1 Dipel Broied h

OF THE

COUNCIL

AND OFFICERS

OF THE

ZOOLOGICAL

SOCIETY OF

LONDON.

1914.

Patron.

His Masesry Tur Kina.

COUNCIL.

His Grace Tae Duke or Beprorp, K.G.,

RicHarD H. Burne, Hsq.,M.A.,

Vice-President. .

Atrrep H. Cocks, Esq., M.A.

THe Rr. Hon. toe Hart or

Crommry / UP.C, > 1GiCiBe

GC NGS KEG Sake Enis.

Vice- Pee

F. G. Dawrrey Drewirt, Esq.,

M.A., M.D.

CuarLtes DrumMonn, Esq.,

Treasurer.

F. Du Cant Gopman, Esq.,

Calis. HRS:

Sir Waurer Roper LAWRENCE,

Bt., G.C.L.E., Vice-President.

Hrnest W. MacBriper, Esq.,

Wis, IDESies5 TRS, 1002

President.

, F.R.S., President.

E. G. B. Mrapn-W Apo, Esa.

Prof. Epwarp A. MINcHIN,

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

12. Cennumes Mircuenn, HsqQ.,

M.A., D:Se., ED) Rakes

Secretary.

W. R. Octrvie-Grant, Esq.

ALBERT Pam, Esq.

Aprian D. W. Poutock, Esq.

Sir Ronatp Ross, K.C.B.,

BER Se

THE Margurss oF SLIGO,

F.S.A.

OLpDFIELD THomas, Hsq., F.R.S.

AntHony H. WHINGFIELD,

Ksaq.

Henry Woopwarp, Esq., LL.D.,

F.R.S., Vice-President.

PRINCIPAL OFFICERS.

P. Cuatmers Mircnrny, M.A., D.Sce.,

Secretary.

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

KDE, IRS,

., Prosector.

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

Resident Superintendent of the Gardens.

D. Sern-Smivn, Curator of birds and Inspector of Works.

Epwarp G. BouLencer, Curator of Reptiles.

Prof. H. Maxweint Lerroy, Curator of Insects.

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

Henry G. J. Pravor, Tnbr Ue tan and Clerk of Publica ions.

JoHN Barrow, Accountant.

W. H. Coxz, C} hief Clerk.

LIST OF CONTENTS.

1914, pp. 1-490.

EXHIBITIONS AND NOTICES.

Page

The Secrerary. Report on Additions to the Society’s

Menagerie during the months of November and

Weenie whol See cece dt « ays yh take SRM te acts eta hore ice 217

Mr. D. Sera-Smuirs, F.Z.S., Curator of Birds. Exhibition

of photographs and skin of Hybrid Birds ... .........-- 218

Mr. G. A. Bounencrr, F.R.S.. F.Z.S. Notice of Report on

Batrachians and Reptiles from Dutch New Guinea ... 218

The Secrerary. Report on Additions to the Society's

Menagerie during the month of January 1914 ......... 219

Mr. Grorce Jenison. Exhibition of a mounted specimen

of a Hybrid Sea-Lion

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

bition of photograph of the Giant Saddle-backed

Tortoise 220

Dr. R. T. Leer, M.B., F.Z4.5., and Surgeon H.

L. Atkinson, R.N. Lantern demonstration of the

Helminthes collected by the British Antarctic (‘Terra

Nova’) Expedition, 1910-1913 222

Page

Mr. C. Tare Recan, M.A., F.Z.S. Notice of Report on

Freshwater Fishes from Dutch New Guinea ............ 226

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

Menagerie during the month of Kebruary O04 27. - eee 475

_G. C. Rosson, B.A. Notice of Report on Mollusca from

IDYonro MUINVEN ye CC MUU AVeEE Sea ceMennencpnusracencsdococcdoadodaace 475

. K. G. Buatr, B.Sc. Notice of Report on Heteromerous

Coleoptera from Dutch New Guinea: 12.2 ee ee 476

- R. H. Burne, M.A., F.Z.8. Palatal growth in mouth

one (Oenonvell, (Uleseneiiernits) 15) oaancceocosoqdaaadudsoocatess00. 476

_R. J. Pocock, F.R.S., F.Z.S. On the Feet of Domestic

Woes.) )(Mlexct tigress 13.) ec ieee eee eee 478

The Secrerary. Exhibition of the photograph of a female

Orang-utan (Sumiasatyrus) ........c0cccucueeeseeseseneuceeens 485

The Secrerary. Exhibition of photographs of large-tailed

una DWourestictS hee piers sere: eee eee reer 485

. W. T. Catan, F.Z.8. Notice of Report on River-Crabs

(Potamonidee) from Dutch New Guméalecees eee 485

:, OLDFIELD THomas, F.R.S8., F.Z.S. Notice of Report on

Mammals from Dutch New Guinea........................ 485

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

Menagerie during the month OL Miarch those eee 486

The Szcrerary. Notice of change in time of Scientific

Meetings «0 idcucestes aie sstiene cided eee ence Oe ante eee eee 486

Mr. D. Seru-Surra, F.Z.S. Exhibition of an Egg of Mantell’s

Kiwi (Apteryx mantel) laid in the Society's Gardens. 487

Mr. Srantey Hirst, F.Z.S. Notice of Report on Arachnida

(other than Spiders) and Myriopoda from Dutch New

(Givaharee, \eAaneadooos WE eR NEL NnBe ante osoridcbs. .Scacc5- 487

Surgeon G. Murray Levick, R.N. Notice of Lecture on

the Manners and Customs of Adélie Penguins (Pygos-

CEILS OM CLLOL) oo tintcleslea a oe oz sa sels 04 cles e> «gee ee 487

Page

Sir Epmunp G. Lopsr, Bt., F.Z.S. Exhibition of Antlers

of Red Deer (Cervus elaphus) CER ORE OLN ALAIN: TORINO) INE 9 0 488

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

Menagerie during the inonth of April 1914 ............ 489

Mrs. R. Hate Tuomas, F.Z.S. Exhibition oF Skulls of Horn-

les Amatclomestiene eee meee tat er see Sh nate) G, 490

Mr. D. M.S. Wavson, M.Se., F.Z.8. Exhibition of Pro-

colophon trigoniceps, a Cotylosaurian Reptile ............ 490

PAPERS.

1. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—XII. Further Observations

upon the Genus Urocystidium Beddard. By Frank KE.

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

Societys: ((lext-foumesy I O)\e heap came pact a teemtes. IL

2. Observations made to ascertain whether any Relation

subsists between the Seasonal Assumption of the

“ Helipse” Plumage in the Mallard (Anas boscas) and

the Functions of the Testicle. By C. G. Sunremann,

F.Z.S., and 8. G. Suarrock. (Text-figures 1-6.) ...... 23

3. The Possible Connection between Spindle-Length and

Cell-Volume. By C. F. U. Munx, M.Sc., F.L.S., F.Z.8.

(IBA Rev ets a1 LES) SS ia Ae een cit a ae eS 45

4, Contributions to a Study of the Dragonfly Fauna of

Borneo.—Part II. The Gomphine and Chlorogom-

phine. By F. F. Larpnaw, M.A., F.Z.8. (Plate I.) . 51

5. Note on an imperfectly developed specimen of the Sea-

Urchin (Zchinus esculentus). By H. C. CHapwicr,

Agnes (hext-neunest U4)! eek Saks eee. 65

6. Spiders from the Montebello Islands. By H. R. Hoec,

Mien Ze over (Ce labese le UND cs RSI Scala ety ees 69

10.

ily

14.

15.

. On the Nests of Pseudoscorpiones: with Historical

Notes on the Spinning-Organs and Observations on

the Building and Spinning of the Nests. By H.

WWidibIS IKnW, IRSA keene boise. ccasteaesemee er. see

. The Structure and Life-History of a Tapeworm (Ichthyo-

toenia filicollis Rud.) Parasitic in the Stickleback. By

F. J. Mecerrr, M.Se. (Birm.), Board of Agriculture

and Fisheries Research Scholar, University of Bir-

mingham. (Plates I.—IV., and Text-figures 1-5.)......

. Trematode Parasites from Animals dying in the Zoo-

logical Society’s Gardens during 1911-1912. By

Wittiam Nicotz, M.A., D.Sc., M.D., F.Z.8. (Plates

1 I) ee ne Pe PBN REA MGSO mae ia cacgaoooa0 4003

On the Skull of a Pariasaurian Reptile, and on the

Relationship of that Type. By D. M. 8. Warson,

M.S8c., F.Z.8., Lecturer on Vertebrate Paleontology im

University College, London. (Text-figures 1-7.)......

Report on the Deaths which occurred in the Zoological

Gardens during 1913, together with a List of the

Blood-Parasites found during the Year. By H. G.

Pummer, F.R.8., F.Z.S., Pathologist to the Society ...

. Some Phases in the Reproductive History of the Female

Mole (Yalpa europea). By Freprric Woop-Jonus,

D.Se., F.Z.8. (Plates I-II1., and Text-figures 1-13.).

. On the Fossil Carnivores Cynodictis intermedius and

Cynodow gracilis from the Phosphorites of Quercy.

By Azperriva Cartsson, Zootomical Institute, Uni-

versity of Stockholm. (Plate I.) ya)2 3-5 eee

On the Genera Ceratocephale Malmgren and Tylorhynchus

Grube. By L. N. G. Ramsay, M.A., B.Sc., Carnegie

Research Scholar, Christ’s College, Cambridge

wer ore cees

On the Annelids of the Family Nereide collected by

Mr F. A. Potts in the N.E. Pacific, in 1911. Witha

Note on the Morphology of Micronereis as a Repre-

sentative of the Ancestral Type of the Nereide. By

L. N. G. Ramsay, M.A., B.Sc., Carnegie Research

Scholar, Christ’s College, Cambridge. (‘Text-figures 1-7.)

Page

113

139

155

18]

191

227

231

237

16.

IS)

23.

. Fauna of Western Australia.

vil

The Structure and Development of the Caudal Skeleton

of the Teleostean Fish, Pleuwragramma antarcticum.

Byean eNwynnn homnon. (Plates ty de IM ye i... 2.

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—XIIT. On Two new Species

belonging to the Genera Oochoristica and Linstowia,

with Remarks upon those Genera. By Frank E.

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

Eneasoctety.) 0 (Mextanounesil= 8. Via cteuyas tse sselc.t os.

The Malay Race of the Indian Elephant, Hlephas mami-

mus hirsutus. By R. Lypexxer, F.RS., F.ZS.

(Me sstetn comme les): een aecem UNE Mle, at ace ON a

Fauna of Western Australia.—I. The Onychophora of

Western Australia. By W. J. Daxin, D.Sc., F.LS.,

F.Z.S., Professor of Biology, University of Western

Mustealianren( Vext- Hewes eee. 5. anaes.

II. The Phyllopoda of

Western Australia. By W. J. Daxin, D.Sce., F.LS.,

F.Z.S., Professor of Biology, University of Western

ivustrabice, @eslabes liad Tee vemere 0 ai: ee NOS oe

. Notes on a Collection of East Afvican Mammals pre-

sented to the British Museum by Mr. G. P. Cosens.

FS ya Ga Uipvars18) UA NWAING Poe Steere ators) oars ela slain fe satiate Salant

2. On the Nature of the Lateral Muscle in Teleostei. By

Epwarb W. SHann, B.Sc., Assistant to the Professor

of Natural History and Lecturer in Comparative

Embryology in the University of St. Andrews. (Text-

AM OMIM E Sill OS) Mrsttacya te are iee aeecanmeee eae oman ene ARR

Note on Aristeus goldiet Macleay, and on some other

Fishes from New Guinea. By C. Tarn Recan, M.A.

(Gtlesa iat teringeis 1s, 45) Be Beno ce aan R Opa so aa eo Rene eer nor eee

4. Crustacea from the Falkland Islands collected by Mr.

Rupert Vallentin, F.L.S.—Part II. By the Rev.

Tuomas R. R. Sreppine, M.A., F.R.S., F.L.S., F.Z.S.

(eden resal PCa Oa eaatian Gn ote aCe aeC Itc H CAce pe eaeene eee Soy.

. Further Contributions to the Anatomy of the Ophidia.

By Joseru C. Tompson, Surgeon, U.S. Navy ......... 5

263

289

293

307

319

339

341

26.

29:

30.

31.

Alphabetical List of Contributors

vill

Page

The Coloration of the African Hunting Dog (Lycaon

pictus). By Major J. Srevenson-Hamitron, C.M.Z.S. 403

. On a New Cestode from an Albatross, Diomedea irrorata.

By H. A. Bayuis, B.A. -(Vext-figures 1-4.) .......53: 407

. On a Remarkable Case of Affinity between Animals in-

habiting Guiana, W. Africa, and the Malay Archipelago.

By OnpriEetp Unomas, oh IRS.. EZIS, en, eee ennenee 415

On Two New Species of Tapeworms from the Stomach

and Small Intestine of a Wallaby, Lagorchestes con-

spicillatus, from Hermite Island, Monte Bello Islands.

By R. C. Lewis, M.A.(Cape), (1851 Exhibition Scholar).

(Plates =X. and Vext-figures 1-32) \jcsnesesaneeeeaees 419

Notes on the Circulatory System of Hlasmobranchs.—I.

The Venous System of the Dogfish (Scylliwm canicula).

By Cuas. H. O’Donoeute, D.Sc., F.Z.8., Senior Assis-

tant in the Zoological Department, University College,

London. (Plates I., II., and Text-figures 1-4.)......... 435

Notes on Colour Development in the Indian Wood-Stork.

By GrorcE Jennison, Belle Vue Gardens, Manchester. 457

. Scent Organs in Trichoptera. By Brucz F. Cummrnes,

British Museum (Natural History). (Text-figures 1-8.) 459

a0 (=I Apa ee Ey Ce I EL ah gon XV1l

AOE HABE Le At i tse

OF THE

CONTRIBUTORS,

With References to the several Articles contributed by each.

(1914, pp. 1-490.)

Page

Arkinson, Surgeon E. L., R.N., and Lerrer, Roperr T., ‘

D.Sc., M.B., F.Z.S.

Lantern demonstration of the Helminthes collected by

the British Antarctic (‘Terra Nova’) Expedition, 1910-

TLS eit ae Br etn epee SORA Ut eC ed ee Baan 222

Bayuis, H. A., B.A.

On a New Cestode from an Albatross, Diomedea irrorata.

(estol cummestl “4a eiteticlathet nites Se Reh <5 GAL ion Min oats 407

Bepparp, Fran E., M.A., DSc., F.R.S., F.Z.8., Prosector

to the Society.

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.— XII. Further Observations upon

the Genus Urocystidiwm Beddard. (Text-figures 1-9.) ... 1

XIII. On Two new Species belonging to the Genera

Oochoristica and Linstowia, with Remarks upon those

Poneto hex i auresi|( 8. ants riscsacssilestst ea stiasis' Je ica' 263

Page

Brair, K. G., B:Se.

Notice of Report on Heteromerous Coleoptera from

MubehsNew. Gaines) ova cce lone eee eee Gee ete eee eee 476

BouLEeNncER, Epwarp G., F.Z.S., Curator of Reptiles.

Exhibition of photograph of the Giant Saddle-backed

WNGIARONISE.R sa npaasncgded sebunanacdaeneendonaacnesas mood desc essa so08 sossc0 220

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

Notice of Report on Batrachians and Reptiles from

Duteh New Guineas. s.030 sachin ect aes een eee 218

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

Exhibition of palatal growth in mouth of Camel.

(Text-figure sd yc cecihs. as asueteckdemne casts: eee ee eee ee eee 476

Catman, WivurAm T’., D.Sc., F.Z.S.

Notice of Report on River-Crabs (Potamonidz) from

Duteh New Guinea)... {05.0.5 ieee eee eer eee eee 485

CarLsson, ALBERTINA.

On the Fossil Carnivores Cynodictis intermedius and

Cynodon gracilis from the Phosphorites of Quercy.

GBla ter.) rs tadecs te cxtvae cee sniinek Mate tnaes eee eee ae ee 22:7

CHapwick, H. C., A.L.8.

Note on an imperfectly developed specimen of the

Sea-Urchin (Hchinus esculentus). (Text-figures 1-4.) ... 65

Cummines, Bruce F.

Scent Organs in Trichoptera. (Text-figures 1-8.) ...... 459

Page

Dakin, Prof, Winu1AM J., D.Sc., F.L.S., F.Z.8.

Fauna of Western Australia.—I. The Onychophora of

Westerm Australias (Text-figure 1.) 0.2.0... 00sec tec esce eee 289

If. The Phyllopoda of Western Australia. (Plates

15 dhs, ALTE Rs ie era ei ce etl 8 ACen nH a Soe oe ae 293

. Dottman, Guy.

Notes on a Collection of Hast African Mammals pre-

sented to the British Museum by Mr. G. P. Cosens ...... 307

Hirst, Stanuey, F.Z.8.

Notice of Report on Arachnida (other than Spiders)

and Myriopoda from Dutch New Guinea ..................... A87

Hoee, Huyry R., M.A., F.Z.8.

Spiders from the Montebello Islands. (Plates I.-IIT.). 69

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

TON, J.

JENNISON, GEORGE.

Exhibition of a mounted specimen of a Hybrid Sea-

OGG WIN Re i BER as 1 Dear eat Ne at ot SONG Neues Ae ir a ACU ea 219

Notes on Colour Development in the Indian Wood-

RSHUOVELS oa glnatere) sees een ate wal Re ica lle ea RR ORR Pg PE 457

Jones, F. Woop-. See Woov-Jonzs, F.

Kew, H. Wautts, F.Z.8.

On the Nests of Pseudoscorpiones: with Historical

Notes on the Spinning-Organs and Observations on the

Building and Spinning of the Nests ...0.....0.....ssseteescees 93

Xil

Page

Larpiaw, Frank F., M.A., F.Z.8.

Contributions to a Study of the Dragonfly Fauna of

Borneo.—Part Il. The Gomphine and Chlorogomphine.

(QElabe adi) Freie ais lace stg ae ae tients ok ial ed oe Se eee ol

Lerer, Rospert T., D.Se., M.B., F.Z.S., and ArTKrnson,

Surgeon H. L., R.N.

Lantern demonstration of the Helminthes collected by

the British Antarctic (‘Terra Nova’) Expedition 1910—

Sd oe ea ane ee MRE ES PeeeMeE och Hor ah <ohhtso Sha dincsacaotona 222

Levicxn, Surgeon G. Murray, R.N.

Notice of lecture on the Manners and Customs of

Adeheseene mins) Gsygoscclis) aolclic) lene aye neers eer nee 487

Lewis, R. C., M.A. (Cape).

On Two New Species of Tapeworms from the Stomach

and Small Intestine of a Wallaby, Lagorchestes conspi-

cillatus, from Hermite Island, Monte Bello Islands.

(Plates 1 —X.; and. Vext-fisures 1-31)... -pa.-peeeoee eee ee ee 419

Lover, Sir Epmunp G., Bt., F.Z.8.

Exhibition of Antlers of Red Deer (Cervus elaphus) ... 488

LypEKKER, RicHarp, F.R.S., F.Z.S.

The Malay Race of the Indian Elephant, Hlephas

maximus hirsutus. (Text-figures 1-3.)............-2.e0eeeee 285

Merx, Cuartes F. U., M.Sc., F.L.S., F.Z.8.

The Possible Connection between Spindle-Length and

Cell-Volume. (Plates I. & LY) sv cnpusesuenecs acetic ee eee eee 45

Mecerrt, F. J., M.Sc. (Birm.).

The Structure and Life- History of a Tapeworm (chthyo-

tenia filicollis Rud.) Parasitic in the Stickleback. (Plates

Tay, .and Vextsfisures 1225.) iaiken aes. Aceh heen ee eae 113

xiil

Page

MrrcHe., P. Caaumers, M.A., D.Sc., LL.D., F.R.S., F.Z.S.,

Secretary to the Society.

Report on Additions to the Society's Menagerie during

the months of November and December 1913 ............... 217

Report on Additions to the Society’s Menagerie during

EMER MOU Ole Jannat al OM Ava see tab came se Gens. oC aN Stes sty 219

Report on Additions to the Society’s Menagerie during

Eermonbh, afew bruary, 1 OA sce. accem a dmceeey ua sees ce dita AT5

Exhibition of the photograph of a female Orang-utan

( SONU, SULIT) ees Rn ASB SGaA Ce a Soe SARC EAE SONS HE HMR RAN 485

Exhibition of photographs of Large-tailed Punjab

IDXOUTNG@SAUS: TSUNEO dried sheer mts aaa Pine creat Seen Hiay a MAN BO 485

Report on Additions to the Society’s Menagerie during

Eieentoutinot, Marche MOVE Gi: ...mracen ce ee suaatione cateoes: levees 486

Notice of change in time of Scientific Meetings ane 486

Report on Additions to the Society’s Menagerie during

Eheamiomt note Acorn acl ONE seb Seidel enero aula det none mae Ne 489

Nicont, Winu1am, M.A., D.Sc., M.D., F.Z.8.

Trematode Parasites from Animals dying in the Zoolo-

gical Society's Gardens during 1911-1912. (Plates

IT EN OS) Hiercties eat eer a ta em aT Soa) EOE te sis)

O’DonoGHUE, CHARLES H., D.Sc., F.Z.S.

Notes on the Circulatory System of Hlasmobranchs.—

I. The Venous System of the Dog-fish (Seylliwm canicula).

Scie eM eee at ewes 435

(Plates I., IT., and Text-figures 1-4.)

Primmer, Henry G., F.R.S., F.Z.8., Pathologist to the

Society.

Report on the Deaths which occurred in the Zoological

Gardens during 1913, together with a List of the Blood-

BAcASIGes LOUMG CUMIN LAG) MOUE actual dog iiatens Ws, kept ds eevee 181

X1V

Pococs, Recinatp I., F.R.S., F.LS., F.Z.8., Curator of

Mammals.

On the Feet of Domestic Dogs. (Text-figures 1-3.) ... 478

RAsAw. iy NG MA RB Se,

On the Genera Ceratocephale Malmgren and T'ylo-

PIOUNUG UNIS (Grealorey: | Soy span saaodAguapedsscs¢cocuotoocdgoqesessonarnonen 231

On the Annelids of the Family Nereide collected by

Mr. F. A. Potts in the N.E. Pacific, i 1911. With a

Note on the Morphology of MWicronereis as a Representa-

tive of the Ancestral Type of the Nereida. (Text-figures

JE (A) src deb en ation Sesaaetaaneeeatron ag i2viso9c0d jakdacks opcawsoaznbe a0: 237

Reean, C. Tarn, M.A., F.Z.8.

Notice of Report on Freshwater Fishes from Dutch

New Guinea ..... gaa ps UI RAD A ka 226

Note on Aristews goldiei Macleay, and on some other

Fishes from New Guinea. (Text-figures 1 & 2) ........:... 339

Rosson, G, C., B.A.

Notice of Report on Mollusca from Dutch New Guinea. 475

SELIGMANN, CHARLES G., I'.Z.S., and SHarrock, 8. G.

Observations made to ascertain whether any Relation

subsists between the Seasonal Assumption of the ‘‘ Kelipse”

Plumage in the Mallard (Anas boscas) and the Functions

onthe Testicle \(WMext-teures 16) )oeieay na. - eee ee eee 23

Sura-Surru, Davin, F.Z.8., Curator of Birds.

Exhibition of photographs and skin of Hybrid Birds ... 218

Exhibition of an Egg of Mantell’s Kiwi (Apterya

Page

SHANN, Epwarp W., B.Sc.

On the Nature of the Lateral Muscle ‘in Teleostei.

