differ-

ences, Which are very marked. Females, which may measure up

to 25 mm. in length, are simply of a dull olive-grey in colour,

usually with a dark patch above the anal fin, while the males,

which do not attain quite so large a size, are most conspicuously

ornamented with red, blue, violet and yellow, with a dark ocellar

spot situated in the middle of the body, in front of or beneath

the dorsal fin, and another at the base of the caudal fin, these

spots appearing at least fourteen days before the brilliant hues.

Males are to be further distinguished from females by the

position and prolongation of their anal fin, which is transformed

into an intromittent pairing organ.

The male is remarkably active, and is perpetually courting the

female, going through all kinds of antics in front of her, and as

breeding goes on all through the year, at least in captivity, the

latter is in an almost permanently pregnant condition, and within

a fortnight of having brought forth a brood (such a period

representing the duration of the gestation, at a temperature

of over 70°), once more brings a generation into the world.

It should be borne in mind, however, that a single impregnation

is sufficient for the fertilization of several broods, the embryos of

the second and third generations being already in an advanced

condition when the first young are born. Hach brood comprises

from five to as many as twenty-five fish.

The young fishes grow rapidly : about 4 mm. long at birth, they

double that length in a fortnight or so, when their sex can be

determined by the shape and position of the anal fin; it is not,

however, until at least six weeks after birth that the males

become adorned with the brilliant colours which characterise the

adult, this period again varying according to the temperature of

the water.

The first brood, containing a male which bred before the

assumption of its secondary sexual characters, was produced on

March 11th‘ of this year, and numbered eleven. As is almost

invariably the case, a number of weaklings died within a few days

of their birth, and eight fish remained at the end of a fortnight,

when I found the sexes to be equally divided, which is remark-

able, for as a rule females outnumber males by about three to one.

On May 7th, six young ones were born of this brood, and at the

time none of the males had developed their adult coloration, the

only markings distinguishing them from the females being the

spots beneath the dorsal fin and at the base of the caudal, and it

was in fact not until six days later that one of these acquired its

full sexual livery. The others followed suit on May 17th and

18th. These broods were kept at an average temperature of 80°.

Tn another case, a brood of fourteen fishes, born on June 5th,

908 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

and of which only seven survived, two males and five females,

reproduced their species on September 29th and 30th, the adult

coloration of the males appearing only on October 7th and 16th.

When we consider the fact that these fishes, owing to the warmer

tanks being already occupied, were kept at the low average

temperature of 65°, we must conclude that, like their growth, the

period of gestation must have been slow, and that the male

undoubtedly bred at least a month before acquiring its full

secondary sexual characters.

A parallel case is well known in the male Salmon, which

occasionally becomes sexually mature in the parr condition,

while the Wrass (Coris julis) and the Dragonet (Callionymus

lyva) have been observed by Mr. E. W. Holt also to become

sexually mature prior to having fully developed their adult

characters. In the two latter cases, however, the question is

one of degree only, the characteristic male livery not being

entirely absent, as in Girardinus.

EXHIBITIONS AND NOTICES.

October 29, 1912.

Prof, E. A. Mincuin, M.A., F.R.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 months of

May, June, July, August, and September 1912.

May.

The registered additions to the Society’s Menagerie during the

month of May were 448 in number. Of these, 207 were acquired

by presentation, 60 by purchase, 43 were received on deposit,

72 in exchange, and 66 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 203.

Amongst the additions special attention may be directed to :—

A large collection of Mammals and Birds from Nepal, including

1 Indian Rhinoceros (Rhinoceros unicornis), 1 Kiang (Zquus

kiang), 2 Barasingha Deer (Cervus duvaucelli), 2 Sambur Deer

(Cervus aristotelis), 3 Hog-Deer (Cervus porcinus), 2 Monaul

Pheasants (Lophophorus impeyanus), 2 Cheer Pheasants (Catreus

wallichi), and 2 Wedge-tailed Fruit-Pigeons (Sphenocercus sphe-

nurus); presented on May 21st by H.M. Tar Kine.

A pair of European Bison (Bison bonasus), from Russia, pre

santed by H.G. the Duke of Bedford, K.G., on May 15th.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 909

1 Black Leopard (felis pardus), from Assam, purchased on

May 21st.

1 Branded Palm-Civet (Arctogalidia stigmatica), from Sarawak,

new to the Collection, presented by Granville J. Altman, Ksq.,

F.Z.S., on May 16th.

1 Civet (Viverra civetta), from EK. Africa, and 2 Hgyptian

Mongooses (Mungos ichnewmon), from Zagazig, presented by

H.E. Hassan Hasib Pasha on May 21st.

1 Ural Owl (Syrniwm uralense), from Northern Europe, pre-

sented by E. G. B. Meade-Waldo, Esq., V.P.Z.S., on May 8th.

A collection of rare Reptiles from India, received in exchange

on May 21st, containing, amongst others, 2 Banded Kachugas

(Kachuga dhongoka) and 3 Variegated Snakes (Zamenis fuscio-

latus), new to the Collection, and 2 Gharials (Gavialis gangeticus).

JUNE.

The registered additions to the Society’s Menagerie during the

month of June were 511 in number. Of these 305 were acquired

by presentation, 66 by purchase, 50 were received on deposit,

13 in exchange, and 77 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 205.

Amongst the additions special attention may be directed to :—

A collection of Malayan Animals presented by the Government

of the Federated Malay States and other Donors on June 24th,

including 1 Indian Elephant (Zlephas indicus), 1 Tiger ((elis

tigris), 1 Clouded Tiger (Felis nebulosus), 1 Binturong (Arectictis

binturong), 2 Argus Pheasants (Argusianus argus), 4 Rufous-tailed

Fireback Pheasants (Acomus erythrophthalmus). and 1 Malayan

Peacock-Pheasant (Polyplectron bicalcaratum), besides the

following, which were all new to the Collection, viz. :—1 Short-

tailed Mongoose (Jungos brachywrus), 10: Charlton’s Tree-

Partridges (Arboricola charltoni), 1 Water-Cock ((allicrea

cinerea), 2 Water-Snakes (Acrochordus javanicus), | Bowring’s

Gecko (Hemidactylus bowringii), 1 Bridled Gecko (Hemidactylus

Jrenatus), and 1 Leschenault’s Gecko (Hemidactylus leschenaultii).

2 Malayan Tapirs (Zapirus indicus) § 2, from Palembang,

Sumatra, received in exchange on June 22nd.

1 Grys-bok (Wototragus melanotis) 3, from Cape Colony, pre-

sented by Dr. Louis Péringuey, C.M.Z.S., on June 3rd.

1 Shoebill (Baleniceps rex), from the Sudan, presented by |

Lt.-Gen. Sir Francis Wingate, G.C.V.O., on June 29th.

2 Ocellated Turkeys (Meleagris ocellata), from Central America,

deposited on June 24th.

2 Chestnut-bellied Rock-Thrushes (Petrophila erythrogastra),

from the Himalayas, new to the Collection, received in exchange

on June 17th.

2 Brown-backed Robins (Thamnobia cambaiensis), from India,

new to the Collection, purchased on June 18th.

910 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

1 Temminck’s Robin (Hrithacus komadori), from Japan, new to

the Collection, presented by Wilfrid Frost, Esq., on June 14th.

2 Grey Thrashers (Zowostoma cinerewm), from Lower California,

new to the Collection, presented by H. D. Astley, Esq., on

June 15th.

6 Chilian Siskins (Chrysomitris uropygialis), new to the Col-

lection, presented by Arthur C. Macdonald, Esq., on June 26th.

2 Himalayan White-throated Ground-Thrushes (Geocichla cya-

nonotus), bred in the Menagerie.

12 Russell’s Vipers (Vipera russellz), born in the Menagerie.

JULY.

The registered additions to the Society’s Menagerie during the

month of July were 289 in number. Of these 155 were acquired

by presentation, 13 by purchase, 57 were received on deposit,

25 in exchange, and 39 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 301.

Amongst the additions special attention may be directed to :—

1 African Buffalo (Bubalus caffer) 3, from the Ishasha River,

Uganda, presented by Capt. E. H. Reid on July 29th.

1 Vaal Rhe-bok (Pelea capreolus) 3, from Basutoland, pre-

sented by M. H. Bosworth Smith, Esq., on July 6th.

2 Yellow-headed Wagtails (Motacilla citreola) and 1 Brown

Shrike (Lanius cristatus), from India, new to the Collection,

presented by W. J. C. Frost, Esq., on July 25th.

1 Changeable Hawk-Hagle (Spizaétus limnactus), from India,

new to the Collection, deposited on July 25th.

_ 1 Hemprich’s Gull (Larus hemprichi), hatched in the Menagerie

on July 22nd.

1 Black-and- White Cobra (aia melanolewca), from Dunkwa, Gold

Coast, presented by Dr. H. G. F. Spurrell, F.Z.8., on July 16th.

1 Snake-Fish (Polypterus senegalus), from the Upper Nile,

presented by the Rev. Archibald Shaw on July 9th.

AUGUST.

The registered additions to the Society's Menagerie during

the month of August were 226 in number. Of these 74 were

acquired by presentation, 40 by purchase, 55 were received on

deposit, 9 in exchange, and 48 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 285.

Amongst the additions special attention may be directed to :—

1 White-thighed Guereza (Colobus vellerosus), from West

Africa, presented by Miss Jean Aylwin on August 30th.

1 Roebuck fawn (Capreolus capreolus), from Ross-shire, pre-

sented by the Earl of Altamont, F.Z.S., on August Ist.