(Gextstiomres 13.) es sia RUS Is OA 0 eA A 319

Suarrock, S. G., and Setiamann, Caarues G., F.Z.5.

Observations made to ascertain whether any Relation

subsists between the Seasonal Assumption of the “‘ Kclipse”

Plumage in the Mallard (Anas boscas) and the Functions

omunelesticles® (lext-tounest 1 -O5)ie pans eet cas bene ene 23

Surry, D. Sera-. See Sera-Surirsz, D.

Sreppine, The Rev. THomas R. R., M.A., F.R.S., F.LS.,

TF ask

Crustacea from the Falkland Islands, collected by

My. Rupert Vallentin, F.L.S.—Part II. (Plates I.-IX.). 341

Stevenson-Hamitton, Major J., C.M.Z.S.

The Coloration of the African Hunting Dog (Lycaon

ADOC. “Bice Dosen colon HOE PA OR OE DEAR nD Ont e eric acon” Sy ccan CAE 403

THomAs, OLDFIELD, F.R.S., F.Z.8.

On a Remarkable Case of Affinity between Animals

inhabiting Guiana, W. Africa, and the Malay Archi-

TWIG eokuteossnaccnocchiokeoten sHbO GbE Roses aS ASEE rapa co neEedC aC aeo AL5

Notice of Report on Mammals from Dutch New

(Gia ee RAR the ee sete We or libre A et Nhs ee Pe 485

Tuomas, Mrs. R. Haie, F.Z.S.

Exhibition of Skulls of Hornless PAN OLO MCSE lsnseiasaie 490

THompson, Surgeon Josepy C., U.S. Navy.

Further Contributions to the Anatomy of the Ophidia . 379

XV1

Page

Torton, A. KNYVETT.

The Structure and Development of the Caudal Skeleton

of the ‘Teleostean Fish, Plewragramma antarcticum.

(Bl ates Wc WN ee ieee enee ce alee aoe ie en cteniene eeierioe erie 251

Watson, Davip M. §&., M.8c., F.Z.5.

On the Skull of a Pariasaurian Reptile, and on the

Relationship of that Type. (Text-figures 1-7.) ............ 155

Exhibition of Procolophon trigoniceps, a Cotylosaurian

VEC) OME ae Ns eae oes “1d ov dialaleatn aie ade atheceate ean een eae 490

Woop-Jonrs, FrepEertc, D.Sc., F.Z.S.

Some Phases in the Reproductive History of the Female

Mole (Talpa europea). (Plates I—III., and Text-figures

EEG) Sane A en PRREMED Sho AScincn gona nn ddd ie sdo oto aede 191

INDEX.

1914.—Pages 1-490.

| New names in clarendon type.

Systematic references in italics.

Z.S.L. indicates additions to the Society’s Menagerie. |

Acanthocephala :

from the Antarctic,

Acanthocycline, 344.

Acunthocyclus albutrossis, 344.

gayi, d44.

—— hassleri, 344.

Acomys ablutus, 3109.

——— igiitus, 315.

Acontivstoma marionis, 356.

Allocreadium fowleri, sp. n. 224. |

Allorchestes patagonicus, 368. |

Ampelisca macrocephalus, 357.

Amphitoé (Melita) inequistylis, 366.

(——) tenuicornis, 366.

Ampithoe brevipes, 571.

999

Anas boscas: physiology: variation |

(Figs. 1-6), 23.

Anatomy (see Structure).

ANNDLIDA :

Ceratocephale & Tylorhynchus, 231.

Nereide frou N.M. Pacitic, 237.

Anomotzenia zederi: ethology, 221.

Anthobothrium wyatti, sp. n.,

225.

Anthodon serraius :

(Fig. 7), 165.

Antilope cervicapra (Z. 8. L.), 486.

Aponurus bowersi, sp. u., 224.

Apteryx mantelli: ethology, 487.

Apus, 303.

ARACUNIDA :

from Moute Bello Islands:

matic, 69.

skull - structure |

syste- |

Proc. Zoot. Soc.—1914, No. XX XIII.

ARACHNIDA (€02.)

from Dutch New Guinea: systematic,

487.

Pseudoscorpiones: structure, 93.

Araneus reversus, sp. n. (PI. I.

fig. 5), 77.

Arctictis binturong (Z. 8. L.), 480.

Argiope haynesi, sp. nu. (PI.

fiy. 3), 73.

trifasciata, 73.

Aristeus goldiet, 339.

Artemisia westraliensis, 296.

Arvicanthis abyssinicus nairobe, 315.

preceps, 316.

rubescens, 316.

——— rumruti pallescens, subsp. n.,

316.

striatus massaicus, 315.

testicularis gebele, 316.

Ascaris osculata: ethology, 221.

--— radiata: ethology, 221.

rectangula: ethology, 221.

Astacilla fulclandicus, 353.

Atyloides, 564.

magellanicus, 360.

Atylopsis magellanicus, 365.

Atylus austrinus, 305.

—— huxleyanus, 362.

AVES:

Anas boscas: physiology : variation,

DOs

Apteryx mantelli: ethology, 487.

Pavo: hybrid, 218.

XVill

Ayus (con.):

Pseudotantalus leucocephalus : colour

development, 457.

Bos taurus (Z. 8. L.), 219.

Bovallia regis, sp. n. (Pl. VIII),

362.

Brachyccelium obesum, sp. n.

(Pl. ILL. fig. 8), 147.

Branchinella australiensis, var.

occidentalis, nov. (Pl. 1. figs. 1-5),

296.

—— eyrensis, 298.

-longirostris, 297.

northamensis, sp. n. (Pl. I.

figs. 6-8), 298.

Burmagomphus vermiculatus

insularis, subsp. n. (PI. I. fig. 2),

55.

Calophasis mikado x C. ellioti (hybrid)

(Z.8. L.), 218.

Camelus bactrianus: palatal appendage,

476.

dromedarius: palatal appendage

(Hig. 1), 476.

Cancer minutus, 345.

—— (Gammarus) faleatus, 371.

Canis culpeeus (Z. 8. L.), 475.

—— famuliaris: structure (feet) (Figs.

1-3), 478.

lateralis, 310.

Caprella acutifrons, 373.

dilatata, 373.

penantis, 373.

Carnivora (fossil): skull-structure, 227.

Cassidina emarginata, dol.

Cassidinopsis emarginatus, 351.

Causus rhombeatus: anatomy, 401.

Cephalophus grimmia lutea,

subsp. n., 318.

Ceratocephale osawai, 231.

Ceratophrys ornata (Z. 8. L.), 218.

Cerberus rhynchops: anatomy, 594.

Cerchneis cinnamomina (Z.8. L.), 475.

Cercopithecus pugerythrus centralis,

308.

Cervicapra chanleri, 318.

Cervus elephas: antlers, 488.

INDEX.

CrsropA:

from the Antarctic, 222.

Cittotenia lagorchestis:

420.

villosa : anatomy, 427.

Ichthyotenia filicollis:

life-history, 113.

Oochoristica & Linstowia; structure :

systematic, 263.

Tetrabothrius strangulatus: anatomy :

systematic, 407.

Urocystidium : anatomy: life-history,

Ih.

Chelodina expansa (Z.8. .), 219.

Chlorogomphing (Pl. 1. figs. 4-8), 51.

Cheeropsis liberiensis (Z. 8. L.), 218.

Chrysotis versicolor (Z. 8. L.), 486.

Cissites maxillosa: geographical, 476.

Cittotzenia lagorchestis, sp. n.

(Pls. I.-VI.), 420.

quadrata: anatomy (Fig. 3),

426.

—— villosa, sp. n. (Pls. V.-X.), 427.

zschokkei: anatomy (Figs. 1, 2),

424.

COLEOPTERA :

anatomy,

structure :

frum Dutch New Guinea : systematic,

476.

Colobus caudatus (Z. 8. L.), 217.

Coluber oxycephalus : anatomy, 389.

Conurus euops (Z. 8. L.), 219.

Corophium cylindricus, 372.

Corynosoma antarcticum: ethology,

221.

—— hamanni: ethology, 221.

Crassicauda, gen. n., 226.

Crax alberti (Z. 8. L.), 490.

Crocidura gacksoni, 309.

~ monax, 309.

CRUSTACEA :

from Falkland Islands: systematic,

341.

Phyllopoda: geographical, syste-

matic, 293.

Potamonide. Dutch New Guinea :

systematic, 485.

| Crypturus noctivagus (Z. 8. L.), 490.

Cureeus aterrimus (4.8. L.), 490.

Cyclinea, d44.

INDEX.

Cyclops varius: ethology (PI. IV. figs.

33, 34), 113.

Cymodocea darwinit, 351.

Cyneelurus (ZaS ss) pee 21;

218.

Cynodictis intermedius: skull-structure

(Pl. I. figs. 1-3), 227.

Cynodon gracilis: skull-structure (PI. I.

figs. 4, 5), 227.

Cyon dukhunensis (Z. 8. L.), 219.

CytTouoey :

Cell-volume in various organisms,

45,

Cyzicus (Estheria) rufa, sp. n.

(Pl. IL. figs. 19-21), 301.

jubatus

Dendromus acreus, 3138.

lineatus, 313.

nigrifrons, 313.

DEVELOPMENT:

Mammalia: Talpa europea (genitalia),

191.

Aves: Pseudotantalus leucocephalus

(colour), 457.

Pisces: Pleuwragramma antarcticum,

251.

Seyllium canicula, 435.

Ichthyotenia filicollis,

Vermidea :

113.

Tetrabothrius strangulatus, 407.

Urocystidium, 1.

Echinoderma: Echinus esculentus

(abnormal), 65.

Dibothriocephalus archeri,sp.n.,

294.

coatesi: ethology, 221.

—— lashleyi, sp. n., 224.

mobilis: ethology, 221.

Dicotyles tajacu (Z. 8. L.), 490.

Dieta isolata, sp. n. (Pl. IL. fig. 7),

80.

Diphyllobothrium perfoliatum :

ology, 221.

rufum, sp. n., 224.

Dipodillus harwoodi, 311.

Dolichotis salinicola (Z. 8. L.), 475.

Doliophis bivirgatus: anatomy, 400.

Dules nitens, 340.

Dynamene darwinit, 352.

eth-

Ecixopurma :

Echinus eseulentus : abnormal, 65.

Echinorhynchus campbelli,

sp.n., 223.

debenhami, sp. n., 223.

—— rennicki, sp. n., 223.

turbinella: geographical :

ology, 223.

Echinostomum aliud, sp. n. (PI.

IV. figs. 9, 9a), 148.

Kehinus eseulentus: abnormal develop-

ment (Figs. 1-4), 65.

Edotia tuberculatus, 358.

Elephantulus dundasi, 309.

Hlephas spp. (Figs 2, 3), 286.

maximus (Z. 8. L.), 490.

— maximus hirsutus, subsp. n.

(Fig. 1), 285.

Hlininius hingwi, 376.

Epimys coucha pallida, sp. n.,

314.

eth-

panya, 314.

—— jacksoni, 314.

—— medicatus, 314.

—— niveiventris, 314.

— walambe amale, subsp. n.,

313.

Epomophorus anurus, 308.

minor, 808.

Equus grevyi (Z. 8. L.), 486.

ErnoLoey :

Mammalia: Otaria pusilla x O. cali-

forniana : hybrid, 219.

Aves: Apteryx mantelli, 487.

Reptilia: Testudo abingdonii (Z.S.L.),

220.

Arachnida: Pseudoscorpiones: nest-

formation, 93.

Vermidea: Helminthes from various

animals, 21.

Trematoda: from Birds and Rep-

tiles, 139.

Cestoda: Cittotzenia lagorchestis

and C. villosa from a Wallaby,

419,

Ichthyotznia filicollis from the

Stickleback, 113.

Linstowia ameivx from a Lizard,

263.

Ernonoay (con.) :

Vermidea:

Cestoda : Ovchoristica

from a Marsupial, 263.

Tetrabothrius strangulatus from

an Albatross, 407.

Urocystidium from the Mus-

marmosz

quash, l.

Nematoda from yarious animals,

181.

fudimnadia badia, 301.

cygnorum, sp. n. (Pl. I. figs. 9-

13), 299.

—— feriensis, sp. n. (Pl. I. figs. 14-

18), 30.

—— rivolensis, 300.

Eupagurus comptus, 3416.

—— forceps, 346.

Eusiroides orchomenipes, 362.

(Z. 8. .),

Eutamias

218.

Exospheroma caleareus (PI. I1.), 350.

coatsit, 850,

quadrivittatus

Felis capensis hindei, 309.

tigris (Z. 8. L.), 490.

geographical :

Filaria crassicauda :

)s)s

ethology, 228.

Forficula auricularia :

(Pl. IT. figs. 138-17), 48.

cell-voluime

Galathea gregaria, 346.

subrugosa, 347.

Gallus domesticus X Pavo’ nigripennis, —

(GES) i) 2

Gammarus barlimanus, 370.

fuegiensis, 369.

Gasteracantha minax, 79.

—-—, var. astrigera, 79.

fig. 6), 80.

, var. lugubris, 79.

Gazella granti raineyi, 318.

Geneita erlangeri, 309.

GEOGRAPUICAL:

Maiunalia: Fast Africa, 307.

——-, var. hermitis, nov. (PI. I. |

Carnivora: Phosphorites of Quercy, |

wal.

INDEX.

GoGRAPMICAL (co7.) :

Mamunalia: Cervus eleplias (antlers),

Wurope; 488.

Elephas maximus hirsutus : Malay

Peninsula, 286.

Lycaon pictus: 8. Afriea, 403.

Sciurillus : Guiana, 410.

Pisces : Dutch New Guinea,

226.

Tusecta: Coleoptera: Dutch New

Guinea, 476.

Odonata: Borneo, 51.

Arachnida: Monte Bello Islands,

69.

Dutch New Guinea, 487.

Myriopoda: Dutch New Guinea,

487.

Onychophora : W. Australia,

289.

Crustacea; Phyllopoda of W. Aus-

tralia, 293; Distribution in Aus-

tralia, 294.

Mollusca: Dutch New Guinea,

475.

Vermidea: Helminthes: Antarctic,

221.

Annelida: Ceratoceplale and

Vylorbynchus: distribution,

231.

Nereidx ; N.E. Pacific, 237.

Gerbillus cosensi, sp. n. 311.

Gomphidia kirschi, 53.

maclachlant, 53.

Gomphine (P\. 1. figs. 1-8), 51.

Gomphus consobrinus, 55.

helantanensis, 55.

——- vermiculatus, 5d.

Goura coronata (Z. 8. L), 217.

Graphiw us brockmani tuternus, 310.

Halirages huxleyanus, 362.

Haplocheira barbimanus, 370.

typica, 371.

Harmotrema infecundum, gen. et

sp. n. (Pl. LV. fig. 10), 150.

Heliosciurus multicolor elegans, 310.

Helix pomatia: cell-volume (PI. IL.

figs. 18-24), 45.

INDEX. XXxl

Helminthes : from the Antarctic, 221.

Helogale percivali, 310.

undulata rufula, 310.

victorina, 310.

Hemistomum canaliculatum,

sp. n. (Pl. IV. fig. 11), 151.

Hemiurus oatesi, sp. n., 224.

Heterogomphus icterops bor-

neensis, subsp. n. (Pl. I. fig. 3),

57.

Hippolyte magellanicus, 347.

Hyalella patagonicus, 368.

Hylocichla ustulata swainsoni (Z. 8. L.),

218.

Hyperia gaudichaudii, 374.

Hystrix africe-australis, 317.

Ichthyotwnia filicollis: structure: life-

history (Pls. IL-IV.; Figs. 1-5), |

113.

Icterus pyrrhopterus (Z. 8. L.), 219.

Ictinus acutus, sp. u. (Pl. L. fig. 1),

51.

decoratus, 52.

melenops, do.

, race swmatranus, 5d.

Ictonyx capensis albescens, 309.

Insucra :

Coleoptera: Dutch New Guinea: syste- |

matic, 476.

Odonata : structure: systematic, 51. |

Trichoptera: seent-organs, 409. |

Iphimedia nodosus, 358.

normant, 399.

Jassa faleatus, 371.

Kathleena scotti, gen. et sp. n., 223,

226.

Kuhlia humilis, 340.

Lachesis neuwiedii (Z. 8. L.), 490.

Lama vicugna (Z S. L.), 218.

Larinia montagui, sp. n. (Pl. I. |

fig. 4), 75.

Lavia frons frons, 308.

Lembos fuegiensis (P). 1X.), 369.

kerguelent, 369.

| Lophvceros erythrorhynebus (Z. 8. L.)

Lepas australis, 376.

Lepidurus viridis, 304.

, var. elongatus, 304.

Lepodora garrardi, sp. n., 224.

Leptodira hotambeeia : anatomy, 397.

Leptogomphus kelantanensis, 5d.

senupert, d+.

williamsoni, 54.

Leptosomatum setosum: ethology,

22),

Leptostraka, 374.

Lepus victorie, 317.

Linstowia, 281.

ameive, sp. n. (Figs, 1-4),

269.

PAE

Lophuromys zena, 315.

Lycaon pictus: variation, 403.

Lycosa clara, €8.

Lynceus tatei, 303.

Lyperosomum direptum, sp. n.

(Pl. III. fig. 7), 147.

—— scitulum, sp. n. (PI. III. fig. 6),

146.

Macrochiridothea, 353.

— stebbingi, 354.

Macrogomphus albarde, 5A.

——- decemlineatus, 54.

quadratus, d4.

| Macrorhinus leoninus (Z. 8. L.), 489.

Mammatia :

from Dutch New Guinea: systema-

tic, 485.

from Hast Africa: systematic,

307.

Camelus: palatal appendage, 476.

Canis fauiliaris: structure (feet),

478.

Carnivora (fossil): skull-structure,

227.

Cervus elephas : antlers, 488.

Elephas maximus hirsutus: varia-

tion, 285.

Lycavon pictus: variation, 403.

Otaria: hybrid, 219.

Sciurillus: systematic: geographical,

415.

XXil

MamMatta (co.): |

Simia satyrus: longevity, 489. |

Talpa europea: structure: develop-

ment, 191.

Marpissa ridens, sp. n. (Pl. II.

fig. 12), 90.

Mediorima propria, gen. et sp. n.

Gab esis Wy Ie

Melita inequistylis, 366.

Merula fuscatra (Z. 8. L.), 217.

Metopa ovata, 358.

Metopella ovatus, 358.

Metopoides ovatus, 358.

Microgomphus chelifer, 5+.

Micronereis variegatu, 243. |

: structure: xtiology (Figs. |

2-5, 7), 245. |

Mimus gilvus (Z. S. L.), 475.

Miturga parva, sp. n. (Pl. II. fig. 8), |

82.

Mera fuegiensis, 369.

Mo.uvsca:

from Dutch New Guinea:

475.

Monoculopsis, 360.

—— vallentini, sp. n. (Pls. VI.,

VIL.), 360. :

Montebello, gen. n., 86.

—— tenuis, sp. n. (PI.

86.

Morpiooey (see Structure),

Mungos albicauda, 309.

sanguineus ibee, 309.

Munida gregarius, 346.

subrugosus, 347.

Mus bellus, 318.

gondokore, 313.

gratus, 313,

——- musculoides emesi, 3138.

triton, 313.

Mustela martes (Z. 8. L.), 218.

Myriopopa :

from Dutch New Guinea: systematic, |

487. |

Pseudoscorpiones: structure: etho-

logy, 93.

systematic,

Il. fig. 10),

Nasilio brachyrhynchus albiventer, 309.

Nauticaris magellanicus, 347.

INDEX.

Nebalia lipes, 575.

chilen:is, 379.

Nematocentris nove-quineeé, 339.

rubrostriatus, 339.

| NEMATODA:

999

from the Antarctic, 222.

from the Society’s Gardens, 181.

Neotragus pygmeus (Z. 8S. L.), 478.

Nephila meridionalis hermitis,

var. nov. (Pl. I. fig. 2), 72.

venosa, T1.

Nereis agassizi, 242.

—— brandti, 241.

eyclurus (Fig. 1), 237.

dumerilii, 242.

dynamusi, 241.

ezoensis, 24V.

heterocheta, 233.

—- kerzuelensis: parapodium (fig. 6),

248.

kobiensis, 242.

peiagica (Fig.

shishidot, 287.

vexillosa, 240.

virens, 241.

(Platynereis) agassizi, 242.

Nicoria nasuta (Z. 8. L.), 490.

Notasellus sarsit, 354.

Numida meleagris x Payo cristatus:

hybrid, 218.

Nycticebus tardigradus (Z. 8. L.), 470.

7), 240.

Odonata: Gomphine & Chlorcgomphine :

from Borneo, 51.

Ogmogaster plicatus :

ethology ; 223.

Olios calligaster, 84.

—— hermitis, sp. n. (Pl. Il. fig. 9),

85.

Ommatobrephus singularis, gen.

et sp. n. (Pl. I. fig. 2), 141.

Onychophora: from W. Australia, 289.

Occhoristica, 281.

Marmose, sp. n. (Figs. 4-8),

280.

Ophidia : anatomy, 379.

Ophiophagus bungarus: anatomy, 398.

Opisthioglyphe adulescens,

sp. n.(Pl. I. fig. 3), 148.

geographical :

INDEX.

Opisthogenes interrogativus,

gen. et sp. n. (PI. I. fig. 4), 142.

Orchestia scutigerula, 3067.

Oreotragus oreotragus schillingst, 317.

Oriana wilsoni, gen. et sp. n., 225.

226,

Orogomphus atkinsent, 61.

dyak (Pl. I. figs. 4-7), 59.

splendidus (Pl. I. fig. 8), 60.

Otaria pusilla x O. californiana: ly-

brid, 219.

Ourebia ourebia cottoni, 317.

Ovis vignei (Z. 8. L.), 475.

Oxyopes mundulus, 88.

Pagurus comptus, 346.

—— forceps, 346.

Papio furax, 308.

Papuina lituus: anatomy, 476.

Paradexamine nanus, sp. n., 366.

Paranera austrinus, 369.

Paraxerus ochraceus electus, 310.

Pariasauwrus: skull (Figs. 1, 2, 4-6),

155.

—— hombidens: skull (Fig. 3), 160.

Pariphimedia, 358.

normant (Pls. IV., V.), 359.

Parimo.oey :

Animals in the Society's Gardens,

181.

Payo cristatus X Numida meleagris:

hybrid, 218.

-nigripennis X Gallus domesticus:

hybrid (Z. 8. L.), 217.

Pelomys fullax tridescens, 317.

Peripatordes gilesti (Fig. 1), 289.

—— occidentalis, 289.

—— woodwardi (Fig. 1), 289.

Peripatus leuckarti, var.. occidentalis,

289.

Perodicticus ibeanus (Z. 8. L.), 486.

Peucetia margaritata, sp. nun.

(Pl. II. fig. 11), 89.

Philothamuus

senlivariegatus: ana-

tomy, oot.

Piryganea: head (diagrammatic),

464.

Phyllopoda: from W. Australia, 298.

XXill

PuysioLoey :

Mammalia :

ity, 485.

Aves: Anas boscas, 23.

Simia satyrus: longey-

Insecta: ‘Trichoptera: scent-organs,

459.

Pilerodius pileatus (Z. 8S. L.), 490.

Pipistrellus deserti, 308.

Piscus :

from Dutch New Guinea: systematic,

226.

from New Guinea: systematic, 339.

Pleuragramma antarcticum: stiuc-

ture: development, 251.

Scyllium canicula: venous system,

4305.

Teleostei : lateral muscle, 319.

Planes minutus, 845,

Platophium brasiliense, 373.

Pleuragramma antarcticum; structure:

developiment (Pls. I., II.), 251.

Podocerus brasiliensis, 373.

— cylindricus, 372.

Podocotyle pennelli, sp. n., 224.

Poliopsar leucocephalus (Z. 8. L.), 217.

Polyodontophis collaris :

38l.

geminatus: anatomy, 380.

Pontharpinia rostratus, 357.

Pontogeneia antarcticus, 364.

magellanica, 365.

Potamonide: from Dutch New Guinea,

485.

Prorozoa :

Parasites from the Society’s Gardens,

181.

Psammophis sibilans: anatomy, 398.

Pseudoscorpiones: Nests and Spinning-

organs, 93.

anatomy,

Pseudvtantalus leucocephalus: colour

development, 457.

Pseudoxenodon sinensis: anatomy, 385.

Pygoscelis adelize (lecture notice), 487.

Pyranga bidentata (Z.8. L.), 475.

Raphicerus neumanni, 317.

Reprivia :

Ophidia: anatomy, 379.

Pariasaurus: skull-structure, 155,

XX1V

ReEprrxia (co7.) :

Testudo abingdonii: variation : eth-

ology, 220.

Rhinochetus jubatus (Z. 8. L.), 217.

Rhombosoma goldiei (Figs. 1, 2), 339.

nove-quine@, 339.

Rhynchotragus cavendishi, 317.

— nasoguitatus, 318,

Sciurillus, gen. n., 416.

Sciurus finlaysoni (Z. 8. L.), 479.

saltuensis bonds (Z.8. L.), 490.

Scoteinus schliefent, 509.

Scotophilus nigrita, 308.

Scyllium canicula: venous system (Pls. |

I., IL; Figs. 1-4), 435.

Sericostuma personatum : scent-organs |

(Figs. 1, 2, 4-8), 409.

Sicboldius grandis, 53.

japponicus, 03.

Simia satyrus: longevity, 485.

Spheroma calcarea, 35.

Stenobothrus curtipennis: cell-volume

(Pl. L., figs. 10-12), 45. |

cell-volume (Pl. I., |

——- viridulus :

fig. 9), 40.

STRUCTURE :

Mammalia: Camelus (palatal ap-

pendage), 476.

Canis familiaris (feet), 478.

Carnivora: (fossil skull), 227.

Sciurillus, 415.

Talpa europea (genitalia), 191.

Reptilia: Ophidia, 379

Pariasaurus (skull), 155.

Pisces: Pleuragramma antarcticum,

251.

Scyllium canicula (venous system),

435.

Teleostei (lateral muscle), 319.

Insecta: Odonata: Gomphinz and

Chlorogomphine, 51.

Trichoptera (scent-organs). 459.

Arachnida: Pseudoscorpiones (spin-

ning-organs), 93.

Vermidea: Annelida: Nereidx, 237.

Cestoda : Cittotenia lagorchestis

and C. villosa, 419.

INDEX.

STRUCTURE (co7.) :

Vermidea :

Cestoda: Ichthyotwnia filicollis,

113.

Linstowia ameivz, Oochoristica

Marmosze, 265.

Tetrabothrius strangulatus, 407.

Urocystidium, 1.

Echinoderma: Echinus esculentus

(abnorinal), 65.

Struthio australis (Z. 8S. L.), 218.

— massiicus (Z. 8.1L), 217.

Styphlodora persimilis, sp. n.

(P1. IL, fig. 5), 144.

Sus scrofa (Z.S8. L.), 475.

Sycalis minor (Z. 8. L.), 490.

Sylvilagus superciliaris (ZS. L.), 490.

Symbranchus bengalensis: geographi-

cal, 226.

Tachyoryctes naivushe, 317.

—-. ruddi badius, 317.

Teenia filicollis: structure: life-history

(Pls. L-IV.; figs. 1-5), 113.

Talitride, 367.

Talorchestia scutigerulus, 367.

Talpa europea: structure: develop-

ment (genitalia) (Pls. 1—I1I.; figs.

1-13), 191.

Tanaide, 348.

Tanais ohlini, sp. n. (Pl. L.), 349.

Taphozous mauritianus, 309.

Tatera niyricauda, 312.

Taterillus emini, 312.

—— lowei, sp. n., 312.

Taurotragus oryx (Z.S. L.), 486.

Teleostei: structure of lateral muscle

(Figs. 1-4), 319.

Terranova antarctica, gen. et

sp. n., 226.

Testudo abingdonii (Z. 8. L.), 219.

—— : variation: ethology (Fig. 1),

220.

Tetrabothrius aichesoni, sp. n.,

225,

—— catherine, sp. n., 225.

—— creani, sp. n., 225.

cylindraceus; geographical: etho-

logy, 225.

INDEX.

Tetrabothrius nelsoni, sp. n., 226.

—— priestleyi, sp. n., 225.

—— strangulatus, sp. n. (Figs.

1-4), 407.

—— wrighti, sp. n., 225.

Tetragnatha angulata, sp. n.

GAM, Ie see, 1) 7):

Thamnomys surdaster, subsp., 315.

Thamnophis ordinoides: anatomy,

831.

TREMATODA :

from the Antarctic: systematic,

222,

from Birds and Reptiles: systematic,

139.

Trematomus borchgrevinki: structure

(Pl. II. fig. 14), 251.

—— newnesii: structure (Pl. II. fig.

13), 251.

Trichoglossus rubritorques (Z.S. L.),

218.

Trichoptera: scent-organs, 459.

Triton cristatus: cell-volume (Pl. I.

figs. 1-8), 45.

Tropidonotus sauteri : anatomy, 385.

vibakari: anatomy, 384.

Tryphosites, 355.

chevreuxi, sp. nu. (Pl. IIT.),

355.

Tylorhynchus chinensis, 231.

Urocystidium gemmiparum: anatomy:

life-history (Figs. 1-9), 1.

Urothoe rostratus, 357.

Vallentinia, gen. n., 351.

darwinti, 351.

Variation and Ar1o1oey :

Mammalia : Cervus elephas (antlers),

488.

Elephas maximus hirsutus, 285.

Lycaon pictus, 403.

Proc. Zoou. Soc.—1914, No. XXXIV,

XXV

VariaTIon and Aitronocy (con.) :

Mammalia: Otaria pusilla x O. cali-

forniana (hybrid), 219.

Aves: Anas boseas, 23.

Calophasis mikado x C.

(hybrid), 218.

Pavyo cristatus X Numida meleagris

(hybrid), 218.

Reptilia: Testudo abingdonii, 220.

Vermidea: Micronereis variegata, 245.

Nereis cyclurus, 237.

VERMIDEA :

Trematoda: Antarctic : systematic,

ellioti

222.

from Birds and Reptiles: syste-

matie, 139.

Cestoda: Antarctic: systematic,

222.

Cittotznia: anatomy: systematic,

420.

Ichthyoteenia : structure, 113.

Oochoristica & Linstowia: struc-

ture : systematic, 263.

Tetrabothrius: structure, 407.

Urocystidium: structure, 1.

Nematoda: Antarctic: systematic,

299

ate

from the Society’s Gardens, 181.

Acanthocephala : systematic, 222,

Annelida: Ceratocephale & Ty!o-

rhynchus: systematic, 231.

Nereide: geographical: syste-

matic, 237.

Xenodon merremii (Z. S. L.), 490.

Xenopeltis unicolor: anatomy, 380.

Xerus dabagala dorsalis, 310.

Zamenis constrictor: anatomy, 386.

—— florulentus: anatomy, 389.

rhodorhacis : anatomy, 387.

Zelotomys hildegardee, 315.

avai

PRINTED BY TAYLOR AND FRANCI

: ie RED LION COURT, FLEET STREET. ,

PROCEEDINGS

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.

1914.

PART’ I.

CONTAINING Paces 1] To 226, witH 16 PLaTEs

AND 45 'TEXxT-FIGURES.

We.

tional Muse

PRINTED FOR THE SOCIETY,

SOLD AT ITS HOUSE IN REGENTS PARK,

LONDON:

MESSRS. LONGMANS, GREHN, AND CoO.,

PATERNOSTER ROW.

[Price Twelve Shiliings. |

LIST OF CONTLENTS.-

1914, Parr I. (pp. 1-226).

EXHIBITIONS AND NOTICKS,

Page

The Srcrnrary. Report on Additions to the Society’s ge during the months of

November and December 1913 ............ eee. cece RI sro ors = LL

Mr. D. Sern-SsarHy, F.Z.8., Curator of Birds. Exhibition of photographs and skin of “

AEliy TG WB UEAS 9 le tecaa Se Cio becn, nie +n <u tecepeh atelier ale Brinda Ss Hcl Meh acoso 218

Mr. G. A. Boutuncer, F.R.S., F.Z.8. Notice of paper on the Batrachians and Reptiles —

collected by the British Ornithologists’ Union and Wollaston a xpos in Dutch

SNe ws Grulimeaius coal Gian tarertre eerie wie sree Re Meh Pea ate tig eee aie ote wiaisbee ae ae ate

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

January 1914 ........ eS by ERO Bee oe SCO ty Cito 5 geo iars Sieiui shes e [eee valwintetee Seek)

Mr, Grorce Jennison. Exhibition of a mounted specimen of a a Hybad Sea-Lion ...... 219

Mr. E. G. Bourencnr, F.Z.8., Curator of Reptiles. Exhibition of photograph of the ee! 2

7 os

ra ee Nee a yee oe” ty a ee Oe ee eee eT.

Saddile-backedMortoise 2 stesrg ciitecs sce wae cua sitwue oe «wae eletoes ena ie eres Ieee 220.

Dr. R. T. Lererr, M.B., F.Z.S8., and Surgeon E. L. Atxinson, R.N. Lantern demonstration

of the Helminthes collected by the British Antarctic (‘Terra Nova’) Expedition,

SOMO The eel rrr eS Gre Ame Rian Sminicde a Anco O.c.c oe eisai sie 8 acere teneremammona

Mr. C. Tarn Ruean, M.A., F.Z.S. Notice of paper on the Freshwater Fishes collected

by the British Ornithologists’ Union and Wollaston Expeditions in Dutch New Guinea. 226

PAPERS.

1, Contributions to the Anatomy and Systematic Arrangement of the Cestoidea,—

XII. Further Observations upon the Genus Urocystidiwm Beddard. By Frann EH,

Brpparp, M.A., D.Sc., F.R.S., F.Z.8., Prosector to the Society. (Lext-figs. 1-9.) .. 1

Contents continued on page 3 of Wrapper.

PROCEEDINGS

OF TIE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF TIIK

ZOOLOGICAL SOCIETY OF LONDON.

PAPERS.

1. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea. By Frank E. Bepparp, M.A.,

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

XII. Furruer OBSERVATIONS UPON THE GENU s<qgonian [i In 1Sti,

Urocystipium BEDDARD. / e “an \

[Received November 12, 1913: Read February 3, 19144 M AY 6

(Text-figures 1-9.)

1 :

Stional MuseY=

Urocystidium : LSE, Page ar ae

INCE CU ROUTAN nn nbdeoe Guay -usoseeb coo eas nooECUECondbemeoneeeneae 1)

SOM MnOL MMe nena er yetahe arc matuomeeecneacereaandssaaciyataiea ory wd

SSUTTUIN ARV geR sere See neeieh nick, Wan ati a ouniuiscaiossiesines, csi nonesayh OU

In December 1912* the Society published a communication

of mine upon a tapeworm from the Musquash (iber zibethicus),

of which I was able to describe both the immature and mature

stages which lived side by side in the liver in cavities which are

pr obably to be regarded as dilated portions of the liver-ducts. In

May of last year I found some more examples of the same or

a closely allied worm in the same situation in another specimen

of the Musquash.

Tn this second case of infection the parasites, as in the first

case, consisted of asexual and sexual examples, thus confirming

my original discovery that the two generations of the worm

lay side by side in the same organ of their host. The asexual

* P,Z.S, 1912, pp. 822-850.

Proc. Zoou. Soc.—1914, No. I. 1

2 DR. F. E. BEDDARD ON

individuals were five in number, of which four are figured in

text-fig. 1; the fifth was an obviously incomplete individual.

T shall commence the present communication to the Society with

an account of

$ The Asexual Forms.

The series of examples which represent the asexual phase of

this Urocystidium showed no budding such as I originally

Text-figure 1.

A. Completely developed asexual stage of Urocystidium. B. Incomplete individual

belonging to asexual generation. OC, D, EH. Three younger individuals of the

asexual generation which represent stages in the development of the asexual

form A.

described as the most salient and novel character in this genus.

The specimens, however, which I am now about to describe, are

not inconsistent with a development of this kind. That is to say,

MAMMALIAN CESTODES,

there are no reasons for the belief that the budding which I

formerly described was in any way abnormal in the species.

For it seems to me to be possible, though, as I shall point out

later, by no means certain, that the three maggot-shaped worms

represented in text-fig. 1 may be looked upon as newly liberated

buds, perhaps being derived from the very long specimen (text-

fig. 1 A) which has since ceased to produce buds. This latter

specimen measured 83 mm. in length by a greatest diameter of

35mm. This greatest diameter was at one end of the body; at

the other end it measured only 2mm. There were intermediate

widths in different regions. Although the greater width of one

end of the body suggested that this was a bladder such as I

described in the original specimen, the segmentation was observed

to be continuous here as elsewhere.

The same text-figure (text-fig. 1) illustrates the three younger

asexual forms, which were of varying size, the largest being

presumably the eldest. In any case I have found that the inter-

mediate sized specimen is older than the smallest. The largest

measured 9 mm. in length by 4mm. in diameter. The propor-

tions of the other two to it and to each other are correctly shown

in the drawing referred to. These embryos show a segmentation

least marked in the smallest example. It isa rather fine cross

wrinkling ; but, as I shall show in the sequel, is a true segmenta-

tion. When these embryos were living and rather transparent,

they exhibited spherical spaces in the interior, visible through the

outer wall, which are shown (and somewhat exaggerated) in the

text-figure which has just been referred to. These cavities

obviously suggest an internal budding of scolices as in Cenurus,

etc.; but it is not by any means clear that they are of that

nature.

I examined the smallest of these three young worms by means

of a complete series of transverse sections, and the middle-sized

one in longitudinal sections; the structure has been compared

with that of the fully grown asexual worm described in my earlier

paper upon Urocystidiwm. I have, however, made some fresh

sections of the fully grown asexual worm for the purposes of this

comparison. The structure of the very immature maggot-like

larvee explains certain features in the structure of the fully grown

asexual form which I was formerly unable to explain. I shall,

however, preface these comparisons by a detailed account of

§ Lhe structure of the Young Plerocercoids.

As will be seen later, the separation of these from the more

advanced stage of the sexless worm is purely arbitrary. But, as

there is an hiatus in- point of size and general appearance, it is

convenient to treat of them alone. Pstraye

I use the term ‘ Plerocercoid” in: the loosest. way, for an

immature tapeworm which can be referred to: neither the (ysti-

cereus nor Piestocystis type. Indeed it cannot be referred with

A. “DR: F. E, BEDDARD ON

more exactitude to the Plerocercus type, but perhaps on the

whole is nearer to that type than to the others. The fact is

that the present form and, for the matter of that, the long known

Cysticercus fasciolaris necessitate the revision of the nomen-

clature of the immature Cestodes. I do not, however, propose

such a step in the absence of full knowledge of the present genus,

Urocystidium, and therefore adopt temporarily the term Plero-

cercoid. It is I think generally held that the name Prestocystis

Text-figure 2.

Transverse section of larva which is figured entire in text-fig. 1, E.

A, Cavity surrounded by gland-cells, B. Bladder-cavity. C. Third kind of cavity

perhaps derivable from A, d. Tubes of water-vascular system.

should be applied to those “ bladder” worms in which the cavity

of the bladder is filled up to a greater depth with parenchy-

matous tissue, thus leaving the central cavity comparatively

small and surrounded by very thick walls. On the other hand,

Piestocystis might be converted into the Plerocercus type by the

complete obliteration of the bladder cavity. An apparently

intermediate condition is recorded by Max Braun in his account

of the Cestoidea in Bronn’s ‘ Thierreich,” in which (Cysticercus

MAMMALIAN CESTODES. 5

spherocephalus Rudolphi) the bladder is divided up by trabecule.

Urocystidiwm is not definitely of any of these types.

Text-figure 2 represents a transverse section through about

the middle region of the youngest of the two individuals which I

have studied. In this region the worm is at its broadest ; there

is no question of a posteriorly situated bladder, as in the.complete

immature worm which I described and figured in my former

paper. The diameter is at most 2°5-2°8 mm. ‘The parenchyma

is Interrupted by numerous oval to cireular and more elongated

cavities which occupy the greater part of it. These cavities vary

in number and size in different regions. The largest number

Text-figure 3.

Portion of a transverse section near to that represented in text-fig. 2, but more

highly magnified.

e. External surface showing cuticle and subcuticular layer. 4. Cavity lined by

cuticle, outside which are cells, and corresponding to cavity 4 in text-fig. 2.

that I have counted in a given section is 17; the smallest

number——and this occurred only at the two ends of the body—

is 1. I have found no direct relation between the width of the

body and the number of cavities; that is to say, the cavities are

more numerous at one end of the body, which measured only

2mm. in diameter, than in the section figured, which measured

nearly 3 mm.

' But although there is no direct relation of this inal it isa fact

that it is only at one end of the body that the cavities are most

numerous. This may possibly, for reasons which wiil be apparent

6 DR. F. E. BEDDARD ON

later, be termed the anterior end. But although there are facts

which point to the view that this is the anterior end, there

appears to be no difference in minute structure between the two

ends of the worm. There is not even the rudimentary scolex

marked chiefly by its pigmentation, which I referred to in

my earlier description of the sexless form of this worm. The

cavities which occupy so much of the inside of the body can be

referred to four categories, and their differences are rendered

clear in text-figure 2. The cavity lettered “A” is distinguished

from the others by the structure of its walls. It is lined by a

structureless layer (see text-fig. 3) which quite resembles the

external cuticle of the worm. Round this cuticle are stalked

Text-figure 4.

Transverse section through the same larva as that represented in text-fig. 1, E,

in a place where the cavities C only are to be seen.

d. Water-vascular tubes.

glandular cells which are quite like those of the subcuticular

layer. These cells are very conspicuous, as is shown in the

figure. The cavities are the expression in transverse section of

tubes which are shortand do not branch so far as I could discover.

The obvious similarity to the external layer of the body which

they show led me to expect that an opening or openings on to

the exterior would be found. I have not, however, obtained

perfectly satisfactory evidence of such, and indeed have found no

actual orifice at all. In the case of one of these tubes, which I

MAMMALIAN CESTODES. tf

followed out from beginning to end, the tube became connected

at one blind extremity with the outer layer of the body by a

scattered group of subcuticular cells. This as it appears to me

may represent a previously existing pore. As I possess no

younger example of the plerocercoid than the one described here,

this point cannot be settled. Cavities having the character just

described are few in number and are not always to be seen in

a given series of sections. The letter B in text-fig. 2 points

to a cavity of a completely different character from that which

has just been described. This cavity is always single, and by

following out its course there is seen to be only one of this

nature in the whole body. Furthermore, it does not traverse the

whole body but is restricted to the hinder end (as I have termed

it), and does not nearly extend to the end which is occupied by

the numerous cavities of the third character (cf. text-fig. 4) to be

shortly described. The cavity is median in position and occupies

about three-elevenths of the entire length of the worm’s body. It

occupies the last third of the worm but does not by any means

extend to the very end. This cavity is not marked off from the

surrounding solid tissues by any definite wall. On the contrary,

in certain places at least the appearance is as if the tissues had

gradually become broken down at the edge to form this central

cavity. It fact it seems to me to be fairly certain that this is the

true bladder-cavity, which agrees in most particulars with the

bladder-cavity of such a form as the Piestocystis from the snake

Hoplocephalus superbus, described by Prof. J. P. Hill*, con-

eerning which that author writes +} :—‘“ The bladder-cavity in this

form is represented by an irregular cavity occupying the centre

of what represents the caudal bladder of ordinary Cysticerci and

not distinctly marked off from the surrounding ground-tissue.

The cavity is filled by a granular material consisting of a homo-

geneous matrix with granules which stain deeply with cochineal,

and which represent the products of degeneration of the original

central tissue.” Jam therefore disposed to compare and regard

as equivalent these spaces in the two immature tapeworms, and

to consider, as a consequence of this comparison, that the area in

Urocystidiwm which is occupied by the space in question is the

hinder end. There is, however, as I have already intimated, no

confirmation of this view by the discovery of a scolex. The

cavities lettered ““C” next require attention. These cavities

vary greatly in size, but are never so large as the bladder-cavity

already described. They are either absent or if present neces-

sarily peripheral where the bladder-cavity is at its largest, and

often come to lie close under the subcuticular layer of the body-

wall of the worm. Hence the clear spherical cavities discernible

in the living worm and duly referred to above.

These cavities, however, contain no trace whatever of any.

“* “ A Contribution to a further knowledge of the Cystic Cestodes.” Proc. Linn. '

Soc. N.S.W. (2) ix. p. 49. + Ibid. p. 52.

8 DR; F E. BEDDARD ON

budding scolices. They generally appear to be empty ; but traces

of an apparently coagulated fluid are to be observed after deep

staining with hematoxylin. They have invariably well defined

edges, and thus cannot be confused with the true bladder-cavity

as I assume it to be. In the section which is represented in text-

figure 2, these larger spaces with their well defined walls are

plainly to be distinguished from the space lettered A in that

figure with its lining of membrane and outer coat of glandular

cells. The difference is most striking and is accompanied also by

a difference in size which is apparent in the same drawing. Iam

not, however, convinced that there is a real difference. For some

of the cavities in question show distinctly a layer of cells im-

mediately outside of the cavity, and traces of a thin membrane

within that are to be noted. An expansion of such a tube as is

represented in A of text-figures 2 & 3 might well lead to such

appearances by a stretching of the lining membrane and by the

pulling apart of a close layer of cells. On the other hand, some of

the cavities lettered C (see text-fig. 4) are undoubtedly without a

distinct layer of cells around them. But here again the continued

expansion and perhaps the degeneration of the cells, if they are

concerned with the production of the fluid within the cavities,

may have produced such a result. ‘As I have no younger stages

than that of which I am now giving the description, the matter

does not as I think admit of a more definite expression of opinion.

The most careful examination has not shown any connection

between these cavities ; they are quite isolated as far as I could

make out; and if connections occur they cannot be abundant.

Nor were there any orifices on to the exterior to be observed. I ©

followed out through a series of sections the course of more than

one of the cavities and could find no outlet from their interior

in any direction. ‘There is, however, some evidence, though not

very strong, that they are formed out of the original bladder-

cavity. When the latter is carefully followed up to its anterior

end (as I have called it for reasons given), the cavity is seen to

acquire walls like those which surround the numerous cavities

lying beyond itsanterior end. It ends in fact in what looks very

much like one of the cavities lettered C. But this cavity does

not end in the parenchyma or in connection with one or

more of the rounded spaces just described. It is seen to lie

within one of the latter cavities and to end in it without

opening into it. But we may have here the last remaining

vestige of such a connection as has been suggested. Here

again the absence of younger stages forbids a more definite

statement.

The last series of cavities are those lettered d. These are

undoubtedly the tubes of the water-vascular system and are

peripheral in position. It is, however, to be noted that these

tubes and spaces do not invariably, though they do generally, lie

outside of the cavities belonging to the category C. The water-

vascular system consists partly of irregular spaces of a tubula

MAMMALIAN CESTODES. 9)

character, which occasionally end below and in contact with

the cuticle covering the body externally. I have not, however,

observed any actual pores on to the exterior. I have also found

the principal longitudinal vessel, which is the ventral vessel, as I

judge, of the completely developed sexless worm. I believe this

to be the case, since a transverse vessel arising from it was

occasionally (see text-fig. 5) quite plainly to be recognised.