1 Mount Morrison Laughing-Thrush (Zrochalopterum morri-

sonianum), from Formosa, new to the Collection, deposited on

August 5th.

MR. R. LYDEKKER ON GAZELLA HAYI. 911

1 Magpie-Tanager (Cissopis leveriana), 3 Australian Crimson

Finches (Veochmia phaéton), 2 Peale’s Parrot-Finches (Hrythrura

peal), and 6 Douglas Quails (Lophortyx douglasi), all bred for

the first time in the Menagerie.

1 Matamata Terrapin (Chelys fimbriata), from Brazil, presented

by Oswald Allen, Esq., on August 10th.

12 Six-banded Cyprinodons (Haplochilus sexfasciatus), from

West Africa, new to the Collection, presented by Dr. H. G. F.

Spurrell, F.Z.8., on August 21st.

SEPTEMBER.

The registered additions to the Society’s Menagerie during

the month of September were 227 in number. Of these 59 were

acquired by presentation, 18 by purchase, 20 were received on

deposit, 24 in exchange, and 106 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 247.

Amongst the additions special attention may be directed to :—

4 Pelzeln’s Gazelles (Gazella pelzelni), presented by Dr. R. E.

Drake-Brockman, F.Z.8., on September 26th.

1 Pelzeln’s Gazelle (Gazella pelzelni), presented by A. Gibbs,

Esq., on September 26th.

1 Capybara (Hydrocherus hydrocherus), presented by Sir Walter

Egerton, K.C.M.G., F.Z.8., and J. J. Nunan, Esq., on September

2nd.

1 White-tailed Gnu (Conncechetes gnu), presented by the

Government of South Africa on September 28th.

1 Kordofan Giraffe (Giraffa camelopardalis antiquorum) ¢ ,

presented by Lt.-Gen. Sir Francis Reginald Wingate, G.C.V.O.,

K.C.M.G., C.B., D.S.O., on September 13th.

1 Brown Tyrant (Myiarchus tyrannulus), from Uruguay, new

to the Collection, presented by Mrs. EH. F. Dickinson on

September 3rd.

1 Great Indian Bustard (Hupodotis edwardsi), new to the

Collection, presented by Sir John Hewitt, K.C.8.1., on September

30th.

2 European Bee-eaters (Merops apiaster), received in exchange

on September 11th. .

2 Snake-headed Fishes (Ophiocephalus punctatus), from India,

new to the Collection, received in exchange on September 7th.

‘* GAZELLA HAYI” = GAZELLA FUSCIFRONS.

Mr. R. Lypekxker, F.R.S., F.Z.8.*, contributed the following

note -—

“ As the result of an unfortunate accident, namely the trans-

position of the registration labels of two gazelles received

simultaneously at the British Museum, I find that I have

described and figured a specimen of the Seistan Gazella fuscifrbns

* By permission of the Trustees of the British Museum.

919 MRS. R. HAIG THOMAS ON THE

as a new African species under the name G. hayi (Proc. Zool.

Soc. 1911, p. 961, text-fig. 193 A). My apologies are due both

to the Society and to naturalists in general for this most unfor-

tunate mistake.

The text-figure of the so-called @. hayi will serve as an

illustration of the male of @. fuscifrons. The gazelle collected

by Mr. Hay in Algeria is, I believe, G. isabella.”

The Rev. T. R. R. Sreppine, M.A., F.R.S., F.Z.S., communi-

cated a memoir on the Crustacea Isopoda collected by the

‘ Porcupine* Expedition in 1869-1870, in which he described

one new family, two new genera, and four new species.

This memoir will be published in the ‘ Transactions’ in due

course.

Mrs. Rosz Hare Tuomas, F.Z.S., exhibited the eggs of Phasi-

anus formosanus, P. versicolor, and their F.1 and F. 2 offspring,

and made the following remarks :—

“ Last February I exhibited to the Society a series of Pheasant

skins of an experimental cross between a Formosan female and a

versicolor male, illustrating the transmission by the male parent

to his female offspring of most of the female characters of his

Text-fig. 126.

Eggs ot P. versicolor, F.1, Fo.X Ve., F.2, Fo.X Ve.X Ve., and P. formosanus.

race: further evidence of this is now exhibited, namely, the eggs

of the two parent species and those of their F.1 and F. 2 offspring.

It is plain that a considerable difference in dimensions exists

between the eggs of P. formosanus, the larger, and P. versicolor

the smaller of the two: the expectation was that the eggs of the

offspring would resemble in size the egg of the female parent

species rather than that of the male parent species ; but, on com-

paring them, it is clear that the eggs of F.1 and F. 2 are simply

EGGS OF PHEASANTS. 913

P. versicolor. A photograph (text-fig. 126) by Mrs. Peter Haig

Thomas of the four eggs shows very accurately the comparative

sizes.

The measurements of these eggs are as follows :—

Girth of length. Girth of breadth.

P. formosanus ......... 57% ins. 412 ins,

de COPSICOLOT 23.2 ae Oe Al2 44. ,,

iB, 1 Rape Vier ae: 413, 45. ,,

F. 2, Fo. x Ve. x Ve. . Als ,, 45. ,,

showing a slight increase in bulk in the eggs of F.1 and F, 2

over that of the versicolor.

Another character of which notice was omitted in February

was the iris colour: unfortunately F.19 died eggbound last May

before it occurred to me to make an examination. The male

and female versicolor differ in the colour of the iris, as do also

the male and female formosanus. The male versicolor has a pale

yellow iris, the female a soft brown centre with an outer ring of

pale yellow: the male formosanus has a red-brown iris, the female

a dark brown centre with outer ring of red-brown. Holding

formosanus, versicolor, and F.1 hens close together, it is seen that

F.1 has the female versicolor iris with the ring-colour a very

slightly deeper shade of yellow. Again, outstretching the wings of

these three hens side by side, it is observed that besides the secon-

daries and primaries the two groups of major coverts overlying

these feathers in F.1 9 also correspond exactly in pattern and

colour with the same groups in the versicolor 9 .”

At the conclusion of the Scientific Business, Sir Epmunp G.

Lover, Bt., V.P.Z.S., showed a large number of slides, including

a fine series of coloured subjects, and a number of living speci-

mens, to demonstrate the capacity of the new electric lantern

which he had recently presented to the Society.

INDEX

SCIENTIFIC NAMES,

Acomus

erythrophthalmus, 909.

Acrochordus

Jjavanicus, 909.

Actinia, 519.

A®lurosaurus

Jelinus, 863.

striatidens, 859, 863,

875.

Agrotera

pyrosticta, O01.

Alee, 774, 778, 779.

Aleyonium, 518, 527.

Allopora, 892, 894.

Alopecorhinus, gen. nov.,

859, 864, 865.

parvidens, 859, 864,

875.

Aloposaurus, 862.

Ambia

—— jovaguinensis, 546, 549,

55D.

Amia, 608.

Ammothea, 5il.

virescens, 527.

Ammothée, 510, 511.

virescens, 511.

Amphilius

grandis, 675.

oxyrhinus, 672,

676.

Ancistrodon

piscivorus, 633.

Andrya, 587.

Anoplocephala

hyracis hepatica, 593,

Od.

Anoplotenia, 584,

677.

dasyuri, 694, 846, 847.

Anthobosea, 696, 714,

724.

ethiops, 745, 748.

603,

Anthobosea

albomacula, 725.

albomaculata, 728, 742,

746, 758.

albopilosa, 727, 732.

antennata, 728, 748,

744 746.

anthracina, 727, 733,

754,

apicalis, 745.

arabica, 720, 728, 739,

740.

argenteocincta, 7217,

australasie, 725, 745—

749.

australis, 727-729, 745,

754.

hidentata,

Widers

bipunctata, 728, 742.

746,

152;

bipustulata, 742, 746, |

78),

carbonaria, 728, TA,

742, 745.

chilensis, 728, 741, 746.

clypeata, 727, 735, 751,

7538, 754.

cognata, 727, 731.

crabroniformis, 'T46.

erassicornis, 736, 745,

751, 753.

donaldsoni, 728, 737.

errans, 746, 751.

erythronota, 728, 737,

739, 746, 753, 754.

erythropyga, 728, '744,

7A6.

fasciculata, 736.

Jastuosa, 727, 729, 753.

flavicornis, 727, 7830,

731, 732.

flavopicta, 746, 752.

Anthobosea

Srenchi, 745, 749, 750.

gilest, 745, 747.

inornata, 736.

insularis, 728,

754,

jheringt, 744.

lacteipennis, 746, 752,

753.

levifrons, 727, 733,

734.

lagardei, 745, 750.

leucospila, 739.

longipalpa, 745, 750.

melanaria, 728, '736.

minima, 728, TAV.

moderata, 45, 749,

oan

natalica, 728,738, 739.

nigra, 745, 748.

nigripennis, T45.

nubilipennis, 727, 730,

(es

occipitalis,

734.

sauakinensis, 728, 740.

signata, 727, 729.

strandi, 727, 731.

torresensis, 745, 747.

wnicolor, 727, 738, 734.

varipes, 745, 747, 748.

Apenesia, 699.

Aphrocallistes

whiteavesianus, 787.

Aphytoceros

grosalis, 554.

Apomatus, 788, 793.

timsti, 784, 787, 805.

Aporina, 849.

Aptenodytes

pennantt, 671.

Ara

severa, 806,

737,

696, 727,

Xvi

Araucaria, 726.

Arboricola

charlioni, 909.

Arcosealpellum, 538, 539.