I could not, however, find any continuous dorsal vessel lying

to the inside of this, such as is very obvious in the later stage of

the plerocercoid to be described immediately.

Text-figure 5.

Part of a transverse section of the same individual.

B. Bladder-cavity. d. Water-vascular tubes. ¢.v. Transverse water-vascular

tube. v.v. Ventral vessel.

The larger specimen, which was examined by means of longi-

tudinal sections, showed no great differences of structure from

the smaller individual. The same four systems of cavities were

distinguishable and presented the same characters. The spaces C,

as shown in the accompanying text-figure (text-fig. 6), are in

form roughly spherical and hardly at all elongate. They seem

to be, on the whole, actually smaller than in the younger stage,

which argues commencing disappearance, or rather reduction ; for,

as will be seen later, these various spaces are quite recognisable

in the completely developed and budding asexual worm. The

bladder-cavity (B) seemed to me to be rather larger in proportion

as well as actually than it is inthe younger plerocercoid ; and one

would of course have expected the contrary. It is, however, not

10 DR. F. E. BEDDARD ON

unreasonable to suppose that there are variations in the degree of |

progress of the reduction of this cavity. As before, the cavities A .

and C are chiefly at the two ends of the worm. iy

Text-figure 6.

Longitudinal section through larva figured in text-fig.1, D.

d.v. Dorsal water-vascular tube coiled and therefore seen cut across in several places.

thie m. Longitudinal muscles.

Other lettering as in text-figs. 2 & 5.

The water-vascular system (spaces lettered d) is more fully

developed in this older plerocercoid, and it differs from that of the

younger example in the presence of a perfectly distinct dorsal vessel’

MAMMALIAN CESTODES. 11

with very muscular walls, in addition to a much larger ventral

vessel. These are shown of the correct relative size in text-fig. 6,

d.v.& v.v. The branches of the ventral trunk which connect the

two ventral vessels of opposite sides of the body open as I have

described in the older worm, by two branches into the ventral

vessel. The transverse trunk traverses the bladder-cavity and is

surrounded on its transit by a layer of tissue which here, as else-

where, shows the commencing obliteration of the bladder-cavity.

In this worm the longitudinal and transverse muscular layers of

the body were very evident, and the worm was much pervaded by

caleareous corpuscles. The latter were often grouped a few

together in sacs not far below the subcuticular lay er. I shall recur

to these in considering the structure of

§ The Fully-developed Plerocercoid.

In my former paper upon Urocystidiwm I gave some account of

the general structure of the sexless worm which bears the buds.

I have now a few additional remarks to make upon this stage.

Text-fig. 7 represents a portion of a cross section to illustrate

the inclusion of groups of calcareous corpuscles in cavities below

the subeuticular layer. The corpuscles contained within are both

larger and smaller, and there is also some granular detritus. The

cavities themselves are rather elongated, but are always separated

by muscular fibres running between them and at right angles to

the longitudinal axis of the worm’s body. I have not studied the

development of these cavities. I refer to them for two reasons.

In the first place, the existence of quite similar agglomerations of

calcareous corpuscles in the younger plerocercoid is one of the

arguments which lead me to regard the various stages which I

am now describing as being actual stages in the development of

this worm—as for ming a true sequence in the order which I have

indicated. In the second place, these saes of calcareous bodies

are to be compared, as I believe, to apparently similar structures

recently described by Prof. J. P. Hill*. The figure’ given by

that author agrees very closely with that appended to the present

paper (text- fig. 7), and the worm with which he deals is a

Piestocystis stage, which thus agrees with the present species in

some other particulars, though, as already stated, I cannot

definitely refer the asexual stage to the Piestocystis type. The -

investigation of the younger worms allows of an explanation of

certain structural peculiarities in the older worm which I was

unable to understand while preparing my former paper. There

is, I think, no doubt that the central cavity is, as I then suspected,

an extension forward of the cavity of the bladder, which we now

know to be not limited to one dilated end of the body in the,

young of this species of tapeworm. The elongated worm-like

* * A Contribution to a further know ledge of the Cystic Cestodes.” Proc. Linn.

Soc. N.S.W. (2) ix. p. 60.

+ Ibid. pl. iii. fig. 6

12h DR. F. E. BEDDARD ON

asexual form is therefore in reality a bladder-worm, and not an_

adult in which sexual organs have not yet appeared.

Text-figure 7.

S.C.

Section through skin of the fully developed asexual worm.

C. Cuticle. s.c. Subeuticular layer below which are the saes containing

calcareous corpuscles.

Furthermore, it is to my mind evident that the ** closed cavities

of very problematical nature ” which occur in the fully-developed

sexless worm, which it will be remembered are not segmentally

arranged and which have a lining of cuticle and epithelial cells,

are the cavities lettered A and C in my figure (text-fig. 2) of the

young plerocercoids. In addition to these there are various

tubular cavities which seem to belong to the water-vascular

system, and which | observed to approach very closely to the

exterior of the body but not to open thereon. ‘These tubes,

present in addition of course to the four longitudinal trunks, are

exactly repeated in the younger stages that have been described

in the present communication. There is therefore no abrupt

break between the much more bladder-worm-looking embryos

MAMMALIAN CESTODES. 105}

and the completely formed sexless worm; the metamorphosis

is obviously a matter of gradual growth. The question next

arises as to what is the relation of the complete plerocercoid to

the adult tapeworm. Is the one converted into the other? To

settle this matter, or rather to contribute to its settlement, we

must again consider the buds produced by the sexless worm.

But I shall first make a

S Comparison of Immature Urocystidium (fully developed

Plerocercoid) with Cysticercus fasciolaris.

In my earlier paper upon this worm I compared these two

forms in certain details; but the description was concerned with

external characters alone. Now that I have been able to ascertain

some more facts about the development of the asexual form, a

more detailed comparison will be made. My information as to

Cysticercus fasciolaris is chiefly derived from Bartels’s memoir

cited below *. Although a primary difference of great importance

absolutely separates the two immature forms, it is rather remark-

able that the number of hooks in the Cysticercus fasciolaris seems

to be identical with that of the sexually mature Urocystidiwm.

Bartels gives 34-36 hooks, arranged, as they are in the present

species, in two circles. But though this statement is made in

the text, only 16 in one row are figured by him, and, moreover,

in two different figures 7. This is the number that is met with

in the adult Urocystidiwm. In the immature form, as I have

mentioned, there is no trace of hooks that I could discover. In

both forms the calcareous corpuscles are very numerous. But

while in the present species these corpuscles are most numerous

in the sexual form, the reverse is the case in Cysticercus fascio-

laris, the mature worm, 7'@nia crassicollis, having fewer calcareous

bodies. ‘The most noteworthy resemblance seems to me to lie in

the disposition of the bladder of the two worms. In the Cysti-

cercus the bladder is situated at one end of the body, and is very

smal] compared with the freeand segmented portion of the worm.

As I have already pointed out, precisely the same is the case with

Urocystidiwm. But in both cases the cavities found in the worm

are not limited to a restricted bladder. Cavities occur in both in

the segmented part; these are in Cysticercus mainly limited to

the posterior end of the body, but they are also found anteriorly ;

so that there is here no essential difference from Uvrocystidium.

Bartels remarks of these cavities that they are some of them con-

nected with the bladder-cavity while others are cut off from it.

He mentions that these cavities are sharply marked off from the

parenchyma in which they lie, but that they cannot be said

to possess definite walls. There is thus a greater resemblance to

the bladder-cavity of the buds of Urocystidiwm than to that of

* Zool. Jahrb. Bd. xxi. 1902, Abth. f. Anat. p. 511.

+ Loe. cit. taf. 39. fig. 21 & taf: 38. fig. 15.

14 DR. F, E. BEDDARD ON

the developing asexual worm which arises, as I suppose, directly

from the egg. There is, however, clearly a general resemblance

to the latter, and, as I think, an important resemblance.

The cavities also contain in both the immature Cestodes now

under consideration a coagulable fluid apparent on staining. The

tubes of the water-vascular system are much alike in the two

worms. In both the dorsal vessel is the smaller and has a thick

muscular wall; while the two worms also show resemblance

in the fact that the transverse vessels which run across the

segments arise by two origins from the larger ventral vessel.

This detailed likeness is very striking. In spite of these points

of likeness there are numerous points of difference between the

two species. Cysticercus fasciolaris would seem to show no such

specialisation of the bladder-cavities, or at any rate no such

variation of structure in the cavities found in its interior, as

does Urocystidium. The total absence of hooks, and indeed of a

well-marked scolex, in Urocystidiwm has been already commented

upon. The Cysticercus under consideration agrees with other

Oysticerct in differing absolutely from Uvocystidiwm in these

features. Dr. Bartels heads one section * of his memoir upon

Cysticercus fasciolaris as follows:—‘t In welcher Beziehung

stehen Cysticercus fasciolaris und Tenia crassicollis za eimander

betreffs der Hohe ihrer Organisation ?” The same query might

well be asked concerning Uvrocystidium. In the former case

the only differences are in the disappearance of the traces of

the bladder, the appearance of the gonads, and, apparently, the

slight inevease of the number of hooks upon the scolex. The

general level of organisation is upon precisely the same level

in the two stages of development. The latter statement can

well be made of Urocystidiwm. Transverse sections of the sexual

adult and the completely developed immature stage, which are

shown in my earlier paper upon this worm, leave little structural

difference between the two forms save the two features in which

the. Cysticercus differs from its Tenia. The only important

addition to be made is of course the absence of a scolex in

the young Urocystidiwum. 1t was, in fact, not merely the

equahity of organisation but also the details of likeness which

led me to assume originally that the one form of Urocystidiwm

was really a stage in the development of the other. So far

aso 1s known, these two forms are the only ones among the

higher Cestodes in which there is a high grade of organisation

in the ‘sexless stage. And it is perhaps this general fact of

likeness which tends to impress upon one the further points

of likeness between this form and Cysticercus fasciolaris, and

to diminish the really important points of difference already

referred to. As to further development, Bartels is of opinion

that the Cysticercus develops into the Zenia with no further

change than the loss of the bladder.

* Toe. oe p. 542.

MAMMALIAN CESTODES. 15

§ Comparison of the Young Plerocercoids with the Buds.

As has been already observed, there are no free living stages of

this worm in my possession which are younger than the specimens

described in the foregoing section. But it hasalso been suggested

as a possibility that the buds formerly described by me may be a

stage antecedent to those which I have termed the young plero-

cercoid. When freed from the fully developed asexual worm they

may drop off into the liver ducts and there increase in size—prin-

cipally in breadth, for the length of the longest buds is not far

removed from that of the smallest free plerocercoid—and, after a

' certain alteration in structure, show the characters of the maggot-

‘like larvee which have been dealt with in detail in the last few

pages. I do not think, however, that this can be the case, failing

at any rate further evidence. For the most “mature” of the

buds which I studied in some detail for my earlier communication

upon this genus has a much more “adult” structure than has

the youngest of the plerocercoids described in the present paper.

And if we are to assume that the young plerocercoids described

here develop gradually into the large sexless worm, as I think is

fairly certain, this possibility is rendered still less likely.

I shall now indicate the differences in structure between the

_ plerocercoids and the buds which oppose themselves to this com-

parison. The various stages in the development of the buds shown

in text-fig. 116 * of my earlier paper seem to show that the bladder

is the first part of the worm to appear and that the vermiform

_region is an outgrowth of this. For the buds are at first bladder-

like outgrowths of the stock, and then develop a process at the free

end which gets to be more important. Now in the plerocercoids

the whole worm is at first also little more than the bladder region ;

but this is converted into the more mature individual by a pro-

portionate suppression of the cavities which together constitute

the bladder-like region, and the gradual conversion of the greater

‘part of the worm into a more markedly vermiform body. Traces

of the bladder-cavities, however, persist in the latter up to the

anterior end. ' This difference of mode of growth does not, how-

ever, exhaust the differences observable between the two series

of young worms. The bladder-cavity itself differs in thetwo. As

already described, it has no proper walls in the free young plero-

cercoids, while in the buds it has not precisely a definite wall,

but the inner layer is thickened and it is sharply marked off from

the empty space which it encloses: it is, in fact, like some of the

cavities lettered C in the young free worms.

There is, however, the tube which I have lettered « in my

figure of a ‘transverse section through a bud}, which is in

structure very like the spaces lettered A in the transverse section

through the young plerocercoid (text-figs. 2, 3). The former is,

* P.Z.S. 1912, p. 824.

+ Ibid. text-fig. 116, p. 833.

16 DR. F. E. BEDDARD ON

however, distinctly tubular and definitely opens on to the

’ exterior, in both of which characters it differs from the cavities

described above in the young free larva. The water-vascular

system presents both similarities and dissimilarities in the two.

In the bud there are but two longitudinal vessels, one on each

side of the body. These are connected by transverse vessels and

are thus in all probability to be looked upon as ventral vessels ;

the dorsal, it is to be presumed, appear later. The youngest free

larva also possesses only two lateral I$ngitudinal trunks, which

are for similar reasons to be regarded as ventral vessels; but it

has in addition other water-vascular tubes which are completely

wanting in the bud. In fact, there are no detailed resemblances

such as would be sufficient to overbalance the general unlikeness

between the buds and the free living asexual parasite; they do

not appear to me to be stages in the same series.

§ Origin of the Sexual Form.

It seems to me to be much easier to believe, on the other hand,

that the buds area stage in the development of the sexual worm.

But here it is only a matter of possibility or probability; there

are but few facts. And these facts are mainly negative. For

there are actually no more positive points of likeness between the

bud and the sexual form than there are between the buds and

the fully developed sexless form; but perhaps the resemblances,

though few, are of a little more importance. In the first place,

the solidity of the body anteriorly to the bladder, save for the

one tube which runs along it, is more suggestive of the adult

sexual worm, than of the sexless worm with its abundant cavities

and continuous bladder-cavity. The gradual tapering off of the

single cell-lined cavity, which the buds possess, as it approaches

the anterior end of the body, indicates the ease with which it

may vanish altogether. In the second place, the bud shows no

series of peripheral tubes which I have regarded, in the case of

the young free worms, as a part of the water-vascular system.

These are absent from the sexually mature worm. Finally, there

are the aggregations of cells, of which I have spoken in my first

paper upon this worm and which may be the commencing sexual

organs.

This is the most important resemblance. The other features

in the structure of the bud do not forbid the view which I

am now supporting by such facts as are available. But they

might with equal reason be urged in favour of a likeness with

the free sexless stage.

There is obviously nothing decisive about the retardation in the

appearance of the smaller dorsal water-vascular tube, and the fact

that there are two layers of bundles of longitudinal muscular fibres

in the cortex points with equal directness to the adult worm and to

the complete immature worm. But the comparison here is with

the fully developed immature tapeworm, and not with the maggot-

MAMMALIAN CESTODES. 7

like earlier stages of this Gf I am right in so regarding them).

It would appear, therefore, that the facts which have just been

referred to in the structure of the bud may be perhaps really

looked upon as arguments in favour of a comparison of the bud

with the fully adult sexual form and a conclusion that the former

ultimately gives rise to the latter. For if we are to assume that

these buds give rise to the plerocercoid, it will have to be further

assumed that the regularly arranged and perfectly differentiated

cortex and medulla of the bud becomes lost, only to be recon-

structed later when the plerocercoid has arrived at the completion

of its development. Of the alternatives, this seems to me to be

‘the less likely. In short, I am obliged to admit that the develop-

ment of the bud into the sexual worm is mainly probable on

account of the difficulties which beset the view that it is an earlier

stage of the plerocercoid.

$ The Sexual Form.

The sexual form appears to differ in several particulars from

that of my first specimen of Urocystidium gemmipavum*, and I

have been able to compare the two placed side by side. It is to

be noted, however, that they agree with each other in differing

from the asexual worms in their browner colour. ‘This is perhaps

rather more strongly marked in the individual which forms the

subject of the present communication to the Society.

In the case of both worms, the segmentation begins immediately

after the scolex and the segments are quite narrow. ‘They get,

however, slightly longer at the posterior end of the body, and in

the new specimen the last three segments are comparatively long

though not nearly so long as broad. The body ended abruptly

with the last of these segments. The present worm contrasts

with the original specimen in its smaller size; it is of about half

the length, measuring some 42 mm., and is not nearly so broad,

the greatest breadth being 4 mm.

It is, however, in the scolex that the most prominent difference

between the two individuals is to be found. Of this region I

have had drawings prepared, which are shown in text-figure 8.

On a superficial view, with a hand-lens only, the scolex of the

second individual is seen to be much smaller than that of the

specimen which I described a year ago. ‘This difference is

heightened when the two are compared under a compound

microscope. ‘The upper right-hand figure in the illustration

(text-fig. 8C) shows that in the new specimen the scolex end is

terminated by a thin collar, in which a row of sets are imbedded

forming a complete circle. A good way below this are the

suckers, which are relatively small and separated from the hooks

. by a larger interval than that which occurs in the type example

of Urocystidium gemnuiparum. It might perhaps be held that

* Loc. cit. p. 841.

Proc. Zoou, Soc.—1914, No, II.

18 DR. F. E. BEDDARD ON

the differences between the two examples is due to the different

state of retraction of the rostellum in both. It might be said

that if we retract the rostellum of the type specimen, a collar

might be formed such as is shown in the second example sur-

rounding the depressed rostellum in which the hooks appear.

While this is quite possible theoretically, it may be pointed out

that the difference in size seems at first sight and without micro-

scopic investigation too great to allow of such a comparison. This

Text-figure 8.

A. Sexual individual. B. Scolex of same with rostellum retracted.

C. Anterior end of sexual individual described in P. Z. S. 1912, p. 841.

will readily be seen on an inspection of text-fig. 8, A & B, which

are drawn of the correct relative size. It may also well be that the

thin collar-like edge which bears the hooks in the fresh example

of Urocystidiwm is a permanent structure, and that there is

nothing to compare with it in the earlier example. But this

point cannot of course be settled in the absence of other specimens.

On the whole it would seem impossible that any differences in

MAMMALIAN CESTODES. 19

the state of contraction of the two examples could account for

the plain lack of resemblance shown in the various figures

referred to. Moreover, the suckers are very much smaller in the

new example of Urocystidiwm, and by no means so prominent as

are those of Urocystidium gemnuparum. So much then for the

scolex of this worm, so far as it can be seen by a mere inspection

of the uninjured worm. I have made a series of transverse

sections through the scolex after the above figures were drawn, in

order to elucidate further the structure of this part of the body.

In spite of the apparent differences enumerated above, I cannot

find any reason after a microscopic examination for regarding

this individual as referable to a second species of the genus. The

hooks are disposed in two rows, the hooks of each row alternating ;

there are sixteen to each row and those of one row are much

smaller. The apparently smaller size of the suckers is due to

their complete retraction. It is very important to note how in-

dividuals may appear to differ if not examined microscopically

and when in a different state of contraction.

In my original description of Urocystidium gemmiparum L

gave a somewhat detailed account of what I then supposed to be

ripe ova scattered thickly through the segments of the body, par-

ticularly towards the end of the body. I have re-examined the

original sections which I made, and have cut fresh sections from the

original material in order to consider the matter afresh. A part

of one of these sections is illustrated in text-fig. 9. It will be seen

that, as I have already reported, the medulla as well as the cortical

layers is stuffed with bodies closely resembling eggs. They are

large and lie loosely in the parenchyma, being often aggregated

into clumps. This section was regarded by two naturalists, to

whom I showed it, as being chiefly made up by these masses of

what appear to be eggs. I confirm my original description as

regards the structure of these bodies. They consist of a nucleated

cell closely surrounded by a transparent homogeneous layer,

which I noted as being the egg-shell. I did not see the nucleus

in all of the supposed eggs, which I carefully examined with

high powers. This matter [am able to correct by the subsequent

observations upon which I report here. The nucleus is present

in all of them; but often it is seen in various stages of degenera-

tion, culminating in freedom from granules, and thus almost

complete transparency. In the second example of this tapeworm,

the same cells were present and no particular description of them

is necessary. The occurrence of the same bodies in a second

individual led me to suspect that they were not eggs; for on the

hypothesis of a dicecious tapeworm it might be expected that a

male would be found. I had realised the likeness of these bodies

to caleareous corpuscles, but had abandoned that view in deference

_to their immense multitude, and if anything greater prevalence

of numbers in the medullary part of the body of the worm.

Moreover, it is usually stated that the nucleus of the cell in

which lime is deposited, and which becomes in consequence a

20 DR. F. E. BEDDARD ON

calcareous corpuscle, is excentric. This is pointed out by Benham *

and in the investigations of v. Janicki upon the development of

these bodies in the genus Davainea + On the other hand, it is

to be admitted that Lénnberg = found the nuclei to be often

centric as well as excentric in the Bothriocephalid Abothriwm

rugosum. There are, however, two strong arguments in favour

of regarding these bodies as calcareous bodies: these are, first,

that they bubble with gas on being treated with dilute hydro-

chloric acid; and, secondly, that I have found in this second

example of the worm rudiments of the real generative organs.

Text-figure 9.

Part ot a transverse section through sexual form of Urocystidiwm gemmiparwmn.

n. Nerve-cord. ¢.m. Transverse muscles. w.d. Dorsal water-vascular tube.

w.v. Ventral ditto. The medulla is crowded with darkly stained calcareous bodies

The second argument is I think conclusive as against the view

that the bodies in question are eggs. The reproductive organs

* ©A Treatise on Hashes; Oxford, pt. iv. p: 107 (1901

+ Arch. de Paras. t. vi. 1902, p. 261. ? .

+ K. Svensk, Vet.-Ak. Handl. Bd, 24, 1891, p. 78, pl. i. fig. 5,

MAMMALIAN CESTODES. 21

are in all the segments which I examined quite immature; but

the cord of cells representing the future ducts were quite visible

and were seen to lie always on the same side. This worm there-

fore appears to have unilateral generative pores. This discovery

renders the novelty of this worm as a genus rather more doubtful.

But the facts now known do not permit of a settlement of the

question. For the structure of the reproductive organs can alone,

at the present day, determine the systematic position of these

tapeworms. It is, as I think, unsafe to base generic identity

upon other characters in the absence of information about the

reproductive system. But it may be pointed out that there are

undoubtedly certain resemblances in the structure of the water-

vascular tubes of this worm and of Tenia crassicollis, which are

mentioned above * in comparing the young of the two worms.

But I do not think generic identity with Tenia proved.

§ Summary and General Considerations.

The fresh material reported upon in the foregoing pages

enables me to define more fully the structure and life-history

of the tapeworm which I described formerly as Urocystidium

gemmiparum t, from Fiber zibethicus.

As for the general structure of the fully grown asexual genera-

tion I have nothing to add to my former description, to which

reference may be made. The individual, however, which I have

described and figured in the present paper, differs from that

previously described in showing no development of buds. Two

earlier stages in the development of the asexual worm are

described in the present paper. ‘These are plump, short, but still

vermiform, and segmented, worms. ‘They differ from the full-

grown asexual worm: in the greater proportionate size of the

bladder-cavity and other cavities perhaps connected with the

bladder-cavity. But all of these cavities persist in the full-grown

asexual stage. ‘They show no hooks or recognisable scolex. In

the earliest stage there is no differentiation of the body-wall into

cortex and medulla; in the more advanced young there are

but feeble traces of this differentiation, quite achieved in the

completely formed asexual worm.