Arctictis

binturong, 909.

Arctogalidia

stigmatica, 909.

Arges

cirratus, 666, 670.

Argulus, 765.

Argusianus

argus, 909.

Arvicola

arvalis, 839.

Asellus

aquaticus, 660.

Astacus, 519.

Aulacodes

brunnealis, 548.

metaloxalis, 548.

Austrotiphia

kirbyi, 739.

Avitellina, 848.

Bagrus

urostigma, 675.

Baleeniceps

rex, 909.

Balanus

nubilus, 798, 800.

Barbus

argyrotenia, 672, 674,

676.

erlangeri, 674.

mimus, 672, 674, 676.".

natrobiensis, 674.

neglectus, 675.

percivali, 674.

zanzibaricus, 674.

Belemnitella

mucronata, 530.

Bertiella, 677, 694, 844.

Bison

bonasus, 908.

Boa, 621, 643.

constrictor, 628, 634.

divinilogua, 628, 633,

634.

reticulata, 644.

Bos

sondaicus, 902, 903, 906.

— birmanicus, 906.

— lowi, 904-906.

— porteri, 906.

Braunsomeria, gen. nov.,

696, 697, 700,

atriceps, 696, 699.

quadraticeps, 696, 699-

758, ‘754,

INDEX OF SCIENTIFIC NAMES.

Brevicella, gen. nov., 546,

549

emarginata, 546, 549,

55D.

Bubalus

caffer, 910.

Calamachrous

albipunctulis, 546, 554,

555.

Calliobothrium, 693.

Callionymus

lyra, 908.

Calliptilus

solitarius, 806.

Callosila, 726.

Camelus

Canis

familiaris, 556.

Capra

pyrenaica, 754.

— victorie, 756.

Capreolus, 774, 778.

capreolus, 910.

Caprinia

diaphanalis, 551.

Cariacus, 774.

Cariama

eristata, 557, 558.

Carijoa

rupicola, 509.

Catreus

wallichi, 908.

Causus

rhombeatus, 634.

Cerastes

vipera, 622.

Ceratodus

forsteri, 607-612.

Cercaria

ordinata, 767, 768, 769.

Cercopithecus, 606.

Cervus, 778.

affinis, 564-575, 781.

albirostris, 558, 574.

aristotelis, 908.

canadensis, 574, 774,

775, 777.

— wardi, 575.

cashmeriensis, 565, 566.

— macneilli, 570.

duvaucelli, 783, 908.

elaphus, 574.

eldi, 776, '780.

hanglu, 565-575, 775,

With

kansuensis, 574.

macneilli, 571-574.

porcinus, 908.

\"W/

Cervus

pygargus, 559.

thoroldi, 574.

wallichi, 556-575.

canthopygus, 575.

Cheetostomus

Jischeri, 667.

lepturus, 666, 667, 670.

marginatus, 667.

palmeri, 666, 667, 670.

paucispinis, 666, 667,

670.

Chapmannia, 600.

lapica, 603.

Chelys

jimbriata, 911.

Chimarrhornis

leucocephala, 556.

Chironephthya

variabilis, 523, 524.

Chitinopoma

fabricii, 790.

greenlandica, 790, 805.

Chloéphaga

poliocephala, 556.

Chonopeltis, 765.

inermis, 766.

Chrysomitris

uropygialis, 910.

Circeis

armoricana, 796, 797.

Cissopis

leveriana, 911.

Cistecephalus

arctatus, 871.

nicrorhinus, 871.

Cittotzenia, 604, 677.

Clarias

lazera, 675.

Clarotes

laticeps, 675.

Clupeosoma

polusalis, 548.

Ceelopeltis

monspessulana, 617.

Colobosila, 726.

Coiobus

abyssinicus occidentalis,

vellerosus, 910.

Coluber

esculapii, 622, 638,

634.

corais, 639.

natric, 613.

Connochetes

gnu, 911.

Corallus

caninus, 643.

cookit, 628, 643.

madagascariensis, 643,

Poe

INDEX OF SCIENTIFIC NAMES.

Coris

julis, 908.

Coronella

catenifer, 626,

getula, 617.

levis, 622.

Corophium

acutum, 664.

lacustre, 656, 658, 664,

668.

triaenonyx, 664,

volutator, 658.

Cosila, 724, 725.

apicalis, 745.

argenteocincta, 732.

australis, 728.

biguttata, 729.

cearbonaria, 741.

chilensis, 725, 741.

donaldsoni, (05, 737.

erythropygia, 744.

tnornata, 736.

insularis, 737.

jheringi, 748.

(Callosila) australis,

728.

(—) fasciculata, 736.

(—) flavicornis, 730.

(—) minuta, 734.

(—) signata, 729.

Crocidolomia

binotalis, 553.

Crucigera

irregularis, 786, 805.

zygophora, 786, 805.

Cryptohelia, 885.

Curena

externalis, 548.

Curicta

oppositalis, 504.

Cyanocorax

affinis, 806.

Cyclorchis, 690.

Cynognathus, 874.

Cysticercus

fasciolaris, 838, 850.

longicollis, 839, 840.

Dama, 777.

dama, 781.

mesopotamica, 781.

Dasyurotenia, gen. nov.,

677, 680, 683, 694,

695, 823.

robusta, 677, 678, 679,

684, 694, 834.

Dasyurus

ursinus, 617.

Dausara

amethystina, 553.

Davainea, 591, 592,

606.

Dendrohyrax

dorsalis, 671.

Dendrolagus

ursinus, 806.

Dendronephthya,

510, 518, 521.

diseiformis, 506, 519,

520, 527.

Diadectes, 860.

Dizlurodon

whaitsi, 868.

Dicroceelium, 854.

infidum, 8)4, 856.

Dicrogenium, 725.

Dicynodon, 871, 872.

laticeps, 859, 876.

leoniceps, 869.

lutriceps, 859, 868, 870,

876

505,

psittacops, 809, 869,

876.

tigriceps, 868.

Didelphis

virginiana, 556.

Dilepis, 847.

Dimorphoptera,

725.

anthracina, 733.

clypeata, 739.

cognata, 731.

Jastuosa, 729.

levifrons, 734.

morosa, 729.

nigripennis, 728.

sabulosa, 733.

scoliiformis, 728.

signata, 729.

wnicolor, 734.

Dioicocestus, 844, 848.

acotylus, 831.

Diplostomum

sirtale, 767, 768, 769.

Dipteropeltis, gen. nov.,

763

724,

hirundo, 763, 766.

Dipylidium, 587.

Discognathus

dembeensis, 679.

Dissemurus

paradiseus, 556.

Distichopora, 892, 894.

Distoma, 856.

Dolops

longicauda, 763, 765.

Dorcephalus, 773, 778.

Heeasaurus

priscus, 875.

Proc. Zoou, Soc.—1912, No, LXI,

xX1X

Echidna, 814.

Echinococeus, 837.

Eirone, 699, 725.

Elaphurus, 773, 778, 780,

783.

davidianus, 773, 74,

777, 179.

Hlaphus

indicus, 909.

Elis, 725.

alicia, "737.

apimacula, 721.

capitata, 711.

combusta, 724, 754.

dimidiata, 716.

dubia, 721.

ephipprum, 724.

Sedtschenkvi, 721.

major, 696, 723.

sellowt, 696, 722.

tricolor, 721.

xanthocera, 709.

(Mesa) abdominalis,

703, 707.

(—) adelogamia, 703,

706

(—) alicie, 696, 703,

704, 753, 754.

(—) ametalla, 704,

713.

(—) apicipennis, 696,

703, 707

(—) apimacula, 714,

HAD!

(—) asmarensis, 704,

(—) auriflua, 696, 703,

705, 706.

(—) bengalensis, 714,

716

(>) burmanica, 714,

ah).

(—) capensis, 709.

(—) capitata, 704,

710.

(—) carbonaria, 718.

(—) claripennis, 714,

717, 718.

(—) clavata, 713.

(—) diapherogamia,

706

(—) dimidiata, 713,

714, 715.

(—) dimidiaticornis,

714, 720, 753.

(—) dubia, 714, 719.

(—) erythropoda, 703,

709

(—) exlensa, 714, 721.

(—) fedtschenkot, 714,

Ge Suscipennis, 713,

Pal ‘and weed 704,

706.

(—) hottentota, 703,

708.

(—) Fs, 704, 706,

ee oe ta, 696, 704,

val

(=) Pihlog 713.

(—) wtnotata, 703,

709

(—) leta, 714, 721

(—) longiventris, 696,

704, 712, 753, 754.

(—) madalensis, 718,

714, 715, 721.

(—) mandibularis, 713,

714, 715.

(—) nodosa, 704, 707,

713.

(—) nursei, 714, 721.

(—) opacifrons, 714,

G19.

(—) peringueyi, 703

708.

(—) petiolata, 713,

714, 717-719, 721.

(— 2 picticollis, 714,

720.

eel re 708,

(— Me cre 713.

(—) rothneyt, 714,

717

(—) ruficeps, 704, 706,

754.

(—) — tes

704, 7

oe — ee ogamia,

“ave — ~ ruficeps, 704.

& rufo-femorata,713.

(=) saussuret, 703,

709

(—) spoliata, 696, 704,

elle

(—) torrida, 703, 708.

(—) tricolor, 714, 720,

753, 754.

(—) ustudata, 714, 718.

(—) xanthocera, 703,

709.

Emydochampsa,

nov., 875.

platyceps, 875.

' Emydops, gen. nov., 859,

‘ 871.

minor, 859,871, 876. |

gen.