The structure of the asexual worm as elucidated by the

younger stages described in the present paper, shows that it is

not referable exactly to any type of asexual tapeworm as yet

described. In spite, however, of its external and internal (trans-

verse water-vascular vessels) segmentation, it is essentially a

bladder-worm, but, as it appears, without a scolex. In com-

plication of structure and in details, this asexual stage is not

far different from the sexual stage. The sexual worm, which

occurs in the same cavities of the liver side by side with

* Supra, p. 14.

+ It is to be borne in mind as possible, but not at all probable, that the two

sets of worms are of different species.

29 ON MAMMALIAN CESTODES.

the asexual stages, is of roughly the same size as the fully-

developed asexual stage. It has a very strongly muscular

rostellum and two rows of sixteen hooks each, the outer row

consisting of smaller sized hooks than the inner. The body of the

sexual worm is crammed with large oval calcareous bodies with a

centrally placed nucleus. Such bodies are also very abundant, but

not so abundant, in the bladder-worms. The gonads, etc. (quite

immature in both individuals) are fairly central in the pro-

glottids, and the ducts are all directed towards the same side of

the body.

It would appear, therefore, that both the structure of the

immature tapeworm and the series of stages by which maturity

is arrived at, are quite unlike anything that is at present known

among the Cestoidea. But as there are obvious lacune in the

information which has been set forth in the present paper, some

uncertainty attaches to the life-history the course of which

is suggested by those facts. There are, as it would appear,

two larval stages following each other and derived from each

other directly. From the egg (as I presume in the absence of

earlier stages) arises the larva which I have described as a plero-

cercoid ; this gives rise by budding to many larvee which differ in

several structural features from the asexual parent; these (and

here there are no positive facts but only inference) give rise to

the sexual worm. ‘There are thus three stages in the life-history

of this tapeworm which are not met with in other Cyclophyllidea,

except as mere multiplication as in Hchinococcus. Bothriocephalus

has three distinct stages, but they are not comparable to those

described here, since the first is a free-swimming ciliated larva

The complication of the life-history in this form is suggestive,

of course, of the Trematoda; but I can make no detailed

comparison.

ON THE “ ECLIPSE” PLUMAGE IN THE MALLARD, 23

2. Observations made to ascertain whether any Relation

subsists between the Seasonal Assumption of the

“ Helipse ” Plumage in the Mallard (Anas boscas) and

the functions of the Testicle*. By C. G. Srniamany,

F.Z.S., and 8. G. SHarrocr.

[Received December 20, 1913 : Read February 17, 1914. |

(Text-figures 1-6.)

INDEX.

Physiology and Variation.

The observations herein set forth were made with the object of

ascertaining whether any relation exists between the condition

of the testicle and the seasonal assumption of the eclipse plumage

in the male of the Wild Duck or Mallard (Anas boscas).

Although the seasonal changes are well known and were fully

described many years ago by Waterton, with whom the term

‘« eclipse ” originated 7, it does not appear that any observations

have been made on the condition of the testes accompanying

the change.

What we wished to determine was, whether the assumption of

the male plumage corresponded with the advent of spermato-

genesis, and whether the occurrence of the “ eclipse” is associated

with retrogressive changes in the sexual gland.

We may point out that the interest attaching to this question

is increased if the pairing habits of the wild duck and some other

common birds be considered. The cockerel of the common fowl,

for example, is sexually potent as soon as its external characters

are declared (so that poultry breeders are careful to separate

the cockerels from the pullets as soon as the former begin to

show male plumage), but in the pheasant, although full male

plumage is assumed in the autumn, no pairing (or any mani-

festation of the sexual instinct) takes place until the spring. In

the wild duck the birds pair in the autumn or early winter, after

the male assumes full plumage, but copulation does not occur

until the spring is advanced.

Our observations fall into two series :-—

1. Simultaneous observations on the plumage and the condition

of the testes in a series of Wild Ducks throughout the year.

2. Observations on Wild Ducks from which the testicles have

been removed.

* The expenses connected with this work were defrayed by a grant from the

Royal Society, London.

+ W. Yarrell, ‘ History of British Birds,’ vol, iii. p. 175,

24 DR. C. G. SELIGMANN AND MR. S. G. SHATTOCK ON THE

Observations on the plumage and the condition of the Testes.

These were made on domesticated wild ducks such as are found

in the London parks and supplied from game-farms. These

birds are commonly slightly larger than those really wild, but

they pass through the seasonal changes of plumage in approxi-

mately the same period, and appear to vary in the time (season)

of their change no more than do wild birds from England and

Scotland respectively. We were able to examine a few really

wild birds caught in decoys in January and February, and found

no substantial difference in the condition of their testes and that

of those domesticated, but we did not extend these observations,

since it appeared probable that the confinement of such birds

might lead to abnormalities in the onset of their plumage

changes, and even affect the condition of their reproductive

glands.

Before dealing with the changes in the plumage and in the

sexual glands of adult birds, it will be well briefly to summarize

the conditions found in young or immature birds, by which we

mean birds of the year, that have not yet assumed full plumage.

So defined, the period covers the first five to six months of the

bird’s life and ends in November or December, In the young

birds the nestling plumage persists until at least the middle of

September, about which time a few speckled feathers appear on

the legs and shoulders, the breast and belly being still unchanged.

By this time the full complement of flight-feathers has appeared,

but the feathers themselves are usually not more than half grown.

The testes of such birds are usually quite small, measuring on an

average about 10 mm. in length and about 2 mm. in breadth;

they are firm on section, and yellowish brown in colour. Under

the microscope the tubuli are found to be small, and with a

relatively narrow lumen lined with a single layer of cells, between

which certain larger spheroidal spermatogonia are intercalated.

In some the cells are two deep. The interstitial stroma is

very cellular, and, contrasted with the condition found.in the

fully functional gland, it appears relatively abundant *. There

is a good deal of individual variation in the external appearance

of the birds at this time, even a few days making a considerable

difference. The growth of the birds and the development of the

plumage proceed rapidly during October, so that by November

the drakes have attained then full size, and have assumed their

perfect plumage. The testes become larger and yellowish white

* We may refer here to the appearance presented by the testes of a freshly-

caught wild Mallard killed during the last quarter of December. This bird weighed

1 Ib. 142 ozs.; its testes, which were firm, yellowish in colour, and dry on section,

together weighed 88 mg: Microscopic sections showed the tubuli to be of small

size, with narrow lumen, lined with a single or double row of cells, and with larger

spermatogonia occasionally occurring between the basal cells. There was no sign of

spermatogenesis.

‘ROLIPSE” PLUMAGE IN THE MALLARD. 25

in colour, as well as softer, but no juice exudes on section, and

spermatozoa are absent. At this time of the year, 7.e.at the end

of November and December, there is no marked and constant

difference between the testes of the young birds of the year that

have but just assumed their winter plumage for the first time,

and the testes of older birds that are passing into their full

plumage for the second or third time. But since our observations

lead us to think that the changes in the testes may take place

rather more rapidly and regularly, and perhaps a little earlier, in

the older birds, we confined our observations to birds which had

passed through at least one full change of plumage.

In order to make clear the significance of the plumage-changes

in normal and in partially castrated adult. birds, recorded

below, we may give asummary of those which naturally occur in

the male wild duck.

Normally in the adult Mallard (Anas boscas), which, it is

assumed, has bred early in the spring, the curly tail-feathers are

lost, and the moult of the body-feathers begins late in May or

early in June. By the beginning of July the assumption of the

dusky, summer, or “eclipse” plumage should be tolerably com-

plete, though the moult of the flight-feathers has not, as a rule,

begun, these being lost usually by the middle of the month. The

eclipse plumage persists throughout August, during which month

the Mallard, the Duck, and the young are externally very much

alike. By the middle of September the curly feathers have

usually appeared in the tail of the Mallard, and the bird passes

from its summer to its winter plumage, which does not, however,

reach its full beauty until about midwinter.

The period of the year at which the plumage changes take

place varies with the latitude. In the British Museum (Natural

History) there is the skin of a bird in full male plumage with

curl feathers in the tail, which was killed at Shanghai (lat. 31° N.)

on August 28th, 1884. The skin of another bird, killed on

December 11th, 1879, at Nagasaki (lat. 33° N.), is in partial

eclipse, with many brown feathers on the vertex, and eclipse

feathers in the breast; while the skin of a third bird, from

Wuhu, on the Yangtsze, killed.in January 1885, has similar, but

less well marked, remains of the eclipse plumage on the breast

and head.

A Table showing the condition of Plumage and that of the

Testicles at each month of the year.

January 30th. Full winter plumage.

Weight of both testes, 3850 mg.:; each gland is about 23 mm. in greatest

length, of a yellowish-white colour and soft; fluid can be scraped from the

cut surface, but it does not ooze naturally.

(In a bird killed January 23rd, the testicular tubuli were small; with well-

defined lumen; cells about two deep; no spermatogenesis.)

26 DR. C. G. SELIGMANN AND MR. S. G. SHATTOCK ON THE

February 14th. Full winter plumage.

Testes soft, 15 mm. in length. Tubuli large and full of well-conditioned

cells, but there are no spermatozoa or spermatids. Small groups of finely

eranular interstitial cells are present.

February 14th. Full winter plumage.

Each gland is about 15 mm. long by 9 mm. in breadth, the tubuli are full of

cells but contain no spermatozoa.

March 7th. Full winter plumage.

Weight of both testes together, 5800mg. Testes equal in size, bean-shaped,

maximum diameter 25 mm. ‘Tubuli thickly lined with cells; a moderate

number of spermatozoa in some tubuli, 7. e. spermatogenesis is in progress.

March 16th. Full winter plumage.

Weight of testes 9150 mg. Tubules of full size, occupied by dense masses

of cells ; centrally there are numerous spermatozoa.

(The testes of another bird, killed on the same day, together weighed

10,170 mg.)

March 22nd. Full winter plumage.

Testes together weighed 32,500 mg. Hach was as large as a pigeon’s egg.

Tubules of full size; typical picture of active spermatogenesis.

April 20th. Full winter plumage.

Testes large, about 50 mm. long; and so soft as to be almost diffluent on

section: total weight, 30,140 mg. Spermatogenesis in active progress.

May 7th. Pull winter plumage.

Total weight of testes, 30,580 mg. Macroscopically and microscopically

they resemble those of the bird killed on April 20th.

May 30th. Full winter plumage.

Testes size of a haricot bean; the tubuli are distended with cells, and

contain large numbers of spermatozoa. The organ is, however, retrograding,

as appears from the absence of spermatid sheaves, and the fact that the

spermatozoa in the centre of the tubules are badly stained.

May 80th. Full winter plumage.

Hach testis, not larger than a small haricot bean, about 11 mm. long and

7mm. broad. Tubuli, not of full size; there is a central area of vacuolated

substance in which there are somewhat thinly scattered cells. No mitoses

and no spermatogenesis ; the whole appearance is one of inactivity.

June 6th. Full winter plumage.

Testes narrow and firm, about 13 mm. long by 4mm. broad. Tubuli of

somewhat small size, each with a peripheral layer of cells; the centre of the

tubuli occupied by a vacuolated material in which cells are thinly scattered.

There are no spermatozoa, but very fine deeply stained granules of chromatin

oceur in moderate numbers in the centre of the material (chromatolysis).

‘¢ RCLIPSE” PLUMAGE IN THE MALLARD. Did

July 4th. Bird in almost full eclipse.

Testes oval, about 6 mm. in transverse diameter, pale yellow in colour, and

firm on section. Tubuli small with well-defined lumen. Cells for the most

part two deep. No spermatogenesis.

July 8th. Full eclipse.

Testes small, scarcely 10 mm. long by 5 mm. broad. Tubuli of compara-

tively small size, furnished with well-developed lumen. Cells two and three

deep. No spermatozoa.

July 12th. Full eclipse.

Testes about 20 mm. long by 12 mm. broad; weight of each gland about

3700 mg. Tubuli of fairly large size, no lumen. Cells in the centre of the

tubuli are small. Here and there the sheaf arrangement of spermatids is

indicated, but the nuclei are mostly badly stained. No properly stained

spermatozoa,

July 12th. Full eclipse.

Testes resemble those of preceding ; no spermatozoa to be seen. Weight

of testes together, 6560 mg.

August 17th. Full eclipse.

Testes small, diameter of transverse section 4mm. Tubuli small, stroma

relatively large in amount. Tubuli furnished with well-defined lumen, cells

about two deep. No spermatogenesis.

August 21st. Hclupse.

Testes small, about 5 mm. in transverse diameter. Tubuli narrow, quite

inactive. A single layer of peripheral cells; a narrow lumen; no trace of

spermatozoa.

September 14th. Eclipse passing off.

Testes about 7 mm. in transverse section; weight 195 mg. Tubuli of

small size, with a narrow central lumen lined with one or two layers of cells ;

no signs of spermatogenesis.

(In a bird killed September 13th, the testes were 6 mm. in transverse

diameter ; the tubuli of fair size, and filled with loosely aggregated cells; no

spermatogenesis.)

September 30th. Eclipse passing off.

Many young feathers of winter plumage coming through on

breast and abdomen.

Testes small, firm, somewhat brown, about 12 mm. long and 3 mm. broad.

Tubuli of medium size, with central lumen ; lined with cells averaging two

deep.

September 30th. Full winter plumage.

Testes small, firm and somewhat brown, about 12 mm. long, but scarcely

3mm. broad. The tubuli are of medium size only, and are filled with cells.

October 9th. Full winter plumage.

Tubuli of medium size and filled with cells. Spermatogenesis is not in

progress, but amidst the central cells there appear here and there a few rod-

28 DR. C. G. SELIGMANN AND MR. S. G. SHATTOCK ON THE

like bodies, apparently obsolete spermatozoa, possibly the residues of a

previous state of activity.

October 25th. Full winter plumage.

Testes 15 mm. in chief diameter. Tubuli of medium size; furnished with

a layer of basal cells, central to which lies a mass of cells filling the lumen of

the tube. No spermatogenesis.

October 27th. Full winter plumage.

Testes about 15 mm. in longer diameter. Tubuli small, a narrow lumen in

most. Cells averaging two deep. No spermatogenesis.

November 1st. Full winter plumage.

Testes 12 mm. in longer diameter. Tubuli narrow, with fine lumen. No

spermatogenesis.

November 7th. Full winter plumage.

Testes about the size of a haricot. Tubuli of medium size; full of cells;

no lumen; no spermatogenesis ; some mitoses in the more central cells.

November 17th. Full winter plumage.

The testes fairly firm, exuding no fluid on section; together they weighed

5920 mg. Tubuli of medium size; fullof cells; nolumen; no spermatogenesis.

November 28th. Full winter plumage.

Testes yellowish; exuded no juice on section; weighed together 720 mg.

No spermatozoa. Tubulismall, well-defined lumen. Cells average two deep.

December 6th. Full winter plumage.

Both testes of medium size, and showing active mitosis.

From the foregoing table, which details the condition of the

testes In a series of birds examined throughout the year, and the

state of their plumage, the following summary may be made :—

The testes attain their maximum size during the breeding-

season, 2. ¢. at the end of March or beginning of April. At this

time each gland is almost as large as a pigeon’s egg, and so soft

as to be nearly diffluent on section, while the juice which exudes

contains enormous numbers of spermatozoa. Although this con-

dition is more or less maintained during the first half of May, by

the end of that month the glands present a very different

appearance. Although the birds are still in full winter plumage,

the testes have shrunken to the size of a haricot bean; no mitotic

figures are encountered in the cells of the tubuli, and spermato-

genesis has ceased ; or, if any spermatozoa are to be found, these

are badly stained and obsolete. During June the glands become

smaller and firmer, and the whole microscopic picture is one

of inactivity ; they diminish still further in size during July and

August, and acquire a yellow or brownish colour. This condition

persists throughout September, during which month the bird

“ BCLIPSE” PLUMAGE IN THE MALLARD. 29

puts off the eclipse for the winter plumage. During October and

November, when the brilliant plumage is fully declared, the testes

increase slowly in size, although they remain of a yellow or

brownish colowr, vetain their firmness on section, and eaude no

fluid when incised. Spermatogenesis does not commence until

December (perhaps the end of November) ; and the testes at first

do not greatly increase in size. There is some variation as to the

precise time at which spermatogenesis begins, but in any case

the bulk of the testes greatly augments during the latter half

of January and February. Evolution proceeds more and more

rapidly during the second half of February, and March, until by

the end of the latter month, or during April, the glands attain

their maximum size.

It must be understood that in the above account we have

endeavoured to give an average picture of the annual evolution

and involution of the testes, but there is no doubt that a con-

siderable range of variation obtains in the time of the onset of

spermatogenesis. Furthermore, in some birds, spermatozoa may

be found in the testes at a somewhat later date than is usual;

in such circumstances it is probable that spermatogenesis arose

correspondingly late.

IT,

Observations made to study the results of Castration

upon the Plumage.

It was our intention to castrate a series of ducks in order to

see whether any aberrations would result in the natural history

of the plumage, but in spite of repeated attempts, it was found

impossible to completely extirpate the glands. Every visible

trace might be removed, after an extensive laparotomy, and yet,

when the bird was killed some months later, a greater or lesser

amount of regenerated testicular tissue was found either in the

normal position or engrafted upon the neighbouring organs. The

castration in every case was carried out under an anesthetic.

The following experiments should all be read in the light of this

fact, viz. that some regeneration of testicular tissue must have

taken place within a few weeks or months of the operation ; and

since we do not know how small an amount of this tissue may be

sufficient to exert an action on the body, we cannot say whether

the results observed within the first few months of the operation

would have persisted had the castration been complete. There

are, however, certain considerations which lead us to believe that

our results, in so far as they apply to the first six or eight months

after the removal of all visible testicular tissue, are essentially

comparable with those which would occur during this time were

it possible to obviate the partial regeneration of the gland. In

the first place the delay in the assumption of the eclipse in the

birds submitted to operation from December to April (when the

plumage is masculine), indicates that the removal of all visible

testicular tissue has produced a definite effect on the bird;

30 DR. C. G. SELIGMANN AND MR. S. G. SHATTOCK ON THE

whereas castration performed in July, while the testicles are

retrograding and the bird is in eclipse plumage, produces

no corresponding delay. Furthermore, the small size of the

“ ovafts,” or regenerated tissue, in birds killed from nine to ten

months after operation, would indicate that the amount of

testicular tissue, regenerated, and exerting its influence on the

bird during the early months after the operation, must have

been extremely small.

Our observations fall into two series according as the testes

were removed, (A) while the bird was in full winter plumage, or

(B) during eclipse.

(A.) The Results of Castration when performed on Birds

whilst in full Winter Plumage.

No. 18.) (Text-fig. 1.) Bird in full winter plumage; castrated in

December 1906. The testes removed showed active mitosis and

spermatogenesis in many tubuli.

June 9th, 1907.—Bird in full plumage, with glossy vertex, and

one very glossy feather in tail*. The only sign of approaching

eclipse 1s a slight brownish-yellow tinting of some of the speckled

feathers on the belly.

July 11th, 1907.—Full plumage unmodified except that there

is a diffuse light brown coloration of the abdomen: this is not

due to any growth of new feathers but to a pigmentary modi-

fication in the old. There are two curl feathers in the tail and a

third forming.

July 28th, 1907.—The general appearance of the bird is still

that of a male in full plumage; and the white ring on the neck

is as marked as ever it was, but there are a few brown feathers

on the cheeks (first noticed on July 23rd), and a strong flush of

eclipse feathers in the maroon area on the breast, which is

becoming somewhat lighter in colour. A number of eclipse

feathers are obvious upon the abdomen; these are mostly old

feathers in which pigment changes have taken place; the great

majority of feathers on the abdomen are still vermiculated.

There are a few new eclipse feathers in the scapular region. The

tail contains two curl feathers and one partially curled.

August Ist, 1907.—Kclipse progressing very slowly ; the head

is in much the same condition as it was when last noted. The

breast has perceptibly lightened, light brown feathers barred with

black alternating with the dark maroon feathers; a few typical

eclipse feathers are coming through the down of the breast.

The scapulars have somewhat darkened, and there are more

barred feathers upon the flank, each showing up as blotched

with dark, upon the grey vermiculated background of the older

feathers. ‘These dark feathers appear to be new. ‘Tail as last

deseribed. Although grey vermiculated feathers predominate

upon the abdomen, they are intermixed with a number of

brownish feathers with black centres: some of the latter are

new; others are due to pigment changes.

* On May 14th there were three curl feathers in the tail.

“ BCLIPSE” PLUMAGE IN THE MALLARD. 31

Text-figure 1.

The Mallard (No. 18) as it appeared on August 27th, 1907; castrated in

December 1906.

The eclipse is far from complete, z. e. it has been delayed by the castration; the curl

feathers have not as yet been lost, nor has the white ring on the neck fully

disappeared, etc.

Text-figure 2.

€ RES

Normal Mallard in full winter, or non-eclipse plumage.

ay DR. C. G. SELIGMANN AND MR. S. G. SHATTOCK ON THE

August 11th, 1907.—Gloss on head limited; small uvegular

areas of it on sides of face and neck; white ring faintly repre-

sented at sides, absent in front and at back. The eclipse

plumage is spreading rapidly upon the breast; this does not,

however, present a typical appearance, although most of the

feathers are brown with black centres, and the new ones coming

through the down of the breast are of the eclipse type. There is

no new flush upon the abdomen, though there are many brown

feathers upon the flanks, where vermiculated feathers still

prevail; it is not clear whether these brown feathers are new

or not. In both wings the primaries, secondaries, and tertiaries

are being rapidly moulted. Curly feathers are present in the

tail.

August 18th, 1907.—No flight-feathers remain in either wing.

August 24th, 1907.—The eclipse on the body is not obviously

proceeding. The curly feathers are still retained in the tail.

September 1st, 1907.—There is still a good deal of gloss on the

vertex and the nape; the white ring is slightly indicated at the

sides of the neck, not at the back or front. The breast is in

almost full eclipse; the change is certainly largely due to the

appearance of new feathers, but in part seems to be due to pig-

mentary change in the old. The feathers of the abdomen are,

some eclipse, some vermiculated; there are feathers of both kinds,

including a plentiful flush of young vermiculated feathers, on the

flanks. ‘There are no curl feathers in the tail, though these were

present five days ago. The two central feathers are dark and

glossy, and are beginning to ridge; the other tail-feathers are

glossy, as they always have been.

September 12th, 1907.—The general aspect of the bird is inter-

mediate; the majority of the breast-feathers that are fully

grown are eclipse; there is, however, an abundant flush of young

winter feathers coming through, although these do not yet affect

the colour of the plumage. The vertex and the nape are glossy,

but the cheeks present only a very few flecks of gloss; there is

no trace of the white ring on the neck. Most of the feathers on

the abdomen and posterior part of the breast are vermiculated ;

practically these are all old, but among them there is an abundant

flush of young vermiculated feathers coming through : the eclipse

feathers are few in number. On the flanks the feathers are

almost entirely vermiculated, but many of them are tipped with

brown, including a number of old (eclipse) feathers ; new vermicu-

lated feathers are coming through the down. The tail-coverts

and back are glossy, with abundant new tail-coverts coming

through. There are no curly feathers in the tail, but the central

feathers are ridging and becoming glossy.

September 21st, 1907.—The general appearance is that of a

bird in winter plumage, with the exception of the head and

breast, for the majority of the breast-feathers are of eclipse type,

in spite of a flush of winter feathers coming through and a con-

siderable number of winter feathers which have already expanded,

“* RCLIPSE” PLUMAGE IN THE MALLARD. 33

The head has only a slight amount of gloss; the cheeks are only

slightly flecked with the same. The feathers of the abdomen are

predominantly vermiculated, a few eclipse feathers being left

among them. The tail has one good curl feather and another

ridging.

September 25th, 1907.—The head and neck are almost entirely

glossy ; the white ring is appearing, but the feathers of the breast

are still largely of eclipse pattern, though many young feathers

have come through. Among the vermiculated feathers of the

abdomen a few eclipse ones are still present; practically all the

young winter feathers have come through. The scapulars are

unaltered. There are two curl feathers in the tail, and a third

is ridging ; the tail-coverts are glossy green, more so than when

last described.

November 12th, 1907.—The bird is in full male winter

plumage, with good curl feathers in tail. The only trace of the

eclipse is to be found on the flanks, where there are still a few

feathers tipped with brown and a few brown feathers which are

not vermiculated at all. This bird was allowed to live until the

end of July 1908, by which time, had it been a normal bird, it

should have been in full eclipse, or even have been passing out

of it.

The following note was made on July 27th, 1908 :—The bird

is in full eclipse, though there is a slight gloss upon the vertex.

The breast is in eclipse, but there are a good many partially

vermiculated feathers in the lower part of the breast and on the

abdomen ; there are no young feathers coming through. There

are no curl feathers in the tail, but the two central are becoming

glossy and beginning to ridge. he primaries and secondaries of

the wing have been shed; the young feathers are not fully

developed ; the feathers of the flank are vermiculated, mixed with

eclipse. The bird was killed and examined on July 30th, 1908.

On the right side there was a series of loosely attached nodular

grafts, which in total volume are about the size of a haricot ;

they were of a dull yellow, and brown on section. On the left side

there were two small nodules, which, together, are the size of a

small pea; there are other grafts, viz. at the root of the liver and

attached to the mesentery; both these are of the size of small

peas. All the testicular tissue is of a dull yellow colour.

(No. 13.) (Text-fig. 3, p. 35) Bird in full winter plumage;

castrated December 20th, 1906. Thetestes removed were brown

in colour, and did not exude fluid on section. Microscopically

the tubules have a wide lumen and a many layered lining, with

smaller cells about the centre. There are no spermatozoa, but

mitotic figures occur in certain of the cells.

July 11th, 1907.—The general appearance of the bird is that

of a male in almost full plumage. The vertex is dark green,

though not definitely glossy ; the cheeks are flecked with brown,

but the white ring on the neck is perfect. There is a slight

Proc. Zoou. Soc.—1914, No, ITT. 3

34 DR. C. G. SELIGMANN AND MR. S. G. SHATTOCK ON THE

lightening of the lower breast-feathers, and the feathers of the

abdomen are generally somewhat browner and less grey than in

the male in full plumage. There are four perfect curl feathers

in the tail; the wing-feathers have not been shed, and are still

quite firm.

September 11th, 1907.—Vertex dark, slightly glossy; only a

few flecks of gloss on the cheeks ; the white ring round the neck,

just beginning to show after having disappeared. The breast 1s

in eclipse; there is a moderate flush of young eclipse feathers

coming through. The abdomen is in eclipse. The flank-feathers

are almost entirely eclipse, but a few are in part vermiculated.

There are no curl feathers in the tail (text-fig. 4).

January 24th, 1908.—Although the general appearance of this

bird is predominantly that of a male in winter plumage, there

are many signs that the eclipse is only slowly passing off. The

vertex is dark, but it can scarcely be described as glossy, though

the rest of the head is generally so, with only a few brown

feathers remaining. The white ring, however, round the neck

has now become well marked. There is much eclipse plumage

left on the breast and abdomen. The flanks are in partial eclipse,

a number of incompletely vermiculated feathers being present.

The tail contains one poorly developed curl feather; the three

other central feathers are dark and beginning to ridge—the first

stage in the formation of the curl.

(No. 2.) Bird in full winter plumage ; castrated December 20th,

1906. The testes removed were brownish yellow, and did not

exude fluid on section. The tubuli were full of cells, and though

no spermatozoa were present there were abundant mitoses.

July 11th, 1907.—Although the general appearance is that of

a bird in winter plumage, it is modified by a considerable flecking

with brown feathers on the sides of the face, and by the occur-

rence in the anterior portion of the grey of the breast, of indi-

vidual feathers of a dull black edged with brown. The vertex is

glossy, and there are four perfect curl feathers in the tail.

The photograph of this bird (text-fig. 5, p. 36), taken on

September 11th, 1907, shows that the assumption of the winter

plumage was delayed. The plumage is predominantly eclipse.

There is no gloss on the head, and although the central tail-

feathers are beginning to ridge, by far the greater number of the

feathers on the head, belly, back, and shoulders exhibit wholly

or in part the dusky coloration of the eclipse.

September 11th, 1907.—The vertex is dark and only slightly

glossy ; a few flecks of gloss are present upon the cheeks. There

is no white ring on the neck. The feathers on the breast are

predominantly eclipse, but here, and on the abdomen, there

are feathers which are partially vermiculated, and there is an

abundant flush of fresh vermiculated feathers coming through ;

among them there are a few eclipse feathers. The flanks contain

many eclipse feathers and some new vermiculated ones. There

‘¢ HCLIPSE” PLUMAGE IN THE MALLARD, 35

Text-figure 3.

E x we

Mallard (13), photographed in July 1907; in almost full winter plumage.

The bird was castrated in December 1906. The eclipse has been delayed.

Text-figure 4,

The same Mallard as shown in text-fig. 3: photographed September 11th, 1907.

The bird was castrated in December 1906. The advent of the eclipse in the summer

of 1907 was delayed. The plumage is now largely of the eclipse kind; the

bird was not fully out of eclipse in January 1908, before which time it should

normaily have been in complete winter dress.

ca)

36 DR. C. G. SELIGMANN AND MR. S. G. SHATTOCK ON THE

is one glossy curl feather in thé tail, probably new; two other

feathers are beginning to ridge. Tail-coverts glossy.

Text-figure 5.

Mallard (2), photographed September 11th, 1907.

The bird was castrated in December 1906. The advent of the eclipse in the summer

of 1907 was delayed. The photograph shows that the passage from the

eclipse to the winter plumage is also delayed; the plumage is still pre-

dominantly eclipse; the vertex was only slightly glossy; the white ring has

not reappeared on the neck; there was but one curl feather in the tail.

January 24th, 1908.—Bird completely male; four good curl

feathers in the tail. On the flanks there is an occasional feather

of a dusky brown colour with obscure vermiculations.

This bird was killed on February 19th, 1908, when a spheroidal

graft, 6 mm. in diameter, was found in the abdomen. The tubuli

of the graft were of full size, and active spermatogenesis was in

progress.

(No. 48.) Bird in full winter plumage; castrated March 6th,

1907.

May 14th, 1907.—Bird in full plumage, with three curl feathers

in the tail. The only premonition of the eclipse is a slight

browning of some of the interscapular feathers and those upon

the lower part of the breast.

July 11th, 1907.—The plumage has undergone but little

‘“¢ROLIPSE” PLUMAGE IN THE MALLARD. 37

change, though the lustre has partly disappeared from the vertex,

which is dark; and there is a little lightening of the lower

chestnut feathers of the breast, which are tipped with white.

There are three perfect curl feathers in the tail, and the old

wing-feathers are quite firm and exhibit no tendency to be shed.

August 18th, 1907.—Much of the gloss on the vertex, and

some of that on the cheeks, is retained; the breast is partly in

winter, partly in eclipse plumage; the abdomen is in full

winter plumage. There are some dark feathers on each flank.

September 11th, 1907.—The general aspect is that of a bird

not quite in winter plumage; the head and neck are partially

glossy, though there is still a good deal of brown upon the cheeks.

There are some eclipse feathers in the breast, but the greater

part are vermiculated and more winter feathers are coming

through the down. The abdomen and the posterior part of the

breast are completely vermiculated. There is one curl feather in

the tail, and the tail-coverts are glossy. This bird was killed on

September 12th, 1907. It presented a single graft the size of a

small haricot: the tubuli were large and full of cells; in every

tubule spermatogenesis was in active progress.

(No. 7.) Bird in full winter plumage; castrated April 5th,

1907. The testes were large and very soft, as when in full

activity during this month.

On April 21st it was noted that there was a brownish wash-

like tinting of feathers on the lower breast and abdomen.

July 27th, 1907.—Eclipse not complete. Some gloss on the

vertex and cheeks ; the white ring has disappeared. The chestnut

area of the breast is only partially in eclipse. A number of eclipse

feathers are coming through, though many look vermiculated.

A large number of vermiculated feathers persist in the flanks,

and there are many new vermiculated feathers in this position.

Vermiculated feathers are coming through at the bases of the

wings. The hinder part of the breast is greyish rather than

vermiculated ; no new feathers are coming through here. There

are no curl feathers in the tail; the central feathers are ridging.

The wing-feathers do not appear to have been as yet shed.

November 30th, 1907.—Bird in full male plumage ; three curl

feathers in the tail. The bird was killed and examined. There

were a few encapsulated blood-clots about the site of operation,

but no trace of the testes except two nodules situated close

together and each the size of a millet seed. There were no

nodules on the intestines or liver. Microscopic examination of

the nodules referred to proved that they consisted of testicular

tissue, some of the tubuli of which were distended with cells.

No spermatozoa were present, and the central cells of the larger

masses were degenerated.

38 DR. C. G. SELIGMANN AND MR. S. G. SHATTOCK ON THE

(B.) The Results of Castration when performed on Birds

whilst in Eclipse Plumage.

(No. 11.) Bird in almost full eclipse plumage; castrated in

July 1907. The head was in full eclipse, except that a few

feathers on the side of the face showed a greenish gloss. The

whole of the breast was in complete eclipse, with young eclipse

feathers coming through in the anterior part. There were a few

vermiculated feathers at the base of the neck behind. The

primaries and secondaries had been shed from both wings, and

there were no curl feathers in the tail, though a little gloss

remained on the tail-coverts. The testes removed were about

22mm. by 8 mm. in diameters, yellow in colour and firm on

section.

September 17th, 1907.—The head has passed into almost com-

plete winter plumage, although a few brown feathers are still

present. The upper part of the chestnut area of the breast is in

full winter plumage, though a considerable number of eclipse

feathers are still present in the lower part of it. The grey

portion of the breast and the abdomen are almost entirely

vermiculated, though a few eclipse feathers are still present.

Over the area plucked for operation in July the feathers are only

faintly vermiculated and are ofa greyish brown. There are two

good curl feathers in the tail.

November 8th, 1907.—The bird was in full winter plumage,

with four curl feathers in the tail. It was now killed. A graft

the size of a small haricot was found at the site of the right

testis.

(No. 12.) Bird in almost full eclipse; castrated July 1907.

The testes were about 22 mm. by 8 mm. in diameters; pale

yellow in colour and firm on section. The head is in almost

complete eclipse, only the slightest glossiness persisting at the

vertex. The chestnut area of the breast is in complete eclipse,

with a few eclipse feathers still coming through the down; the

rest of the breast is predominantly eclipse, although a few of

the old vermiculated feathers persist. On the flanks there are

both vermiculated and eclipse feathers. The wing-feathers have

not been shed; there are no curl feathers in the tail. A good

deal of gloss persists upon the upper tail-coverts and the feathers

of the saddle.

September 17th, 1907.—The condition of this bird resembles

that of the preceding (No. 11), with the following exceptions :—

The head is not quite so advanced towards winter plumage, and

the ring on the neck is not so well marked; there are fewer

eclipse feathers in the lower part of the chestnut area; and there

are three curl feathers in the tail.

November 8th, 1907.—Bird in full male plumage, with four

curl feathers in the tail. It was now killed; at the site of the

right testis there were three small grafts, one about the size of

“CRCLIPSE” PLUMAGE IN THE MALLARD. 39

a carraway seed, and two others about the size of a grain of

maize and of a hemp seed respectively.

(No. 19.) Bird in full eclipse; castrated July 12th, 1906.

The testes removed were of medium size, 15 mm. in longer

diameter; the weight of each was about 3700 mg. The tubuli

were fairly large, and distended with cells, without lumen. Here

and there the spermatid sheaf arrangement was indicated. There

were, however, no properly formed and stained spermatozoa.

September 11th, 1906.—Bird in almost full male plumage,

except that the stippling on most of the vermiculated feathers

has not attained its full darkness, and that the majority of these

feathers are tipped with white.

January 24th, 1907.—Bird in full male plumage; there are

two curl feathers in the tail.

April 4th, 1907.—Bird in full male plumage.

July 27th, 1907.—Bird in almost full eclipse, though there is

a slight gloss on vertex. ‘The grey upper portion of the breast

is in eclipse, which must, however, be considered to be passing

off, since there are many winter feathers coming through the

down. The abdomen and the hinder part of the breast, eclipse.

The wing-feathers have been moulted, but the new primaries and

secondaries are not yet fully grown. There are no curl feathers

in the tail.

The bird was killed on July 30th, 1907. On the left side

there was a nodule the size of a small haricot, loosely connected

with the great veins below the normal site of the testis. A group

of grafts. together as large asa filbert, was loosely attached to the

back of the liver.

(No. 13.) Bird in full eclipse; castrated July 8th, 1907. The

testes removed were quite small, scarcely 10 mm. long by 5 mm.

broad. The tubuli, of comparatively small size, were furnished

with a well-differentiated lumen ; cells two or three deep: no

spermatozoa.

September 17th, 1907.—The bird still retains much of the

eclipse plumage; there are some glossy feathers on the vertex

and round the eye, but the rest of the head and neck are in

definite eclipse, and the neck shows only the faintest remains

of a white ring. The breast and the abdomen are in eclipse,

though in both positions a few vermiculated feathers are coming

through. The area which was plucked for the castration is

slightly darker than elsewhere. On the flank many of the brown

eclipse feathers are becoming vermiculated, and new vermiculated

feathers are, in addition, appearing. ‘There are no curl feathers

in the tail; the moult of primaries and secondaries in the wings

has been much delayed, as the new feathers have not yet attained

their full size, and their shafts are still vascular.

November 18th, 1907.—The head is glossy, but many feathers

on the cheeks are still partially brown; the white ring on the

40 DR. C. G. SELIGMANN AND MR. S. G. SHATTOCK ON THE

neck is well developed. The chestnut area of the breast contains

so many eclipse feathers that the general appearance of this part

is eclipse, but there are also a few winter-plumage feathers inter-

mixed. The lower part of the breast, though generally vermicu-

lated, shows a small number of eclipse feathers. The area on the

abdomen plucked in July for castration is covered with feathers,

darker than elsewhere and only partially vermiculated ; it might

perhaps be described as ‘‘semi-eclipse” in character. A few

eclipse and partially eclipse feathers persist in the flank. The

tail contains two good curl feathers, and two others are ridging ;

all are glossy, as well as the tail-coverts. ‘The bird was killed on

this day. On the left side there was no trace of testicle, but on

the right there was a nodule the size of a large haricot at the

site of the gland, and below this a second, no larger than a millet

seed.

Summary of the Effects of Castration upon the Plumage.

If we take the condition of No. 18 for the seven months

succeeding the castration carried out early in December 1906,

whilst the bird was in full male plumage, it appears that at the

end of July 1907 (when the normal mallard has been in eclipse

for some weeks) this bird still remained in almost full winter

plumage. A careful water-colour drawing, made on July 23rd,

1907, by My. Norman H. Hardy, shows a bird in complete

winter plumage, except for some small areas of brown on the

cheeks and round the eyes, and a slightly diffuse tinting with

brown of some of the vermiculated feathers on the abdomen.

This bird, however, had passed through its eclipse, and had

reassumed the full male plumage by September 21st. In July

1908, 2. e. the following year, it passed into eclipse like a normal

bird. The record of No. 13, castrated during the latter half of

September, when the bird was in full male plumage, shows a

similar result; but in this case the delay in the appearance of

eclipse feathers was even more marked than in No. 18; while in

both the birds not only was the eclipse delayed, but it was

incomplete.

The behaviour of No. 48, castrated in March 1907, resembled

that of No. 18, though the delay in the onset of the eclipse was

not so marked ; in No. 7, the testes of which were removed in

April 1907, there was comparatively little delay.

Whilst there can be no doubt that the removal of the testes

during the period of the eclipse does not retard the assumption

of the proper male or winter plumage; castration carried out, on

the contrary, whilst the bird is in full male plumage delays the

appearance of the eclipse. This delay is not so much a positive

delay as an abnormal persistence of the winter plumage. For if

the winter feathers be plucked (7. e. artificially shed), the new

feathers that replace them are of the typical eclipse kind.

Text-fig. 6 shows a bird concerning which it is no exaggeration

‘“RULIPSE” PLUMAGE IN THE MALLARD. Al

to say that half the area on the breast and abdomen, which

is normally vermiculated, exhibits the eclipse plumage, while

the rest is in male or winter plumage. This bird (No. 75) was

castratedin May. In the middle of July its general appearance,

whilst standing, was quite male, for the only obvious sign of the

oncoming of the eclipse was a slight loss of the gloss on the

vertex of the head and a browning of the cheeks; the curl

feathers in the tail were still unshed—. e¢., the appearance of

the eclipse was delayed, as usual, after castration. But on

lifting up the bird, it was seen: that the abdomen and breast,

where they had been plucked for castration in May, were covered

with a thick growth of buff and black eclipse feathers. Although

this is the most marked example of the growth of eclipse feathers

in plucked areas of castrated birds otherwise in winter plumage,

we have seen the same thing in a less degree in other cases.

Text-figure 6.

A Mallard (75) castrated in May.

The abdomen and lowest part of the breast (plucked under ether anesthesia for

castration) have become (middle of July) covered with a growth of buff and

black eclipse feathers, the plumage being otherwise of the winter kind, 7. e. the

appearance of the eclipse elsewhere has been delayed.

The ducks distinguished as Nos. 18, 48, and 69 were killed

during late summer or early autumn, 7. ¢. at a time when the

testes are normally in an inactive condition. We unfortunately

omitted to examine the histological condition of the “ grafts” in

No. 18, killed July 30th, 1908; but in the case of No. 48, killed

42 DR. C. G. SELIGMANN AND MR. 8S. G. SHATTOCK ON THE

on September 12th, 1907, spermatogenesis was in full progress

in the graft, a condition of activity which does not occur under

normal conditions at this period of the year.

In the case of two additional castrated birds (not further re-

corded in this paper) which were killed between the end of June

and September 1907, spermatogenesis was likewise taking place in

similar grafts : 7. e., at a time when in normal birds the testes are

functionless, at least as regards their external secretion. The

histological condition found in the grafts in these different birds

is shown in the following table :-—

(No. 8m.) Killed: June 1907 (late).

Condition of grafts :

One graft about the size of a small gooseberry ; the tubuli are of large size

and distended with cells; mitotic figures fairly abundant. In the centre of

one tubule is a group of deeply-stained filaments with bulbous ends, which

must be considered spermatozoa.

(No. 39.) Killed: September 12th, 1907.

Condition of grafts :

One graft about the size of a large pea. This consists of the epididymis

and testicular tissue, the tubuli of which are of full size; spermatogenesis is

in active progress.

(No. 48.) Killed: September 12th, 1907.

Condition of grafts :

One graft the size of a small haricot. The tubuli are large and distended

with cells; in every tubule spermatogenesis is in full progress.

(No. 22 bis.) Killed: Mid-September 1907.

Condition of grafts :

Graft consists of closely-set tubuli of large size and fullof cells. Spermato-

genesis with well-developed spermatozoa present in some of the tubes: in

other tubuli the centre is filled with a vacuolated mass of cell-débris

without spermatozoa.

Conclusions.

I. In the male of the Wild Duck the testes undergo annually

a series of seasonal changes (as in many other birds), and are

spermatogenic only during the winter months and early spring.

Il. The periods of activity and non-activity do not coincide

with the two seasonal changes in the plumage.

ITI. The normal passage of the bird from full winter (breeding)

plumage to its dusky summer (eclipse) plumage is, however,

delayed if castration is carried out during the months whilst the

gland is assuming, or has attained, its activity.

One bird (No. 18) which was castrated in the winter, and in

which the advent of the succeeding eclipse was delayed the

following summer, was kept until the summer of the next year.

“RCLIPSE” PLUMAGE IN THE MALLARD. 43

In this case the second eclipse occurred at the normal period.

As, however, small nodules of regenerated testicular tissue

were found at the autopsy (as indeed they were in every other

case), we are not at present in a position to say whether a

Mallard which is absolutely without testicular tissue will con-

tinue to pass through the same seasonal changes of plumage as

the normal bird.

It is a remarkable fact that the grafts were fully spermatogenic

in the month of September, an occurrence altogether abnormal

in the testicle of the entire bird. We can offer no explanation of

this anomaly.

The delay above referred to has its parallel in the well-

established fact that if a colt is castrated when shedding its

winter coat, the shedding is for a time arrested, and then

proceeds only very slowly.

It is of interest to observe here that in the case of the Wild

Duck, when females assume the male plumage (a phenomenon

well known also in the Common Pheasant and other birds), the

spurious males undergo the seasonal eclipse, although this is some-

what incomplete and aberrant.

IV. Removal of the testes during the eclipse does not produce

any constant, appreciable effect upon the next passage of the bird

into winter plumage.

It would appear from these observations that the seasonal

change of plumage in the Mallard is not connected with the

spermatogenic function of the testicle.

But whether a second function of the organ, viz. the production

of an internal secretion, or hormone, is responsible for the change,

could only be proved by castration so effectively carried out as to

exclude absolutely any reproduction of testicular tissue.

The only method of ensuring this is to reopen the abdomen

after the castration, and remove any nodules of reproduced

tissue. Our results in this direction we may lay before the

Society on some future occasion.

HOP Waee Fe irre ee : Vie

- 2 MARS ERIE: Ae ! PUP

IL,

1914, Meek. PI.

leah Dupre:

SPERMATOCYTE CELLS IN VARIOUS ORGANISMS,

SIP IRIAN OC Ws, Cia IGILS

Pe. Z. 8, WIa&, Meelk, Pi.

VARIOUS ORGANISMS.

WM.

ON SPINDLE-LENGTH AND CELL-VOLUME. 45

3. The Possible Connection between Spindle-Length and

Cell-Volume. By C. F. U. Murex, M.Sc., F.L.S.,

F.Z.S.

[Received October 31, 1913: Read February 17, 1914.]

(Plates I. & II.*)

INDEX.

Cytology.

Introduction.

I have stated that in Porficula auricularia and Helix pomatia

the ratio between the lengths of the mitotic spindle at the

conclusion of the two spermatocyte metaphases is identical,

or almost identical, with the ratio between the radii of two

spheres of which the volume of one is equal to twice that of the

other; and the same ratio has been observed by von Winiwarter

in the spermatocyte metaphases of Man. Since each primary

spermatocyte divides to form two daughter secondary spermato-

cytes, and since no period of growth seems to separate their

mitoses, the volume of the primary spermatocyte cell in the

metaphase is presumably equal to twice that of the secondary

spermatocyte. Connection is therefore suggested between the

spindle-length and cell-volume at this stage.

Although this ratio has been observed in organisms repre-

senting three phyla of the animal kingdom, the inference is

speculative; for coincidence may be responsible for the apparent

connection. i pointed out in an earlier paper that only one

generalisation seemed to have been established concerning the

mitotic spindle, namely, that it is not a figure formed entirely

by the action of forces at its poles. We have since found that

its length at the conclusion of spermatogenetic metaphases

cannot be correlated with the volume of the chromatin; and,

if we can eventually prove that the length at this stage is

or is not connected with the volume of the cell, we shall have

succeeded in establishing another generalisation.

In a paper on chromosome dimensions, published in 1912,

I stated that increasing somatic complexity of the organism

seemed to be accompanied by increase. of chromatin volume in

the germ-cell. The measurements given, however, proved that

no theory depending entirely upon a quantitative analysis can

suffice; for, in certain cases, the difference of chromatin volume

in widely separated organisms was found to be less than that in

organisms belonging to sister families.