Endothiodon, 871.

bathystoma, 867, 875.

platyceps, 859, 867,

875, 876

uniseries, 866, 867, 875.

Endothiodon

whaitsi, 859, 866, 867,

875, 876.

Entephria

glaucias, 5d0.

gr isealis, 546, 550, 555,

mioswart, 546,550, 55d.

Epimys

norvegicus, 671.

Equus

burchellt, T57.

— chapmanni, 757.

— selousi, 757.

— transvaalensis, 757,

760.

— wahlbergi, 757.

kiang, 908.

Erithacus

komadorz, 910.

Errina

antarctica, 894.

capensis, 894.

carinata, 878, 880,

893.

cochleata, 880, §898,:

894.

dabneyi, 880 893, 894.

echinata, 880, 8&1,

893.

fissurata, 878, 880,

887, 894.

glabra, 878, 880, 893,

894.

gracilis, 878, 881, 894.

horrida, 878, 880,

881, 893.

labiata, 878, 880,

888. 893.

macrogastra, 878, 879,

894.

moseleyt, 894.

nove zealandie, 876,

877, 894.

pourtalesti, 880, 881,

ramos, 878-881, 893.

(Labiopora) — antare-

tica, 881, 8&2, 887,

890.

(—) aspera, 881, 888-

891, 894, 895.

(—) capensis, 878, 886,

890-893, 895, 896.

(—) gracilis, 889, 890,

(—) moseleyi, 881, 887,

888, 890.

INDEX OF SCIENTIFIC NAMES.

Errina

(Labiopora) nove ze-

landie, 882-884,

889-893, 895, 896.

(Spinipora) echinata,

881i, 895

Erythrura

pealer, 911.

Eryx

conicus, 634.

Jaculus, 633, 634.

Johni, 633, 634.

Hsoterodon

uniseries, 875.

whaitsi, 875.

Hunectes

marina, 854, 856.

murinus, 628, 631,

634.

noteus, 634.

Eupodotis

edwardsi, 911.

Eurytrema, 854.

Eutropius

depressirostris, 675

Felis

nebulosus, 909.

onea, 0955, 806.

pardus, 909.

tigris, 909.

uncia, 555.

Fiber

zibethicus, 822,

840.

839,

Galechirus, 861.

Galeops, gen. nov., 859,

860, 861.

whaitst, 859, 860, 87d.

Galepus, 861.

Galesaurus, 874.

Gallicrex

cinerea, 909.

Gammarus

duebenti, 657, 661.

locusta, 657, 661-665,

pulex, 657, 658, 660,

661.

zaddachi, 656-658,

660, 661, 663, 665.

Gavialis

gangeticus, 809.

Gazella

fuscifrons, 911, 912.

hayi, 911, 912.

isabella, 912.

pelzelni, 911.

Geocichla

cyanonotus, 910.

Geotiphia, 724, 726.

Giratta

camelopardalis

- quorum, 911.

rothschildi, 771,

772.

— schillingsi, 772.

— thornicrofti, 771,

PRR

—- tippelskirchi, 771,

Ti2.

Girardinus

peciloides, 906.

Gly phodes

cesalis, 552, 553.

magnificalis, 046, 552,

595.

nigricincta, 546, 52,

555.

pfeiffere, 546, 554,

555.

polyzonalis, 552.

pseudocesalis, 546, 552,

553, 505.

anti-

Gorgonia

suberosa, 524.

Gorgonops

torvus, 861.

Govia, 770.,

Graphiurus

spurrelli, 555,

(rypaeétus

barbatus, 806.

Hancockia

eudact ylota, 770.

Haplochilus

sexfasciatus, 911.

Hemidactylus

bowringti, 909.

trenatus, 909.

leschenaultii, 909.

Hemistomum, 767, 770.

Hemitragus

jemlarcus, 556.

Hesperornis, 819.

Holothuria, 519.

Hyzena

hyena, 559.

Hyalopomatopsis, 792.

langerhansi, 791.

marenzlleeri, 791.

occidentalis, T91.

Hydrocherus

hydrocherus, 911.

Hydropotes, 774.

Hymeuolepis

acuta, 687.

murina, 822.

reticulata, 603.

villosus, 331.

INDEX OF SCIENTIFIC NAMES.

Hyracotxnia, gen. noy.,

576, 579, 593, 604.

capensis, 576.

hyracis, 576, 594, 596,

597, 601, 6005.

procavie, 604, 605.

Hyroides

dianthus, 804.

gronlandica, 790.

norvegica gronlandica,

Icterus

axanthornus, 806.

Tetidvgnathus

hemburyi, 859, 865,

875.

parvidens, 865, 866.

| Ictidopsis, gen. nov., 859,

872, 874.

elegans, 859, 872, 873,

876.

Ictidosuchus, 861.

Inermicapsifer, 581, 582,

590, 59-4, 595, 599,

603, 607, 688, 692,

847.

sp., 593, 605.

capensis, 576, 577, 580,

586, 588, 591, 592,

593, 600, 602, 606.

criticus, 593.

hyracis, 577, 578, 579,

585, 586, 587, 588,

591, 592, 606.

interpositus, 592.

pagenstecheri, 593.

settii, DTT, 578, 592;

606.

TIswara, 715.

Jassa, 656.

Kachuga

dhongoka, 909.

Kistecephalus

arctatus, 871.

Labeo

cylindricus, 673.

neumanni, 673.

percivali, 672, 673,

676.

_ Labiopora, 876, 879, 889,

881. 884, 885.

antaretica, 887, 890,

893, 894.

‘oe

Labiopora

moseleyi, 878, 888, 893,

894.

Lacerta, 614, 626,

631, 644.

muralis, 643, 807.

Lachesis

gramineus, 625,

Leospira, 793, 799.

Lama

pacos, 526.

Lanius

cristatus, 910.

Larus

hemprichi, 910.

Leptocheirus

pilosus, 656, 658.

Ligula, 824,

Linstowia, 677.

Licheterodon

madagascariensis, 634.

Lithophyton

arboreum, 511.

Lithophytum, 511, 513.

arboreum, 511, 51%,

Dit

Lobophytum, 527.

Lomanotus

genei, 770.

Lophophorus

impeyanus, 908.

Lophortyx

douglast, 911.

Lycosuchus, 862.

Lystrosaurus, 873, &74.

627,

Macalla,

arruensis, 546, 547,

55d.

mioswart, 8546, 547,

BO},

olivalis, 546, 547, 555.

syrichtusalis, 547.

Madrepora, 519.

Margarosticha

plumbealis, 546, 549,

005.

Maurillus, 725.

Mazama, 778.

bricenti, 783.

Meleagris

ocellata, 909.

Melitodes

albitincta, 506, 525.

Mera

pusilla, 797.

Meria

abdominalis, 703.

Merops

apiaster, 911.

Xxil

Millepora

aspera, 879, 881.

Moniezia, 604, 677, 692,

695.

Mormyrops

deliciosus, 673,

Mormyrus

kannume, 673.

Motacilla

citreola, 910.

Mungos

brachyurus, 909.

ichnewmon, 909.

Myiarchus

tyrannulus, 911.

Myzine, 697, 725.

abdominalis, 696, 703,

754.

albomaculata, 742.

anthracina, 714, 733.

apimacula, 719.

bipunctata, 742.

bengalensis, 716,

braunsi, 696, 700, 753,

754.

burmanica, 722.

capitata, 710.

carbonaria, T41.

ceylonica, 717.

claripennis, 718.

clavata, 713.

combusta, 696, 714,

724.

constrictiventris, 696,

701, 754.

dimidiata, 715.

dimidiaticornis, 720.

erythropyga, 144.

Ffastuosa, 729.

flavicornis, 741.

fuscipennis, 717.

hortata, 718.

insularis, 737.

leta, T2\.

lecointei, 742.

levifrons, 734.

madraspatana, 715.

nodosa, 713.

orientalis, 714.

pallida, 714.

petiolata, 717.

rothneyt. 717.

ruficeps, 700.

rufifrons, 702.

sabulosa, 733.

sauakinensis,

740.

signata, 729.

stigma, 696, 699, 753,

754.

swalei, 700, 701.

696,

Myzine

tricolor, 720.

umbratica, 696, 702.

unicolor, 734.

violaceipennis, 715.

axanthocera, 709.

Nacoleia

perdentalis, 551.

Naia

melanoleuca, 910.

Naja

tripudians, 851.

Neochmia

phaéton, 911.

Neogensis

flavoplagialis, 548.

Nephthée, 510, 511.

Nephthya, 510, 512, 518,

519.

albida, 516.

bed fordi, 505, 506, 514,

516, 517, 527.

celosa, 517.

chabrolii, 511, 512, 517,

518, 527.

cordierit, 511,

elongata, 516.

Noorda

arfakensis, 546, 558,

555.

nigripunctalis, 553.

Nototragus

melanotis, 909.

Numida

ptilorhyncha,

605.

Narsea, 725.

Nymphula

chryseis, 548.

polystictalis, 548,

Nytbosaurus, 873.

browni, 859, 874,

876.

larvatus, 872, 874.

576,

Odocoileus

sp., 773, 774, 778, 780,

782

americanus, 783.

hemionus, 783.

savannarwmn, 781.

Odontothynnus, 696,

724.

bidentatus, 752.

lacteipennis, 752.

Omphisa

repetitalis, 553.

INDEX OF SCIENTIFIC NAMES.

Omphisa

variegata, 546,

595.

Oochoristica, 677, 847.

Ophiocephalus

punctatus, 911.