I now intend to compare the volumes of spermatocyte cells in

Helix pomatia, Forficula auricularia, Triton cristatus, and Man—

organisms representing three phyla; and, in order that a com-

parison may be possible also in organisms belonging to allied

* Wor explanation of the Plates see p. 49.

46 MR. C. F. U. MEEK ON

families, I have included in PJ. J. photo-micrographs of

spermatocyte cells of two species of Stenobothrus, of which the

family is sister to that of Forficula. We know that the length

of the spindle in spermatocyte metaphases cannot be correlated

with the degree of somatic complexity of the organism; and,

from the investigations now to be carried out, we shall know if

the volumes of these cells cannot be so correlated. Moreover,

the photo-micrographs of cells in Yorficula auricularia and Helix

pomatia will afford opportunity of verifying my original mea-

surements of spindle-lengths at the conclusion of the metaphase.

Material and Methods.

All material was fixed in Flemming’s strong chromo-aceto-osmic

acid fluid, in which it remained for twelve hours. It was then

washed for twenty-four hours in running water; and, after being

passed through successive strengths of alcohol, was embedded in

parafiin. Sections were cut 8 or 10 » thick with a Cambridge

rocking microtome.

The slides were stained for either twelve hours in Heidenhain’s

iron hematoxylin, or fifteen hours in iron brazilin. In the

former’case the mordant was an aqueous solution of ferric

alum, and the slides remained in it for four hours; in the latter

case they were put for two hours into a solution of ferric alum in

70 per cent. alcohol. The iron brazilin enables spindle fibres to

be seen distinctly, and is a useful stain when camera-lucida

drawings or photo-micrographs are required.

The preparations were studied with a Zeiss apochromatic oil-

immersion objective of 3 mm. focus and N.A. 1:40, and the

various compensating oculars. The light was obtained from

a Graetzin lamp, and was passed through the holoscopic oil-

immersion substage condenser made by Messrs. Watson & Sons °

of London. With one exception, all photo-micrographs shown

were made at the same magnification with a Zeiss camera, the

apochromatic objective mentioned above, and compensating

ocular No. 4. The camera extension was 90cm. The magnifi-

cation was estimated with a stage micrometer graduated to

read one hundredth part of a millimetre, and a photo-micrograph

of this scale is included in Pl. II. The negatives and prints

have not been retouched.

A Comparison of the Volumes of Spermatocyte Cells in

different organisms.

Figs. 1-24 of the Plates are polar and lateral views of cells in

the metaphase or earliest anaphase.

Figs. 1-8 inclusive represent spermatocyte cells in 7Z'riton

cristatus. Fig. 9 represents a primary spermatocyte cell in

Stenobothrus viridulus, and cells of this generation in S. curti-

pennis ave shown in figs. 10, 11, & 12; in fig. 12, which is a

lateral view, the odd or heterotropic chromosome is seen passing

SPINDLE-LENGIH AND CELL-VOLUME. 47

undivided to one pole. The primary and secondary spermatocyte

cells of Yorficula auricularia are respectively represented by

figs. 13, 14 and 15, 16, 17. Figs. 18 to 21 show the primary

spermatocytes of Helix pomatia, and the secondary spermatocytes

are shown in figs. 22 to 24.

Now, exact measurements of cell-volumes cannot be made; but

it is evident from the photographs that, in the metaphase, the

primary spermatocyte cells of Zvriton cristatuws, Stenobothrus

viridulus, S. curtipennis, and Helix pomatia differ from one

another only slightly in size, and are considerably larger than

those in Forficula auricularia. Moreover, drawings sent to me

by Dr. von Winiwarter show that in Man these cells are smaller

than those of Z'riton, Stenobothrus, and Helix. And the same

results are obtained if we compare the secondary spermatocyte

cells.

In the circumstances, we must realise that cells of these two

generations may be of similar sizes in widely separated crganisms,

and of very different sizes in organisms that are closely allied ;

and increasing somatic complexity of the organism is not

necessarily accompanied by increase of the volumes of these

cells.

The Length of the Mitotic Spindle at the Conclusion of the

Spermatocyte Metaphases of Helix pomatia (Pl. II. figs. 18—-

24) and Forficula. auricularia (PI. II. figs. 13-17).

We will deal first with spindle-lengths in Helix pomatia.

Figs. 18 & 19 are polar views of the equatorial plate in the

primary spermatocyte metaphase. Figs. 20 & 21 are lateral

views, showing constriction of the chromosomes in progress.

I estimated the spindle-length at the conclusion of this meta-

phase to be 15°3 w; and, since the length found from these two

photographs for the slightly earlier stage is 15 4, my original

measurement seems to have been accurate.

Fig. 22 is a polar view of the equatorial plate in the secondary

spermatocyte metaphase. Fig. 23 is a lateral view of the spindle

at the conclusion of this metaphase. The spindle-length at this

stage was said to be 12:1 u, and this measurement is now verified ;

for I am not attempting in this paper to express spindle-lengths

in terms smaller than half a micro-millimetre, and the length

found from the photograph is 12 yp. Fig. 24 shows a slightly

later stage, when the anaphase has begun; the length of the

spindle in this cell is found to be 12°5 pe.

Let us now consider the spindles of Forficula auricularia.

Fig. 16 is a polar view of the secondary spermatocyte complex ;

ten chromosomes are arranged on the periphery of the spindle,

and two lie within it. Figs. 15 & 17 are lateral views of this

mitosis; the chromosomes are constricting in the equatorial

plane, and the stage depicted therefore immediately precedes

the conclusion of the metaphase. The length of the spindle,

A8 MR. C. F. U. MEEK ON

estimated from the magnification, is 8 uw in each cell, and this

confirms my original measurement; for the length was said

to be 7°8 x during constriction and 8-1 » at the moment when

constriction was complete.

Fig. 13 is a polar view of the primary spermatocyte complex,

and all the chromosomes are shown. Fig. 14 is a lateral view at

the conclusion of the metaphase; the chromosomes have com-

pleted constriction, and the daughter rods are ready to move

towards the two poles. This photograph has been made at a

magnification greater by s'-th than that of the remaining figures.

In my earlier paper the length at this stage was said to be

10-4 »; and, since the length, estimated from the photograph,

is 10°5 p, my original measurement is seen to have been accurate.

There is, however, a new factor that must be considered in the

case of this organism. J remarked in my paper that, at the

conclusion of the primary spermatocyte metaphase, certain cells

showed a spindle-length greater than that required by the ratio.

Such spindles seemed to be distorted, and, after careful con-

sideration, I assumed that they were abnormal in that their

true form had been destroyed in the process of section-cutting.

T have recently studied new preparations of this material, and

have again found spindles of excessive lengths. These occur

in cells that are closely packed together ; but, since many of the

spindles show no sign of distortion, we are not justified in

assuming abnormality in every case.

Four explanations can be put forward. First, the volume of

these cells in the metaphase may vary, and our proposition may

still be valid. In this case, however, various lengths will pre-

sumably be found at the conclusion of the secondary spermatocyte

metaphase; and I have not observed such lengths. Secondly,

the daughter chromosomes may remain apposed to one another

in the equatorial plane for a considerable time after constriction

is complete: if centrosome divergence continues during this

period, the various and excessive lengths may be explained.

This, however, cannot always occur; for, in this organism,

I have found and drawn primary spermatocyte cells in which

the daughter chromosomes have begun to move towards the

poles when the spindle-length is only slightly greater than that

estimated for the conclusion of the metaphase. Thirdly, our

proposition may require modification in that the length of the

spindle may be affected by the shape of the cell. My original

measurements in orficula and Helix were made from cells that

were approximately spherical, and this may explain the constant

lengths observed. When, however, cells are closely packed

together in a cyst, the spherical form disappears, and, if our

modification is valid, the spindle-length will vary with the shape

assumed. Lastly, the length of the spindle at this stage may be

connected with neither the volume nor shape of the cell; and,

in this case, our proposition is entirely disproved. If, however,

SPINDLE-LENGTH AND CELL-VOLUME. 49

this is so, why has the ratio in question been observed in Helia:

pomatia and Man 2

I hope to deal with these explanations in a subsequent paper.

In the meantime, the proposition remains a suggestion.

BIBLIOGRAPHY.

Meer, C. F. U.

1911. The Spermatogenesis of Stenobothrus viridulus. Journ.

Linn. Soc., Zool. vol. xxxii.

1912. A Metrical Analysis of Chromosome Complexes, show-

ing Correlation of Evolutionary Development and

Chromatin Thread-width throughout the Animal

Kingdom. Phil. Trans. Roy. Soc. ser. B, vol. cciii.

The Correlation of Somatic Characters and Chromatin

Rod-lengths, being a further Study of Chromosome

Dimensions. Journ. Linn. Soce., Zool. vol. xxxii.

1913. The Problem of Mitosis. Quart. Journ. Mier. Sci.

vol. lviii. Part 4.

The Metaphase Spindle in the Spermatogenetic Mitoses

of Forficula auricularia. Idem, vol. lix. Part 2.

» Ihe Length of the Mitotic Spindle in the Spermatocyte

: Metaphases of Helix pomatia. Proc. Roy. Soc., ser. B,

vol. Ixxxvii. No. 594.

EXPLANATION OF THE PLATES.

PrateE I.

Figs. 1-8. Polar and lateral views of spermatocyte cells of Tiiton cristatus, in

the metaphase and earliest anaphase.

9. Polar view of equatorial plate in primary spermatocyte metaphase of

Stenohothrus viridulus.

10,11. Polar views of equatorial plate in primary spermatocyte metaphase of

Stenobathrus curtipennis.

12. Lateral view of spindle in primary spermatocyte metaphase of Sfeno-

bothrus curtipennis; the odd or heterotropic chromosome is seen

passing undivided toward the lower pole of the figure.

Prats II.

Fig. 18. Polar view of equatorial plate in primary spermatocyte metaphase of

Horficula auricularia.

14, Lateral view of spindle at conclusion of primary spermatocyte metaphase

of Forficula auricularia.

16. Polar view of equatorial plate in secondary spermatocyte metaphase of

Forficula auricularia.

15,17. Lateral views of spindle in secondary spermatocyte metaphase of

Forficula auricularia.

18,19. Polar views of equatorial plate in primary spermatocyte metaphase of

Helix pomatia.

20,21. Lateral views of spindle in primary spermatocyte metaphase of Helia

pomatia.

22. Polar view of equatorial plate in secondary spermatocyte metaphase of

Helix pomaéia.

23. Lateral view of spindle at conclusion of secondary spermatocyte

metaphase of Helix pomatia.

24. Lateral view of spindle in earliest secondary spermatocyte anaphase of

Helix pomatia.

25. Divisions of stage micrometer, 10 « apart, showing magnification of

figs. 1-13 and 15-24.

Proc. Zoou. Soc.—1914, No. Was 4

P.Z.S. 1914. Laidlaw. Pl. 1.

University Press Cambridge.

BORNEAN DRAGONFLIES.

ON BORNEAN DRAGONFLIES. 51

4, Contributions to a Study of the Dragonfly Fauna of

Borneo.—Part II. The Gomphine and Chlorogomphine.

By F. F. Laiptaw, M.A., F.Z.S.

[Received October 18, 1913: Read February 17, 1914. ]

(Plate I.*)

INDEX. Page

Structure:

WEEN OE OFM DIO Socacec cae cdg coe see bas bor ocB cedor ace Goksdn pre cou eos DOF

Geographical Distribution :

Gomphinz and Chlorogomphine of Borneo...................:c000:.0. 51

Systematic :

Ictinus acutus, sp. i. Rae RAE sete nace 51

Burmagomphus ver mmiculatae Qtr sila SEOs Woces coos 55

Heterogomphus icterops Martin, borneensis subsp. n.? ............ 57

Introductory remarks.—I have dealt at some length with the

venation of the various species discussed below, more especially with

the species belonging to the Chlorogomphine ; chiefly because all

of them, and especially the latter, are very rare in collections,

and because they are of particular systematic interest to the

student of the Odonata.

With regard to the distribution of these members of the

Dragonfly fauna, too little is known to admit of dogmatic state-

ments or even of useful discussion. The genera which seem to

be essentially characteristic of the Great Sunda Islands are

Macrogomphus and Microgomphus, the latter unknown beyond

their limits; all the other: genera here recorded have a wide

distribution in the Orient, and in the case of /ctinws beyond it.

At the same time the Gomphine fauna of Tropical Asia is as

sharply characterized as that of any other quarter of the earth.

Ictinus AcUTUS, sp.n. (Selys nom.) (PI. I. fig. 1.)

26 6. Baram, Oct. 1910.

Length of abdomen 45 mm. + 3 mm.; hind wing 36 mm.;

pterostigma 5 mm.

Venation :—

19 or 20 13 or 14 3-4-celled ¢ e celled 2

ANTM AI ae AT a ol fea Dee OEE, wast Se: se

14 or 15’ Pn i Peasant 7 eal .3-celled’ supr.t “9

Belongs to the group /. decoratus de sei | :

Wings tinged with brown ; orange at the base—in the fore wing

for about one-third the length of the median space, but excluding

the cubital space ; in the hind wing almost up to the arculus.

Pterostigma brown.

Head: Upper lip yellow, edged with black, the black reaching

* For explanation of the Plate see p. 63.

52 MR. F. F. LAIDLAW ON

the base of the lip in the middle line; rhinarium yellow, marked

below with a transverse black line. Nasus and frons brownish

black ; the former has a small round yeliow spot on either side,

the latter carries a pair of transverse yellow spots at its summit,

these are separated narrowly in the middle line by black. Vertex

and occiput entirely brownish black.

Prothorax very dark brown, its under surface and base of the

prothoracic limbs paler.

Thorax rich chestnut-brown, marked with yellow as follows :—

A mesothoracic half-collar divided by the black median dorsal

carina ; a dorsal stripe on either side, incomplete below, and an

antehumeral stripe constricted at its middle; a narrow band on

the mesepimeron and a second band, likewise narrow, on the

metepisternum. Nearly the whole metepimeron is yellow, its

posterior border narrowly marked with black, whilst the meta-

sternum is entirely black.

Legs : Femora dark brown, the first pair with a narrow internal

stripe; tibie and tarsi black.

Abdomen largely black, segments 1, 2 brown below and at the

sides, auricles brown, 2 with a yellow dorsal band narrowing to a

pointat the apex of the segment ; 3, 4, 5,6 black, each with a small

basal yellow mark dorsally, which is triangular in shape with its

apex directed backwards (in the more adult of the two specimens

these markings are of a brown colour rather than yellow).

Anterior third of 7 yellow dorsally, 8, 9, 10 largely brown at

the sides; the distal half of 10 is also brown dorsally ; laminee

of 8 black.

Anal appendages black; upper pair fully equal in length to

9, 10 together, curved inwards a little and crossing each other

like the blades of scissors. Each bears a strong sharp spine on its

outer margin, rising at the junction of the inner and middle

third of its length, and the inner margin of each is finely serrated

near its apex. Lower appendage very short, almost concealed by

the anal tubercle, abruptly truncate, with a small hooked spur on

either side.

This fine form is well characterized by the anal appendages,

which are unlike those of any of the described species. The

type-specimen has been examined by M. Réné Martin, who has

very kindly informed me that it is identical with an example in

the de Selys collection labelled 7. acutws, which has not been

described.

(The type is in the British Museum; co-type, Sarawak

Museum.)

*ICTINUS DECORATUS de Selys.

Ictinus decoratus Selys, Hagen, Monogr. Gomph. p. 273, pl. 14.

fig. 4; Kirby, Cat. Odonata, p. 77; Kriiger, Stettin. Entom. Zeitg.

* Species marked with an asterisk are those of which I have not received examples

from Mr. Moulton.

BORNEAN DRAGONFLIES. 53

1898, p. 315; Williamson, “‘Gomph. ete. of Burma,” Proc. U.S.

Nat. Mus. xxxili. 1907, p. 279; Martin, Mission Pavie, p. 14

(sep.) (1904).

Recorded from Borneo (Jartin), Sumatra (Ariiger), Java

(Selys), and Tonkin (Martin).

IctINuS MELZNopes de Selys.

Ictinus melenops Selys, Hagen, Monogr.:Gomph, pp. 532, 686,

pl. 15. fig. 1; Selys, Addit. Synops. Gomph., Bull. Acad. Roy.

Belg. 2. vii. 1859, p. 548; Karsch, Entom. Nachr. xvii. 1891,

p. 244; Kirby, Cat. p. 77; Martin, Mission Pavie, p. 14 (sep.)

(1904); Williamson, loe. cit. p. 280, figs. 7, 8,29; Ris, Ann. Soc.

Entom. Belg. lv. 1911, p. 239.

Ictinus melenops, race sumatranus, Kriiger, Stettin. Entom.

Zeitg. 1898, p. 315.

I have examined one female taken at Kuching i in December

1899 (Sarawak Museum Collection).

The species is recorded also from Sumatra (Karsch, Kriiger)

and Malacca (Martin), Cambodia and Tonkin (Martin).

*GOMPHIDIA MACLACHLANT de Selys.

Gomphidia maclachlani de Selys, 2™e Addit. Synops. Gomph.,

Bull. Acad. Roy. Belg. 2. xxviii. 1869, p. 767 ; Kirby, Cat. p. 76;

Krier, Stettin. Entom. Zeitg. 1898, p. 314; Martin, Mission

Pavie, p. 14 (sep.) ; Williamson, loc. cit. pp. 281-282.

Recorded from Borneo (Selys), Sumatra (Ariger), Tonkin and

Anam (JJartin).

*GOMPHIDIA KIRSCHI de Nelys.

Gomphidia kir Sa Selys, 4me Addit. ee Gomph., Bull.

Acad. Roy. Belg. 2. xlvi. 1878, p. 673; id., Anal. Soc. Hspan. de

Hist. Nat. xi. 1882, p. 18; Kirby, Oni “4 76; Martin, Mission

Pavie, p. 14 (sep.) ; Wallimnson,. loc. cit. p. 983.

Recorded from the Philippine Is. (Selys), Borneo (Selys), and

Tonkin (Martin).

SIEBOLDIUS JAPPONICUS de Selys.

Sieboldius japponicus Selys, Hagen, Monogr. Gomph.

as i Seehese oi Kerby.) Cate ps0 “Williamson, loc. cit.

OF sb rldins grandis Kriiger, Stettin. Entom. Zeitg. 1898,

pp. 311-314; Laidlaw, Proc. Zool. Soc. Lond. 1902, i. p. 81,

pl. vi. fig. 33 a.

I have examined a male of this fine species, collected by

Mr. Moulton. It agreed closely with the male taken by myself

in Perak. Williamson is probably correct in regarding Kriiger’s

species as not distinct from that of de Selys.

Ranges from Japan to Borneo.

me) ~

54 MR. F. F. LAIDLAW ON

*MACROGOMPHUS ALBARD# dle Selys.

Macrogomphus albarde Selys, 4° Addit. Synops. Gomph., loc.

cit. pp. 416-418; id., Ann. Mus. Civ. Genova, xxvil. 1889,

p. 469; Kirby, Cat. p. 63; Karsch, Entom. Nachr. xvii. 1891,

p- 224; Kriiger, Stettin. Entom. Zeitg. 1898, p. 300; William-

son, loc. cit. p. 289; Ris, Ann. Soc. Entom. Belg. lv. 1911, p. 238,

Li GSH ieee

Recorded from Sumatra (Selys) and Borneo (fis).

MACROGOMPHUS DECEMLINEATUS Cle Selys.

Macrogomphus decemlineatus Selys, 42° Addit. Synops. Gomph.

pp. 418-419: id., Ann. Mus. Civ. Genova, xxvii. 1889, p. 469;

Kirby, Cat. p. 63; Kriiger, Stettin. Entom. Zeitg. 1898, p. 203;

Williamson, loc. cit. p. 289.

2 5 & from Kuching, May 1896 (Sarawak Museum Collection).

Recorded from Sumatra (Selys) and Borneo (Selys).

*M ACROGOMPHUS QUADRATUS de Selys.

Macrogomphus quadratus Selys, 4™° Addit. Synops. Gomph.

p. 415; id., Ann. Soc. Entom. Belg. xxvii. 1884, p. x; id., Ann.

Mus. Civ. Genova, xxvii. 1889, p. 469; Maclachlan, Ann. Soc.

Entom. Belg. xxviii. 1884, p. vii; Forster, Ann. Soc. Entom.

Belg. xliii. 1889, p. 65; Kirby, Cat. p. 63; Kriiger, Stettin.

Entom. Zeitg. 1898, pp. 296-297 ; Williamson, loc. cit. p. 287.

From Borneo (Selys) and Sumatra (Selys).

MicRoGOMPHUS CHELIFER de Selys.

Microgomphus chelifer Selys, Hagen, Monogr. Gomph. p. 364 ;

Selys, Addit. Synops. Gomph. p. 533; Kirby, Cat. p. 63;

Kriiger, Stettin. Entom. Zeitg. 1898, p. 302; Laidlaw, Proc.

Zool. Soc. Lond. 1902, i. p. 79; Williamson, loc. cit. pp. 295-296,

figs. 21, 22.

Sy acioass mol (z:

A new record for Borneo. Known from Malacea (Selys) and

Sumatra (A7riiger).

* LEPITOGOMPHUS SEMPERI cle Selys.

Leptogomphus semperi Selys, 4me Addit. Synops. Gomph.

pp. 443-444; Martin, Mission Pavie, p. 11 (sep.); Kirby, Cat.

p- 70; Williamson, loc. cit. pp. 292-293, fig. 17; Ris, Suppl.

Entom. Deutsch., Entom. Mus. no. 1, 1912, p. 69.

Recorded from Borneo (Martin), Philippines (Selys), Tonkin

(Martin).

LEPTOGOMPHUS WILLIAMSONI Laidlaw.

Leptogomphus williamsont Laidlaw, Journ. Str. Br. R. Asiat.

Soc. [no. 63] Dec. 1912, p. 94, figs. 1, 2.

This species belongs to section B of the genus as defined by Ris.

BORNEAN DRAGONFLIES. 55

Tt has the basal subcostal nerve of the second series present on all

four wings; the hamuli are large and the “ Penischale ” is small.

The upper pair of anal appendages are very similar to those

figured by Ris for LZ. perforatus Ris and L. sauteri Ris.

It is characterized by the possession of a single row of cells

only in the whole anal area, and by the rather striking yellow

spot on the dorsum of segment 10 of the abdomen. (For figures

of genital appendage of this species see Laidlaw, loc. cit.)

The type: d is in the British Museum.

[LEPTOGOMPHUS KELANTANENSIS (Laidlaw).#?f

Leptogomphus kelantanensis Williamson, loc. cit. p. 291 (1907).

Gomphus consobrinus Laidlaw (nom. preoccup.), Proc. Zool. Soc.

Lond. 1902, i. p. 80, pl. v.-fig. 5.

Gomphus kelantanensis Laidlaw, Proc. Zool. Soc. Lond. 1902,

li. p. 382 (footnote).

This species belongs to section A of the genus, according to

Dr. Ris’ arrangement. There is no basal postcostal of the second

series, the hamuli are small, and the “ Penischale” large. The

single male captured is fully adult and its sober colouring is

excellently shown in Mr. Wilson’s figure, where the venational

characters are also satisfactorily exhibited (Proc. Zool. Soc. Lond.

1902, pl. v. fig. 5).

Like Z. williamsoni, this species has its anal area composed of a

single row of cells. The upper anal appendages are rather small,

with a single well-marked but small tooth on their outer side at

about the middle of their length; the apppendage terminates in

a fine upturned point; each extremity of the lower appendage is

hooked upwards rather abruptly at its termination.