Ophiophagus

bungarus, 617, 625.

Opisthoctenodon, 859,

868.

Ornithorhynchus,

866.

Otiditzenia,

591.

eupodotidis, 583.

Otocinclus

maculipinnis, 666, 668,

670.

nigricauda, 668.

perforatus, 668.

Oudenodon, 859, 872.

bolorhinus, 868.

magnus, 866.

Ovis

burrhel, 556.

5d3,

814,

584, 590,

* Oxyloricaria

leightont,

670.

tamane, 666, 669, 670.

666, 669,

Pagyda

botydalis, 551.

Jumosa, 546, 5o1,

555.

Paradexiospira, 793.

Paraleospira, 799.

Paraspongodes

crassa, 506, 523.

Pareiasaurus, 860, 861,

863, 869.

Pelea

capreolus, 910.

Pelias

berus, 643.

Pelophilus

madagascariensis, 612,

633.

Peltidocotyle, 694,

Perameles, 819.

Perisyntrocha

suffusa, 546, 548, 355d.

Petrophila

erythrogastra, 909.

Phasianus

Jormosus, 540,541, 545,

912, 913.

genneus, 540,

torquatus, 540.

versicolor, 540-542,

545, 546, 912, 913.

Phryganodes

analis, 551.

centrabalis, 551.

erebusalis, 551.

tetraplagalis, 551,

usitalis, 551,

Phyllobothrium, 694.

Piletocera

diplatyla, 548.

eyimisatis, 548.

torsicutalis, 548.

Piloerocis

angulifora, 546, 553,

DOV,

Platophrys

podas, ad6.

Platynosomum, 854.

Plecostomus

honde, 666, 670,

Plerocercus, 839.

Plesia

abdominalis, 707.

asmarensis, 712.

carbonaria, 710, 718.

continua, 696, 703.

dubia, 719.

erythronota, 696, 738.

fedischenkoi, 719.

tneisa, 712, 713.

leucospila, 696, 739.

mandalensis, 716.

melanaria, 696, 736,

nurset, 721.

picticollis, 720.

reticulata, 713.

rufo-femorata, 713.

tartara, 719.

(Mesa) abdominalis,

707.

(—) adelogamia, 706.

(—) atopogamia, 706.

(—) diapherogamia,

706.

(—) disjuncta, 706.

(—) erythrepoda, 709.

(—) heterogamia, 706.

(—) hottentota, 708.

(—) hova, 707.

(-—) tnnotata, 709.

(—) mandibularis, 715.

(—) opacifrons, 719.

(—) peringueyt, 708.

(—) purpureipennis,

715

Ld.

(—) saussurei, 709,

Pecilotiphia

albomaculata, 719,

Pollicipes, 530, 537,

Polyplectron

hicalcaratum, 909.

chinguis, 557.

INDEX OF SCIENTIFIC NAMES.

Polypterus

senegalus, 910.

Porella

antarctica, 881, 887.

Pristerodon, 859, 868.

Pristerognathus

platyrhinus, 859, 863,

864, 875.

polyodon, 864.

Procayvia

capensis, 576, 593.

Procolophon, 874.

Prodicynodon

beaufortensis, 859, 867,

376.

pearstonensis, 868.

Proptopterus

annectens, 807.

Prorodes

mimica, SOI.

Prospirorbis, 792.

Proterogynotznia, 685.

Protolwuspira, subgen.

nov., 784, 792, 7ys.

Protula

atypha, 790.

capensis, 790.

diomedee, 790.

pacifica, 784, 785, 805.

superba, 79U.

Pseudomeria, 699,

Pseudotantalus

ibis, 556.

Psilacabaria

gracillima, 506, 525,

Pygospila

tyres, 58,

Pyrausta

alentialis, 554.

deductalis, 554.

jlammealis, 546,

555.

occultilinea, 554,

Python

molurus, 638.

reticulatus, 806.

reguus, 628.

sebe, 633. 634.

spilotes, 626.

Rangifer, 774, 778.

Rehimena

cissophora, 551,

Rhinoceros

unicornis, 908.

Romanchella, 793.

Rucervus, 778.

duvaucelli, 776, 780.

Rusa, 776, 778, 780.

aristotelis, 775, 781.

XXil

Sameodes

polythliptalis, 553,

Sarcophytum, 527.

Scealpellum

accumulatum, 528-530,

537-539.

arcuatum, 528-531.

comptum, 537, 538.

darwintanum, 680,

539,

Jossula, 531, 583, 537,

538, 539.

gulicum, 538.

hastatum, 530, 539,

maximum, d31.

— cylindracewm, 530,

538, 539.

simplex, 528, 580,

538, 539.

solidulum, 530, 537,

538, 539.

trilineatum, 528, 5380,

531, 538, 539.

Sceliodes

grisealis,

555.

Sclerogorgia

suberosa, 524.

Selerophytum, 509.

palmatum, 507, 508.

pinnulatum, 505, 506,

507, 508, 527.

viride, 508.

Seylacosaurus, 861, 864.

Scymnognathus, gen,

noyv., 859, 861, 862.

whaitsi, 859, 861, 875.

Secymnosaurus, 863,

Semnopithecus

pileatus, 806.

Serpula

columbiana, 784, '785,

786, 801, 802, 805.

Jukes, 786.

spirillum, 796.

splendens, 784,

786.

triqueter, 780.

vitrea, 793.

zygophora, 786.

Sierolomorpha, 725,

546, 550,

785,

Siphonogorgia

variabilis, 506, 523,

Sphenocercus

sphenurus, 908.

Sphenodon, 643.

Spinipora, 876, 878, 879,

880, 881, 891,

Spirorbis

affinis, 802.

aggreyatus, 795.

XXIV

Spirorbis

ambilateralis, 784,792,

795, 798, 799, 805.

argutus, 795.

armoricanus,

797.

bellulus, 794.

beneti, 795.

bernard. 795.

borealis, 796, 804.

cancellatus, 792,

796.

carinatus, 802.

claparedet, 795,

comptus, 794.

cornu-arietis, 795, 798.

corrugatus, 794,

evolutus, 793.

fabricii, 802.

Joraminosus, 794.

jformosus, 794.

granulatus, 802.

koehleri, 795.

levis, 795.

langerhansi, 795, 801,

805.

lebruni, 795.

levinsent, 795,

lucidus, (96.

malardi, 795.

marion, 794.

mediterrancus, 795.

medius, 784, 793, 795,

800, 805.

militaris, 795.

morchi, 795, 801, 805.

- pagenstecheri, 794.

patagonicus, 795.

perrieri, 795.

pseudocorrugatus, 794.

pusilla, 794, 797.

pusilloides, 794, 797,

805.

quadrangularis, %95,

802.

racemosus, (84, 795,

793, 799, 805.

rugatus, 794.

semidentatus, 794.

similis, 795.

spirillum, 794,

797, 8085.

spirorbis, 795.

validus, 795, 800, 805.

variabilis, 795, 802.

verruca, %93, 799,

800.

violaceus, 794.

vitreus, 793, 794, 805.

794,

(Dexiospira) spirillwm, |

796.

INDEX OF SCIENTIFIC NAMES,

796, |

Spongodes,

€pirorbis

(Leospira) asperatus,

800.

(—) langerhansi, 801.

(—) medius, 800.

(—) morchi, 801.

(=e),

802.

(—) validus, 800.

(—) variabilis, 802.

(Paradexiospira)

vitreus, 793.

(Protoleospira) ambi- |

lateralis, 798.

Spizaétus

limnaétus, 910.

510, 511,

518; 519, 520, 521.

ceclosia, 512, 513, 518.

nephihyeformis, 512,

518

Sponodia, 518, 520, 521.

Stereonephthya, 510, 518,

519, 520.

lutea, 505, 506, 521,

528, 527.

Stericta

548,

rurealis,

509.

546,

Styphlodora

bascaniensis, 855, 856.

condita, 855, 856.

horrida, 851, 854.

najeé, 851, 852, 854.

serrata, 854, 855.

similis, 855.

Suberogorgia

suberosa, 506, 524.

Sylepta

dinawa, 546, 551, 555.

leucodonta, 551.

polydonta, 551.

| Synodontis

geledensis, 675.

schall, 675.

Syrnium

wralense, 909,

Tachypterus

crassicornis, 'T51.

Teenia

crassiceps. 839.

crassicollis, 838.

ghondhorensis, 598.

paronai, 593.

spatula, 5Y3.

(Anoplocephala),

gondokorensis, 605.

(—) spatula, 605.

quadrangularis,

‘'Taognathus

megalodon, 867.

Tapinocephalus

atherstonei, 859.

. Tapirus

indicus, 909.

Taurops, gen. nov., 859. ~

macrodon, 859, 875.

Taurotragus

oryx, 806.

Telesto

prolifera, 509, 510.

' rtiser, 509.

rupicola, 506, 509,

510.

(Carijoa) rupicola,

509.

Tetrabothrium, 585.

Thamnobia

cambaiensis, 909.

Thaumalea

amhersti, 544.

obscura, 544.

picta, 544,

Thecla

bertha, 897, 902.

biblia, 901.

campa, 901, 902.

castrena, 900, 9O2.

datitia, 901, 902.

esmeralda, 900, 902.

Jancia, 897, 902.

Fernanda, 899, 902.

gaina, 898.

gispa, 896.

hamila, 896, 902.

janias, 900.

Jjapola, 898, 902.

lemona, 899.

molena, 8Y9, 902.

nora, 899, 902.

nubilum, 899.

phrosine, 900.

punctatum, 897.

schausa, 898, 902.

sicrana, 897, 902.

tadita, 901.

Thelognathus, 874.

Thrasaétus

harpyia, 806.

Thynnus, 725.

antennatus, 743.

cathreinii, 746.

jischert, 747.

reischit, 748.

stolzii, 748.

Thysanotenia, 576, 584,

gambiana, 579, 581,

582, 583, 585, 586,

587, 590, 602, 606.

INDEX OF SCIENTIFIC NAMES,

Thysanotenia

lemuris, 583, 585, 606,

686.

Tilapia

alcalica, 676.

grahami, 676,

nilotica, 675.

percivalt, 672, 676.

Tiphia

capensis, 709.

bipunctata, 742.

Totanus

calidris, 647.

Toxostoma

cinereum, 910,

Triplotznia, 677.

Trochalopterum

morrisonianum, 910.

Tropidonotus

fasciatus,

633.

natrix, 612, 613, 632,

634, 640, 643, 646,

647.

ordinatus sirtalis, 767.

624,

Tropidonotus

tesselatus, 622.

Tympanuchus

americanus, 596,

Urocystidium, gen. nov.,

840),

gemmiparum, 822, 824,

826, 828, 833, 840,

841.

Ursus

maritimus, 556.

Vipera

berus, 622.

russelli, 910.

Viverra

civetta, 909.

Wrightella, 526.

robusta 505, 506, 525,

527.

XXV

Xanthomelzna

schematias, 550.

Xenia, 509.

Xenocara

bufonium, 668.

heterorhynchus,

668, 670.

multispinis, 666,

666,

668,

stigmaticum, 668.

Xenopharynx, gen. nov.,

851.

solus, 851, 852.

Zamenis

fasciolatus, 909.

gemonensis, 631,

634.

Zinckenia

lophoceralis, 551.

Zschokkea, 590.

Zschokkeella, 576,

602, 603, 606,

688, 692.

linstow?, 590.

633,

591,

607,

IND

ILLUSTR

Atlurosaurus striatidens, Pl. XCI.

p. 859.

Alopecorhinus parvidens, Pl. XCT.

p. 859. |

Ambia novaguinensis, Pl. LXVIII. |

p. 546.

Ammothea virescens, Pl, LXI. p. 505.

Amphilius oxyrhinus, Pl. LXXIX.

p. 672.

Anthobosca albomaculata, Pl. LUXXX1.

p. 696.

anthracina, Pl, UXXXTII. p. 696.

australasie, Pl. LXXXTIT. p. 696.

— australis, Pls. LXXXII.,

LXXXIILI., p. 696.

clypeata, Pls. LXXXI,-LXXXTII.,

p. 696.

~—— crassicornis, Pl. LXXXT. p. 696.

erythronota, Pls. LXXXI.,

LXXXIL,, p. 696.

Fastuosa, Pl. LXXXT. p. 696.

-—— insularis, Pls. LXXXII,

LXXXIII., p. 696.

moderata, Pl, UXXXT. p. 696.

Apomatus timsit, Pl. LXXXVILI. p. 784.

Barbus argyrotenia, Pl, LXX1X. p. 672.

mimus, Pl, LXXIX. p. 672.

Bos sondaicus, Fig. 123, p. 903.

lowi, Figs. 124, 125, pp. 904,

905.

Brawnsomeria quadraticeps,

Pls, LXXXT., LXXXIL., p. 696.

Brevicella emarginata, Pl, LXVIII.

p. 546.

Calamachrous albipunctalis (vay.),

Pl. LXVIILI. p. 546.

Cariama cristata, Pl. LXIX. p. 546,

ATIONS.

| Ceratodus forsteri, Figs. 84, 85, pp. 610,

611

Cercaria ordinata, Fig. 107, p. 768.

i=)

| Cervus, Fig. 110, p. 778

-—— canadensis, Fig. 108, p. 775.

hanglu, Figs. 70, 108, pp. 566, 775.

—— macneilli, Fig. 71. p. 571.

-—— wallichii, Figs. 66-69, pp. 559,

561, 568, 565.

Chetostomus lepturus, Pl. LXXY.

p- 666.

— palmeri, Pl. LXXV. p. 666.

— paucispinis, Pl. LXXYV. p. 666,

Chitinopoma greenlandica, PI.

LXXXVIII. p. 784.

Corophium lacustre, Pl. LXXIV. p. 656.

Crucigera irrequaris, Pl. LXXXVII.

» isk

z zygophora, Pl. UXXXVILI. p. 784.

Dasyurotenia robusta, Figs. 92-101,

pp- 677-679, 681, 682, 684, 686, 688,

690, 691.

Dendronephthya disciformis, Pl. LXII1.

. 505.

Dicynoion laticeps, Pl. XCII, p. 859.

lutriceps, Pl. XCIL. p. 859.

psittacops, Pl. XCII. p. 859.

Diplostomum sirtale, Fig. 107, p. 768.

Dipteropeltis hirundo, Pl. LXXXIYV.

p- 763.

Eccasaurus priscus, Pl. XC. p. 859.

Elaphurus, Fig. 110, p. 778.

davidianus, Fig. 111, p. 779.

Elis combusta, Pl. LXXXII. p. 696.

(Mesa) alicie, Pls. LXXXL,

LXXXIII., p. 696,

INDEX OF ILLUSTRATIONS.

Elis (Mesa) dimidiaticornis, Pl. LXXXI.

p- 696.

—— (——) longiventris, Pls. LXXXI.,

LXXXTITL., p. 696.

(——) ruficeps, Pls. LUXXXII.,

LXXXIII., p. 696.

) tricolor, Pls.

LXXXIL,, p. 696.

Emydops minor, Pl. XCIII. p. 859.

LXXXL.,

Endothiodon platyceps, Pl. XCIII.

p- 859.

whattsi, Pl. XCITII. p. 859.

Entephria grisealis, P|, LXVIIL., p. 546.

mioswart, Pl, LXVIII. p. 546.

Equus burchelli, Figs. 102-106, pp. 758-

762.

Errina (Labiopora) aspera, Pl. XCV.

p- 876.

— ( ) capensis, Pl. XCV. p. 876.

—— ( ) nove zelandie, Pls. XCLV.—

XCVI., p. 876.

(Spinipora) echinata, Pl. XCY.

p- 876.

Galeops whaitst, Pl. XCI. p. 859.

Gammarus locusta, Pl. LX XIII. p. 656. |

zaddachi, Pls. LXXIII., LXXIV.,

p- 656.

Giraffa camelopardalis

Pl. LXXXVI. p. 771.

Glyphodes magnificalis, Pl. LXVIII.

p. 546,

nigricincta, P|. UXVIIT. p. 546.

— pfeiffere (var.), Pl. LXVIITI.

p- 546.

—— pseudocesalis, Pl. LX VIII. p. 546.

thornicrofti,

Hancockia eudactylota, Pl. LXXXV.

p. 770.

Ayracotenia hyracis,

pp. 594, 596, 597, 601.

Figs. 79-83,

LIctidognathus hemburyi, P\. XC1I, p. 859. |

Lctidopsis elegans, Pl. XCIIL. p. 859.

Inermicapsifer capensis, Figs. 72-78,

pp- 577, 580, 586, 588, 592.

Labeo percivali, Pl. LXXVILII. p. 672.

Lithophytum arboreum, P\. UXT. p. 505.

Macalia arruensis, Pl. LXVILII. p. 546.

- mioswart, Pl. LXVITT. p. 546.

— olivalis, Pl. LX VIII. p. 546.

Margarosticha plumbealis, P|. LX VIII.

p: 046.

Myzine abdominalis, Pls. LXXXIL.,

LXXXIII. p. 696.

—— braunsi, Pls. LXXXI., LXXXII,, |

p- 696.

constrictiventris, Pls. LXXXILI.,

LXXXIIL., p. 696.

Proc. Zoou. Soc.—1912, No. LXII.

XXV1l

Myzine stigma, Pls, UXXX1., LXXXI1.,

p- 696.

Nephthya bedfordi, Pls. LX11., LXII1.,

p. 505.

chabrolit, Pls. LXI., LXII., p. 505.

Noorda arfakensis, P\, UXVIIL. p. 546.

| Nythosaurus browni, Pl. XCIII. p. 859.

Odocoileus, Fig. 110, p. 778.

sp., Hig, 112, p. 782.

Omphisa variegata, Pl. UXVIII. p. 546.

Otocinclus maculipinnis, Pl. LXXV LL.

p- 666.

Oxyloricaria leightoni, Pl. LUXXVII.

p- 666.

—— tamane, Pl, LXXVII. p. 666.

Pagyda fumosa, Pl. LXVITILI., p. 546.

Perisyntrocha suffusa, Pl. LXVIII.

p. 546.

Phasianus formosanus, Pls. LX1V.—

LXVIL., p. 539; Fig. 126, p. 912.

x Phasianus versicolor,

Pls. LXIV—LXVII., p. 539;

Fig. 126, p. 912.

——— versicolor, Pls. LUXIV.-LXVIL.,

p- 589 ; Fig. 126, p. 912.

Pilocrocis angulifera, Pl. LXVIIT.

. 546.

Plecostomus honde, Pl. LXXVI. p. 666.

Pristerognathus platyrhinus, Pl. XCI.

p- 859.

Prodicynodon beaufortensis, Pl. XCIII.

p- 859.

Protula pacifica, Pl. LXXXVIL. p. 784.

Pyrausta flammealis, Pl. UXVAILI.

p. 546.

Rucervus duvaucelli, Fig. 109, p. 776.

Rusa, Fig. 110, p. 778.

aristotelis, Fig. 108, p. 775.

Scalpellum uccumulatum, Fig. 64,

p. 529.

—— arcuatum, Figs. 64, 65, pp. 529,

oo4.

hastatum, Fig. 64, p. 529.

simplex, Fig. 64, p. 529.

trilineatum, Fig. 64, p. 529.

Sceliodes grisealis, P\. LXVIII. p- 546.

Sclerophytum pinnulatum, Pls. LXIt.,

LXIILZ., p. 505.

Scymnognathus whaitsi, Pl. XC. p. 859.

aes columbiana, Pl. LUXXXVII.

84.

Sororts ambilateralis, Pl. LXXXVIII.

p. 784.

asperatus, Pl. LXXXIX. p. 784.

—— langerhansi, Pl. LXXXIX. p. 784.

medius, Pl. UXXXIX. p. 784.

XXVill

Spirorbis mérchi, Pl. LXXXIX. p. 784.

—— pusilloides, Pl. LXXXVIII.

p- 784.

racemosus, Pl, LXXXIX. p. 784.

spirillum, Pl. LXXXVIII. p. 784.

validus, Pl. LXXXIX. p. 784.

vitreus, Pl. LXXXVIII. p. 784.

Stereonephthya lutea, Pl. LXIII. p. 505.

Stericta rurealis, P|, LX VIII. p. 546.

Styphlodora naje, Fig. 122, p. 852.

Sylepta dinawa, Pl, LXVIII. p. 546.

Taurops macrodon, Pl. XC. p. 859.

Thecla bertha, P|. XCVII. p. 896.

campa, Pl. XCVILI. p. 896.

castrena, Pl. XOVILI. p. 896.

datitia, Pl. XCVII. p. 896.

esmeralda, Pl. XCVII. p. 896.

— fancia, Pl. XOVIT. p. 896.

jfernanda, Pl. XOVII. p. 896.

hamila, Pl. XCVII. p. 896.

japola, Pl. XCVITI. p. 896.

INDEX OF ILLUSTRATIONS.

Thecla molena, Pl. XCVII. p. 896.

nora, Pl. XCVIL. p. 896.

schausa, Pl. XCVII. p. 896.

sicrana, Pl. XOVII. p. 896.

Tilapia percivali, Pl. LXXIX. p. 672.

Tropidonotus natria, Pls. LXX.—LXXII.

p. 612; Figs. 86-91, pp. 616, 617,

619, 620, 627, 629.

Urocystidium gemmiparum, Figs. 113-

121, pp. 824, 826, 828, 833, 885, 841,

843, 845, 846.

Wrightella robusta, Pls, LXIT., LXTIL.,

p. 500.

Xenocara heterorhynchus, Pl. UXXVI.

p- 666.

multispinis, Pl. LUXXVI. p. 666.

Xenopharynx solus, Fig. 122, p. 852.

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

No. 11S:

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

October 29th, 1912.

Prof, HE, A. Mincutn, M.A., F.R.S., Vice-President, in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secretary read a Report on the Additions that had been

made to the Society’s Menagerie during the months May to

September, 1912.

Mrs. Rosr Haig Tuomas, F.Z.S., exhibited the eges of

Phasianus versicolor, P. formosus, and of the F, and F, offspring of

an experimental cross between a male P. versicolor and a female

P. formosus. She drew attention to the resemblance in size of

the eggs of the offspring and of the male parent species, whereas

the expectation was a likeness to those of P. formosus, thus

showing the descent through the male to his female offspring of

the small egg of his species.

My. R. Lypuxxer, F.R.S., F.Z.8., contributed a note pointing

out that, owing to an unfortunate mistake in labelling, the

Gazelle he described in the Society’s ‘ Proceedings’ for 1911 as a

new species, Gazella hayi, was really G. fuscifrons.

Mr. LyDEKKER also described the Bornean Bantin as a new

race to be distinguished from the typical Javan Bantin by the

* 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,

more upright direction and less outward curvature of the horns

of the male.

Mr. E. G. Bounencrer, F.Z.8., Curator of Reptiles, read a

paper dealing with the breeding-habits of the “ Millions” Fish

(Girardinus peciloides) from observations made in the Society’s

Gardens, and recorded cases of the male of this species breeding

before assuming the livery of its sex. The author drew attention

to parallel cases among fishes, in which, however, except in one

case, the question was one of degree only.

The Rev. T. R. R. Srespine, M.A., F.R.S., F.Z.8., read a paper

entitled “The Crustacea Isopoda of the ‘ Porcupine’ Hxpedi-

tion.” In the tribe Flabellifera, family Gnathiide, two new species

were assigned to the genus Gnathia, and a new genus was

instituted to receive a new species called Akidognathia cedipus.

In the tribe Asellota a new family, Thambematide, was proposed

for a new genus with a new species named Thambema amicorum.

Gnathia cristatipes, sp. n., owed its name to the striking processes

on the second and third joints of the second perzeopods, its head

in dorsal view being almost circular. Gnathia schistifrons, sp. n.,

had a head much wider than long, with a rounded notch in the

middle of the front, a peculiarity which it shared with G'. ceca,

Richardson, 1911, without sharing the blindness of that species.

Akidognathia edipus, gen. et sp. n., deserved its specific name alike

by the gouty appearance of the second perzeopod and the problems

which required solving in regard to its mouth-organs; while,

contrary to the custom of the family, the adult male appeared

to have at least one pair of maxilla. In the Asellota Thambema

amicorum, gen. et sp. n., seemed to be anomalous in the tribe by

having only four pairs of pleopods in the male and no uropods;

its narrowly cylindrical shape was also unusual.

This memoir will be published in the ‘Transactions’ in due

course.

Dr. F. E. Bepparp, M.A., F.R.S., F.Z.8., Prosector to the

Society, read a paper on the Anatomy and Systematic Arrange-

ment of the Cestoidea, in which he gave an account of six species

of Tapeworms from Reptiles, belonging to the genus Jchthyotenia

(s. 1.).

Mr. E. Duxinrretp Jones, F.Z.S., F.E.8., communicated a

short paper containing the descriptions of thirteen new species of

Butterflies of the genus Zhecla which he had collected at various

localities in §.E. Brazil.

LANTERN DEMONSTRATION.

At the conclusion of the Scientific Business, Sir Epmunp G.

Loprr, Bt., V.P.Z.S., showed a large number of slides, including

a fine series of coloured subjects, and a number of living speci-

mens, to demonstrate the capacity of the new electric lantern

which he had recently presented to the Society.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, November 12th, 1912, at half-past Hight

e’clock P.m., when the following communications will be made :—

H. R. Hoge, F.Z:8.

Some Falkland Island Spiders,

Bruce F. CuMMINGS.

On some Points in the Anatomy of the Mouth-parts of the

Mallophaga.

F. F. Laiptaw, M.A., I.Z.S8.

Contributions to a Study of the Dragonfly Fauna of Borneo.

—Part I. The Corduliine: the Genus Amphicnemis.

H. LysTER a M.A., D.Se ae _F.ZS. -- and WILLIAM

Nicors, MA, D.Sc, MD, PAS,

On some Pinas oi the ae Duck (Edemia nigra) and

their relation to the Pearl-inducing Trematode in the Hdible

Mussel (Mytilus edulis).

G. A. Boutencer, F.R.S., F.Z.8.

PSSST rts Vase isdbedacra anihes te orem cattle a

Descriptions of Three new Fishes discovered in the Gold

Coast by Dr. H. G. F. Spurrell, M.A., F.Z.8.

The following papers have been received :—

G. W. Sits, M.A,, and E. H. J. SCHUSTER, M.A., D Sce., F.Z.S8.

“The Genus Engeus, or the Land Gama of eae

The Hon. Paut A. Mrruven, F.Z.S.

Description of an Amphipod belonging to the Family

Talitride from the Woodbush, Transvaal.

Dr. F. D. WEtcH.

Observations on living Specimens of the Capped Langur

(Semnopithecus pileatus).

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Socrery or Lonpon,

RecEnNt1’s Park, Lonpon, N.W.

November 5th, 1912.

No. 118.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON,*

November 12th, 1912.

Dr, A. Smita Woopwaprp, F.R.S., Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

The SzcrETARY read a report on the Additions that had been

made to the Society’s Menagerie during the month of October,

1912.

The Secretary exhibited a photograph of the Hainan Gibbon

(Hylobates hainanus) in natural colours, taken by Messrs. Elhott

& Fry, by the Lumiére process.

Mr. G. A. Boutencer, F.R.S., F.Z.8., exhibited a specimen,

155 mm. long, of the African Cichlid Perch (Hemichromis bimacu-

latus Gill), which had recently died in Capt. Vipan’s aquarium.

The specimen was remarkable for its large size. Although the

species is common over a considerable part of Africa, and hundreds

of specimens have been collected in various localities, no wild

example is known to exceed a length of 100 mm.

This Hemichromis is tinged with bright red about the head

and on the fins, and, as was noticed in the ‘ Fishes of the Nile,’

p. 462, this red is soluble in spirit, which, a few hours after the

immersion of the fish, acquires a bright orange-red colour.

Mr. Epwarp GERRARD exhibited the skull of an Indian

Rhinoceros (thinoceros wnicornis) which had recently died in

the Society’s Gardens. He pointed out that abscesses had been

formed at the base of the lower incisors, which had been much

* 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 Stic Shillings per annum, payable in advance.

rubbed down so that the nerves had become exposed. The

animal must have suffered great pain, and this no doubt was the.

cause of his dashing his head against the walls and bars of his

den in the violent manner he at times did.

Mr. E. G. Boutencsr, F.Z.8., Curator of Reptiles, exhibited a

living specimen of the Salamander, Amblystoma tigrinuwm, which

he had obtained from an Axolotl placed in special conditions

with the object of forcing the metamorphosis.;

Mr. D. Seru-Suiru, F.Z.8., Curator of Birds, gave a lantern

demonstration of photographs of the young of Hemprich’s Gull

(Larus hempricht), the Black-necked Swan (Cygnus melano-

coryphus), and the White Stork (Ciconia alba) which had been

hatched in the Society’s Gardens during the present year. He

also showed slides of the nest, in a tree near the Apes’ House,

made by the Orang-utan which escaped from its cage on

November 8rd, 1912.

Mr. H. R. Hoee, M.A., F.Z.8., read a paper entitled ‘‘ Some

Falkland Island Spiders,” based on a small collection of Spiders

formed by Mr. Rupert Vallentin during a two years’ stay in the

Falkland Islands.

Of some of the species there were a fair number of specimens,

but they comprised only six species of Spiders and one of the

allied suborder Opilio.

The species were all apparently new, but the genera were all

to be found either in Patagonia, Tierra del Fuego, or the islands

about Cape Horn. The ancestors of the Spiders might all have

been transported aerially at an early period and therefore afforded

no evidence for or against a former land-connection, but in the

event of the latter there should be many more species. The

Opilio might have been conveyed under the bark of floating trees.

Mr. G. A. Boutencsr, F.R.S., F.Z.S., read a short paper con-

taining the descriptions of three new Fishes which had been

discovered by Dr. Spurrell in the vicinity of Bibianaha, near

Dunkwa, Gold Coast, and presented by him to the British

Museum.

A paper was communicated by Dr. H. Lyster Jamuson, M.A.,

Ph.D., F.Z.S., and Dr. Wittiam Nicott, M.A., D.Sc., F.Z.S.,

which contained an account of some parasites of the Scoter Duck

(demia nigra), and discussed their relation to the pearl-inducing

Trematode in the Edible Mussel (J/yitilus edulis).

A paper was received from Mr. F. F. Larpiaw, M.A., F.ZS.,

dealing with some Dragonflies from Borneo belonging to the

subfamily Corduliine, and to the genera Disparonewra and

Amphicnemis of the subfamily Agrionin, with an account of a

’ number of new species.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, November 26th, 1912, at half-past Hight

o'clock P.m., when the following communications will be made :—

G. W. Sura, M.A., and E. H. J. SCHUSTER, MA. D Sey E.ZS.

The Genus Engeus, or the Crayfishes of Australia.

E. 8. Goopricn, M.A., F.R.S., F.Z.S.

On the Structure of Bone in Fishes: a Contribution to

Paleohistology.

C. L. Bovtencrr, M.A., D.Se., F.Z.S.

__ Report on the Myzostomida collected by Mr. Cyril Crossland

in the Red Sea in 1905.

The Hon. Paut A. Meruuven, F.Z.S.

Description of an Amphipod belonging to the Family

Talitridz, from the Woodbush, Transvaal.

Bruce F. Cummines.

Pesta Dib create

On some Points in the Anatomy of the Mouth-parts of the

Mallophaga.

The following papers have been received :—

G. P. FARRAN.

Plankton from Christmas Island, Indian Ocean.—II, On

Copepoda of the Genera Oithona and Paroithona.,

F. KE, Bepparp, M.A., D.S8c., F.R.S., F.Z.8.

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.—VIIT. On some Species of Jchthyotenia and

Ophidotenia from Ophidia.

H. L. Hawxins, M.Sc., F.G.S8.

The Anterior Ambulacrum of /chinocardium cordatum Penn.,

and the Origin of Compound Plates in Echinoids.

H. B. Preston, F.Z.S.

Diagnoses of new Species and Varieties of Agnathous

Mollusca.

R. Lyprexxer, F.R.S., F.Z.S.

The Dwarf Buffalo of Southern Nigeria: with a Revision of 3

the Dwarf Buftaloes of Western Africa.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,

Recent’s Park, Lonpon, N.W.

November 19th, 1912.

Papers (continued).

44, A Wats. on the rare British Nudibranch Hancockia eae Gosse. . By Sir Cirarius

Eniot, K.C.M.G., O.B., F.Z.S. (Pl. LXXXV:) . Sein shines jo enststort croteiereel ener cels 770

45, The North Rhodesian Giraffe. By R. LyexKER, E.R.S., F.Z.S. (Pl. UXXXVIL) .. 7/1

46, On Antler-Growth in the Cervide, with special reference to Hlaphurus and Odocoileus

(Dorcelaphus). By R. I. Pocock, F.R.S., F.LS., F.ZS., Superintendent of the

Gardens and Curator of Mammals, (Text-figs. 108-112.) ........+e seer er seeeee 173

47. Polycheta from the Pacific Coast of North America.—Part T, Serruips, with a

Revised Table of Classification of the Genus Spirorbis. By Huumn L, M. Prxext,

B.Sc., F.Z.S., Demonstrator of Zoology and Reid Fellow, Bedford College, University

of London. (Pls, LXXXVII-LXXXIX.) ......-- cece cece ee ee neces Weise, are oe 784

48, The One-sided Reduction of the Ovaries and Oviducts in the Amniota, with Remarks

on Mammalian Evolution. By Hans Gavow, M.A., Ph.D., F.R.S., F.Z8. ..-...-- 808

49, Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.— VI. On

an Asexual Tapeworm from the Rodent, Fiber zibethicus, showing a new form of

Asexual Propagation, and on the supposed Sexual Form. By Frank EK. Bepparp,

M.A., D.Sc., F.R.S., F.Z.8., Prosector to the Society. (Text-figs. WB SIP SeReace yiode 2

50. On Two new Trematode Parasites from the Indian Cobra, By Wivutam Nicott,

M.A., D.8c., M.D., F.Z.S8., Lister Institute of Preventive ae London. ee

HR ML s)he Wiel vis arses Ste elon heise elem Cale gt alatanteLas PE ARO UID DD apie SABO Dor NoEt OF 851

51. Statistical Note on the Worm Parasites collected from the Animals dying in the

Zoological Gardens, from December 1910 till April 1912. By Wrpriam Nrcoxt,

M.A., D.Se., M.D., F.Z.S., Lister Institute of Preyentive Medicine, London ........ 856

52, On some new Fossil Reptiles from the Permian and Triassic Beds of South Africa.

By R. Broom, D.Se., O.M.Z.S. (Pls. XC—XCIUM.) .. cc ee eee e cere eee cree cees 859

53. Ou the Hydrocoralline Genus, Hrrina. By Professor S. J. Hickson, F.BS., F.Z.8.,

The University of Manchester. (Pls. XCIV.—XCVI.) .-.----+ esses eee cece seeeee 876

54, Descriptions of new Butterflies of the Genus Thecla from §.H. Brazil. By E. Duxin-

FIELD Jonns, F.Z.S., F.E.S. il. KOWAL Reap a ocrs Seis ane eal ree ela eee eee . 896

55, The Bornean Bantin. By R. Lypexxnr, E.R.S. cd ZS. (Text-figs. 123-125.) ...... 902

56. Notes on the Breeding of the “ Millions” Fish (Girardinus peciloides), By Epwarp

G. Bourmncer, F.Z8., Curator of Reptiles ......0:0+ sees seen ence ete rene ee cere 906

Titlepage ...s.-ceeeeeeee Re agin Ris ate, cine featereyey aaahere teeta e eee e ener ee ee ec snes cees i

List of Council and Officers ......++--.20e eee A eee eI re REIT PICK marker POR ; il

Print of Contenter. oiic oa dena weein wore a cinta e's cies iwtaleie(e om mimi mInjo/Mcin:lainie)o\ Semin Winam = mieelnie = Sea

New Generic Terms PEGS UENCE aad onli i Dae mE, SR reac Maes teense eunoien Xvi

Index of Scientific Names ..-1-- seers eee eeeeee rece eter ie nena teen ee sere tances aa OXVie

Index of Illustrations ........ caihei eh che ats wie sialon sist’ e wieipinieyn airs iatavenee Gterpiowere tort ess <xyieg

LIST OF PLATES.

1912, Parr IV. (pp. 757-913).

Plate Page

LXXXIV. Dipteropeltis hirundo .. “ i pice: gps “aaa 763

LXXXV. Hancockia eudactylota Mie as th ge eee

LXXXVI. Giraffa pease thornicr ofti ee nee yon ieee

LXXXVIL. es Be

LXXXVIII. North American Serpulide ........... cat No ore rt aae

LXXXIx. } ; 2

XC.) er

pa | South African Fossil Reptiles ..5...50...ssecesreecs 859

XCIL. | . se,

-XCIIL.) |

XOIV.

XCV, }Hydroids of the Genus Erring ......eccececeseee sees 876.

XCVI. : Peak |

XCVII. New Species of Thecla from 8.E. Brazil ............. .. 896

NOTICE.

The ‘ Proceedings * for the year are issued in four parts, paged consecutively,

so that the complete reference is now P. Z. 8.1912, p.... The Distribution

is as follows:— :

yA dd Part JI. issued in March.

etm ae et June.

eae OD September,

BS Vict, hn ce eee Dee. aa

« Proceedings,’ 1912, Part III. (pp. 505-756), were published on

August 29th, 1912,

1a 4 w

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