The type ¢ is in the Zoological Museum of the University of

- Cambridge. |

BURMAGOMPHUS VERMICULATUS (Martin), subsp. INSULARIS, nov.

Ceti) 2)

Gomphus vermiculatus Martin, Mission Pavie, p. 11 (sep.).

Burmagomphus vermiculatus Williamson (nee Martin 2), loc.

eit. pp. 298-301, figs. 27, 28, 29 (10); (Ris, Tijdschrift v. Entom.,

Deel lv. 1912, p. 164).

Sie

An interesting addition to the fauna of Borneo. In size it

agrees closely with Williamson’s specimens from Burmah, in

most other respects it appears to approach the individuals de-

scribed by Martin from Tonkin. I am disposed to believe that

Willamson’s examples represent a species distinct from the true

B. vermiculatus of Martin; but cannot be certain on the point

without a good supply of material from the two localities.

In any case the present specimen agrees rather with Martin’s

specimen than with those described by Williamson.

_ * Not recorded fram Borneo.

+ {The parentheses around the names of authors placed after scientific names in

this paper are used in accordance with Article 23 of the International Rules of

Nomenclature. (Proc. 7th Int. Cong. Boston, 1907, p. 44 (1912) —Eprror. |

56 MR. F. F. LAIDLAW ON

The individual here recorded shows well the generic characters

tabulated by Williamson. The wing characters are almost

identical with those shown in his figure (loc. cit. fig. 27).

A difference which may be merely individual, but if not,

one that I should regard as of specific importance, is that the

anal triangle of both hind wings of the Bornean specimen is

divided into two cells only, by a cross-nerve running parallel to

the long axis of the wing. Williamson’s figure might well have

been taken from this representative specimen from Borneo

excepting for this, for the difference in the number of antenodal

and postnodal nerves, and for the fact that in the Burmese

specimen figured the area included between Cu, and A, in the

hind wing is a little shorter and broader than in the individual

under consideration ; and, lastly, for a slight difference evident

in the relative size of the pterostigmata.

Details are as follows :—

Length of abdomen 28 mm.

Length of hind wing 28 mm.

Venation :—

13—12 9—9

An.n. = Pn, ——-

9—8 8—10

Pterostigma a shade longer and narrower than in the type of

the genus, covering 4 cells in the front wing.

Head: Anterior surfaces black, with a rectangular yellow mark

on either side of the upper lip, and a yellow spot at each angle

of the lip; a transverse yellow band along the crest of the frons

divided by a fine median line into two lateral halves.

Thorax black above, marked with a yellow mesothoracic half-

collar interrupted in the middle line, tapering laterally. A pair

of narrow dorsal stripes of the same colour, not reaching to the

base of the femora; a very small superior antehumeral spot on

either side. Laterally, from immediately behind the humeral

suture, the thorax is yellow marked with a black line which rises

below at the level of the first lateral suture, includes the stigma,

curves backward to join a second black line which follows the

course of the second lateral suture, but bifurcates above to enclose

a small yellow space.

Abdomen black marked with yellow; 1 with a lateral spot on

either side; 2 with a dorsal triangle, its apex directed backwards

covering the first two-thirds of the segment, sides including the

auricles also yellow ; 3-7 each with a fine yellow, basal, transverse

mark dorsally, occupying about the first eighth of the length of

the segment; 8 entirely black; 9 with the trace of a yellow ring

at its apex; 10 entirely black.

Legs black, the first pair of femora yellow on their inner side, the

second pair with a small yellow mark on the same surface distally.

Appendages black. The genital structures on 2 are almost

identical with those shown in Williamson’s figure (Williamson,

loc. cit. fig. 28.c), the second pair of hamuli not quite so prominent.

BORNEAN DRAGONFLIES. Da

Anal appendages relatively shorter, the limbs of the lower one

stouter and less divaricate (doc. cit. fig. 28,4,B. Cf. also Martin,

quoted by Williamson, loc. cit. p. 301).

[Ris loc. cit. has recently described a species from Java which

he refers to this genus under the name B. jacobsoni. This species

differs from B. vermiculatus in the colouring of the thorax, in the

shape of the genital hamuli, and of the anal appendages. It is

also a little larger.

He is inclined to regard the form described by Williamson as

distinct from the true B. vermiculatus of Martin. |

The specimen described above is deposited in the British

Museum.

HErEROGOMPHUS ICTEROPS Martin, subsp. BORNEENSIS, nov. ?

(EG JES wiles, a5)

Heterogomphus rcterops Martin, Mission Pavie, p. 9 (sep.); id.,

Bull. Mus. d’Hist. Nat. 1902, no. 7, p. 506; Williamson, loc. cit.

p. 316.

1g. Matang Rd. 28:3:10.

Length of abdomen (without appendages) 51 mm.

Length of hind wing 45 mm.

The cross-nerves on the wings on the right side of the single

specimen are highly irregular; it would seem as though that

during development those wings had suffered from a ‘ cell-

storm” which did not interfere with the general symmetry of the

wings nor yet largely with their main structural features, but

considerably disturbed the number and arrangement of the cross-

nerves, especially on and near the costal spaces.

Martin’s description of the species, based on an example in

de Selys’ collection, from Java, is very brief. Hence, without

actually confronting the specimens, it is impossible to say how

far the Bornean form here described is distinct.

Venation :—

13—15! 12—15

In addition, there are several incomplete antenodals on the

right fore wing, and two cross-nerves in the submedian space,

whilst there is also a single crogs-nerve in the supra-triangle of

the hind wing of that side.

The pterostigma has a well-developed brace on all four wings.

Head: With the exception of the vertex and occiput which are

very dark brown, and of the eyes which in the dead specimen are

brown, the head is light brownish yellow in colour; the pro-

thorax is brown, lighter at the sides.

The thorax is yellow marked with brown as follows :—A broad

humeral band continuous above with the brown of the antealar

sinuses and below witha narrow stripe running along the anterior

margin of the mesothorax ; a broad band runs down along the mid-

dorsal carina, narrowing below and just meeting the posterior

58 MR, F. F. LAIDLAW ON

margin of the mesothoracic half-collar, but not coalescing with the

brown stripe along its anterior border. Laterally, there isa narrow

stripe along the second lateral suture continuous over the back.

Legs: Femora red-brown, tibiz and distal parts black.

Abdomen rather pale brown, each segment, excepting the first

two and the last two, having a black ring distally. 1, 2 are

largely yellow at the sides and have yellow markings dorsally ; ;

1 this is confined to the posterior half of the segment, except is

a very fine line extending forwards to the anterior end of the

segment in the mid-dorsal line. In 2 the dorsal yellow colour

takes the form of an irregular longitudinal band. On 7 the

brown of the anterior three-fifths of the dorsum of the segment

carries a square yellow mark. 9,10 are unmarked, 9 dark brown,

10 lighter. The latter segment is very short, not half the

length of 9.

Anal appendages light brown, the extreme points finely tipped

with black. In general resembling those of H. smithi Selys, long

and slender, the upper pair as long as 9, 10 together, the limbs a

the lower appendage about four-fifths as long. The upper pair

are straight, their tips hooked downwards to a very trifling extent.

The limbs of the lower appendage arealso straight, except at their

apices which have a slight curving upwards, whilst each carries

at about one-sixth of its length from the apex a small sharply-

pointed spur, directed inwards at a right angle to the axis of

the hmb.

This large and handsome insect represents a genus new to

the fauna of Borneo, and is one of the most interesting of

Mr. Moulton’s many “finds” amongst the Odonata of the

island. ;

The.specimen will be deposited in the British Museum.

Subfamily CHLOROGOMPHINE.

In describing the venation of the species of this subfamily,

fuse the term “anal loop” to indicate the very definite area

lying below the cubital space bounded by branches of the anal

vein ae by An,+Cu,. The name is used in the same sense by

Needham (Proc. U.S. Nat. Mus. xxvi. p. 733) for Chlorogomphus.

The loop, as well as the area between Cu, and An, in the hind

wing, seems liable to considerable individual variation in the

genus.

J have figured the penis of Orogomphus dyak (P1. I. fig. 7),

and would call attention to its close resemblance to that of

Cordulegaster. The structure of the antenne is also well worth

remark,

Ris has noted the occurrence of tibial ridges in the males of

this subfamily (Ris, Coll. Zool. Selys, ix. p. 9, 1909). These

ridges, so far as one can judge, are scarcely adaptive structures,

and their presence would appear to me to indicate a real if

remote relationship to the Corduline.

BORNEAN DRAGONFLIES. 59

OroGoMPHUS DYAK Laidlaw. (PI. I. figs. 4-7.)

Orogomphus dyak Laidlaw, Journ. Straits Branch R. Asiatic

Sree IMINO jon, AU aN)

@ Gc

Venation :—

The triangles of the hind wings have their upper and outer

sides subequal, the inner side distinctly shorter, resembling that

of the fore wing.

Formula :—

Anal |

of hind ad Ann. Pn.n. M. Cu. t. (cells). suprat.

Dim ATT) | RE) GE esl es

di(type).... ——

eer MS 16a l= 15e OSS Go Talli beady

5—4

21—24, iM-—ahl 3—3 8

ior)

GB copsssoso === =

8—6 16—18 12—13 3—3 U1 Nal 5—4

The wings have a slightly yellow tinge, most marked when

seen with reflected light, between Cu and M, proximal to the

pterostigma, especially in the specimen marked @,,.

Head: Uabium brownish yellow, labrum black; rhinarium dark

brown, nasus bright lemon-yellow ; frons black, but with a fine

yellow line along the crest which is surmounted by a number of

fine black hairs, not so numerous as in O. atkinsoni. Vertex and

occiput jet-black. Ocelli and antenne as in O. atkinsoni.

Prothorax black above, the posterior margin edged with yellow,

and the same colour on the sides.

Thorax relatively small, black, marked with yellow lines. On

either side is a narrow dorsal thoracic stripe, widening a little

at its extremities, touching the mesothoracic half-collar below, a

little curved with its convexity inwards; followed by a broader

antehumeral stripe. There is a narrow stripe laterally on the

metepisternum, and the metepimeron is finely edged with yellow ;

the under surfaces are brownish, and there is a yellow interalar

spot and a fine yellow spot on each alar sinus.

Legs black, coxee and base of under side of femora of the first

pair yellow.

Abdomen black, marked with golden yellow as follows :—A fine

transverse line at the distal end of 1; the distal half of 2, but

this is largely obscured by a broad transverse black band lying

within it; the auricles are yellow. There is a very small pair of

lateral spots about the middle of 3, and a small terminal trans-

verse spot on the same segment. On 6 there is a terminal ring,

occupying about the last one-eighth of the length of the segment.

The anal appendages are black, rather longer than 10. They

resemble generally those of O. splendidus as figured by Dr. Ris,

but differ in detail; the upper pair is much slenderer than in

that species and has the ventral teeth much smaller; the lower

60 MR. F. F. LAIDLAW ON

appendage is much less deeply cleft and its limbs are pointed, not

truncate.

2 2. Matang Rd.

In details of venation these two specimens differ considerably

from one another.

Formula :—

Gana An.n. Pn.n. M. Cu. t. (cells). suprat.

22—23 12—11 38—3 8—7 2—2 5—6

A (type) ... —_——. — — ——

10—10 19—20 14—14 3—4 9—9 3—3 6—6

- 22—22 12—10 3—3 8—9 2—2 5—5

14—14 20—20 14—15 4—3 9—9 3—3 6—5

The difference is most marked in the anal area. Not only is

the number of cells here greater in specimen B than in A, but

there is also a marked difference in the number of cells lying

between the fork of Cu,: in A these cells lie in two rows, and,

excluding the marginal cells, are 9 in number; in B, on the other

hand, they are disposed in three rows and number 13 and 11.

As in the males the inner side of the triangle of the hinder

wings is the shorter, though in this sex the difference is not very

strongly marked.

The type-specimen, A, evidently the more mature, has the

wings suffused with a golden-brown tinge throughout; most

marked at the bases and apices. The colour is richest about the

periphery of the cell, the central part, especially at the apices of

the wings, being often distinctly paler. B, the younger specimen,

has the wings almost colourless, but with a very faint yellow

tinge between the nodus and pterostigma extending down the

wing as far as M,.

The colouring of the head, thorax, and body scarcely differs

from that found in the male.

The following are the principal measurements :—

Length of abdomen: ¢ 53mm., 9 56 mm.

Length of hind wing: ¢ 38mm., 2? 42 mm.; breadth ?

14 mm.

In both sexes a basal postcostal nerve is present.

Type ¢ & Q will be deposited in the British Museum.

OROGOMPHUS SPLENDIDUS de Selys. (PI. I. fig. 8.)

Orogomphus splendidus Selys, 4™° Addition Synops. Gomph.

_p. 681 (1878); id. Anal. Soc. Espah. Hist. Nat. xi. p. 16

(1882); Kirby, Cat. Odonata, p. 79; Martin, Mission Pavie,

p. 14 (sep.) (1904); Williamson, loc. cit. p. 278 (1907); Ris,

Suppl. Entom. Deutsch., Ent. Mus. No. 1, 1912, pp. 77, 79,

fig. 15a, b, Taf. iii. figs. 1-6, Taf. v. fig. 5.

Mr. Moulton has forwarded me two female specimens pre-

sumably belonging to one and the same species though showing

BORNEAN DRAGONFLIES. 61

some rather marked differences in venation. These two specimens

are, I believe, to be referred to O. splendidus de Selys. It is

evident from Ris’ study of the venation of three males belonging

to this species, that there is a considerable amount of individual

variation to be looked for, and the agreement between them and

de Selys’ type is close in other respects.

One of the specimens has been returned to the Sarawak

Museum, and unfortunately I did not before returning it make

full notes of the venational formula for both pairs of wings.

ah ony An.n. Pn.n. M. Cu. t. (cells). suprat.

—26 —13 —2 —9 —3 =)

i —20 —19 iG le Sie Shc —4

26—25 14—13 3—2 8—8 2—2 s—5

cf 138—14, 20—18 17—18 ae 8—8 3—3 56—4

(de Selys’ 23—25 14 ep Cand fee ome

type.) aes Pus BY ie

The most marked difference between Mr. Moulton’s specimens

is in the anal loop. In Ris’ photograph of the type g¢ from

Kosempo that area contains 19 cells.

Further, in Moulton’s specimens in 1. there are only 2 rows of

cells in the space between Cu, and A, almost to the margin of

the wing, whilst in 2. there are on the right side 4 rows and on

the left 6 rows; in the type male there appear to be 6 rows.

The wings of the Bornean specimens are coloured as follows :—

Base and apices of wings suffused with bright golden brown, on

the fore wing reaching to the inner angle of the triangle, and

on the hind wing one cell beyond the arculus; at the apices the

colour begins rather nearer to the pterostigma than to the nodus,

and is fainter on the anal margin of the wing.

The basal postcostal nerve is absent in these female specimens;

it is present in the male figured by Dr. Ris.

The colouring of the head, thorax, and body is as described for

the male.

The chief measurements are :—

Length of abdomen circa 56 mm.

Length of hind wing 48 mm.; breadth 17°5 mm.

[OROGoMPHUS ATKINSONI de Selys.*

Orogomphus atkinsoni Selys, 4™° Add. Synops. Gomph. p. 682

(1878); Kirby, Cat. p. 79; Selys, Ann. Mus. Civ. Genova, 2. x.

(p. 49 sep.) (1891); Williamson, loe. cit. p. 278, figs. 5, 6 (1907).

1 2, Bhowali.

(Indian Forest Research Institute per Dr. Imms.)

* Not recorded from Borneo.

62 MR. F. F. LAIDLAW ON

The following is an account of the single specimen of this

species that I have been able to examine :—

Venation: The triangle of the hind wing, as in the specimen

figured by Williamson from the de Selys collection, has its upper

and inner sides subequal, distinctly shorter than is the outer side.

The male, judging from Williamson’s figure, has the inner side

distinctly shorter than the other two sides, so that the apex of

the triangle is at its outer angle.

The triangles of the Intnl wings In my specimen are bisected

in each wing by a nerve running from its inner angle to

the middle of the outer side, whilst in de Selys’ specimen, on the

left side at any rate, the triangle is divided into 3 cells. Other-

wise the venation agrees in detail between the two specimens,

There is no basal subcostal nerve.

The wing-formula of the two female examples is as follows :—

Te Ann. Pn.n. M. Cu. t. (cells). suprat.

19—19 9—10 Jl 6—7 2—2 3—3

Bhowali, Kumaon...... ss

10—10 14—13 12—12 1—1 6—6 2—2 3—3

Bengal. Coll. Selys. 19— 12— 1— 7— 3— 3

(From Williamson’s —

tigure.) 12— ‘le 13 (2) 1 8 3 3 (24)

The wing-formula of the male is

Bengal. Coll. Selys. 20— 10— 1— 7— 2— 3—

(From Williamson’s

figure.) $= = 14— 1 5 2 2

In the specimen from Bhowali the extreme base of both pairs

of wings has a golden tinge, this does not extend so far as the

first cross-nerves.

Head: Labium dull brown. Labrum brown edged with black.

Rhinarium dark brown; nasus and frons brownish yellow, the

frons at its vertex car ries a line of fine black hairs, and is as high

as the summit of the occiput. In front it nes flattened.

Vertex black; the ocelli lie in a slightly curved line, the median

one being placed a little in advance ‘of the lateral pair.

The antenne have the second joint relatively very large and

stout, cylindrical in shape, and equal in length to the distal part

of the organ which consists of five or six slender Joints. The

large brown eyes meet at a point; and the occiput is small, dark

brown in colour, with a fringe of fine yellow hairs.

The prothorax is small, dark brown above, its posterior margin

lemon-yellow.

The thorax is relatively small, black, and thickly covered with

silky brown hairs; there is a pair of dorsal humeral stripes, rather

wedge-shaped, with their apices directed forwards, not quite

e touching the margin, of a bright lemon-yellow colour. ‘Two

broad bands of the same colour lie on either side of the thorax,

BORNEAN DRAGONFLIES. 63

the first on the mesepimeron, the second, the broader, on the

metepimeron ; the under surfaces are brown, and there is a yellow

spot on the interalar sinus.

The legs are black, the coxee and base of under surface of the

first pair are lemon-yellow.

Abdomen: Segments 1, 2, 3 and 5, 6, 7 a little dilated.

Black with golden-yellow markings as follows :—A fine transverse

band at the distal end of the dorsum of 1. The terminal half

of 2, but this band carries a black median dorsal spot, which has

vather the shape of a three-pointed ivy-leaf directed backwards,

the yellow band is also incomplete below. 3, 4, 5 with a pair of

lateral spots at their middle, and a terminal half-circle, inter-

rupted by the mid-dorsal carina. 6, 7, 8, 9 with the terminal

half-circle only.

Measurements of specimen from Bhowali :—

Length. of abdomen 57 mm.; hind wing 42°5 mm.

Length of pterostigma 3mm. ; width of head 9 mm.

Breadth of wing 15 mm. |

Norr.—I have omitted from the list Macrogomphus abnormis

de Selys as its provenance is doubtful.

EXPLANATION OF THE PLATE.

Fig. 1. Ictinus acutus, sp.n. Superior anal appendages of male, seen from above.

2. Burmagomphus vermiculatus (Martin), insularis subsp. n. ‘Thoracic colour

pattern (diagrammatic).

3. Heterogomphus icterops Martin, borneensis subsp. n.?

4. Orogomphus dyak Laidlaw. 6, wings of lett side.

5. 5 a A 2 , wings of left side.

6. # 5 3 6, anal appendages.

ds ae ms 43 6, penis and vesicle.

8. Orogomphus splendidus de Selys. 2, wings of left side.

(For the photographs from which figs. 3, 4, 5, and 8 are reproduced I am

indebted to the kindness of Messrs. F. W. & H. Campion.)

ON AN ABNORMAL SEA-URCHIN. 5D

5. Note on an imperfectly developed specimen of the Sea-

Urchin (Eehinus esculentus). By H. C. CHapwiok,

Aol Se

[Received October 30, 1913: Read February 17, 1914.}

(Text-figures 1-4.)

The subject of this note was collected and handed to me

for description by Prof. W. J. Dakin, who found it upon the

ruined breakwater at Port Erin during the autumn of 1912.

Prof. Dakin’s attention was attracted to it by two well-marked

depressions in the test, which were evident while the animal

was living and covered with spines. The depressions are in inter-

ambulacra 1 and 4; and, denuded of its spines, the test presents

the appearance of a lump of plastic material which has been

pinched by the thumb and forefinger. The spines of the entire

test were distinctly larger and more densely crowded than those

of a normal specimen of the same size from the same locality.

The apical system presents two abnormalities. Genital 4 is

of normal shape, but consists of two distinct plates united by

suture. Two closely approximated but distinct pores occupy the

position of the normal one on genital 1.

As shown. in text-fig. 1, the composition of the first seven

or eight plates of both series in ambulacrum II is irregular,

that of the first four especially so. The plates numbered 5

and 6 respectively in series @ ie to be composed of two

primaries, the adoral of number 5 being imperforate. In the

case of the elements which I have interpreted as plates 7 and 8,

I conclude that dp’ is the demi-plate of plate 7; and but for the

complete reversal of the position of the pores of the pair borne by

the small plate lying between it and the aboral primary of plate 6,

this might have been regarded as the adoral primary of the

same plate. dp” is probably the demi-plate of plate 8. The first

recognisable plate in series 6 is most probably number 5.

Number 6 1s clearly defined, but there are no traces of sutures

to indicate its composition. The position of the single pore-pair

suggests that it is that of a demi-plate. Number 7 appears

to consist of two primaries without a demi-plate. ‘The peripode

of the element which I have interpreted as the aboral primary

lies at right angles to the normal position and encloses only

one pore. It is, however, possible that the pore-pair which

lies avorally and to the right of this may be an element of

plate 7, and that the large ellipsoid tubercle occupies the position

of the pore-pair of the demi-plate. Should this view be correct,

the next. plate, number 8, is practically a normal one.

Proceeding now in the direction of the apical pole, all the

* Communicated by Dr. F. A. Barner, F.R.S., F.Z.S.

Proc, Zoom. Soe—_ 1914, No. V- +s)

66 . MR. H.C. CHADWICK ON

plates of series w are normal until number 46 is reached. Then

follow a series which, regarded from their inner, tuberculate

ends, apparently number seven plates, and which exhibit striking

irregularity of composition (text-fig. 2). It may be that the two

numbered 47 and 47a respectively compose one but slightly

imperfect plate, that portion of the suture which traverses

the large tubercle, and is represented by the dotted line, being

Text-figure 1.

Oral end of ambulaerum II. The Arabic numerals indicate the numbers of the

plates, reckoned from the peristome. a and 6, series in ambulacral area.

obscure. Number 48 consists of five elements, none of which ranks

among the primaries, while only one 1s imperforate. The next two

plates are numbered 49 and 49 @ respectively, on the assumption,

which the positions of three of the pore-pairs supports, that they

represent one ordinary plate. The appearance of the next plates,

numbered 50 and 50a respectively, suggests that they also

AN ABNORMAL SEA-URCHIN. ' 67

represent one ordinary plate. The presence of a fourth pore-

pair in this and the preceding plate is difficult to account. for,

except on the assumption that both represent small interpolated

elements.

Text-figure 2.

Abnormal plates in series a of ambulacrum II.

For explanation of lettering, see text-fig. 1.

In series 6 of ambulacrum III (text-fig. 3) plates 49 and 50,

while of normal composition, are exceptional in that the pore-

Text-figure 3.

Abnormal plates in series & of ambulacrum ILI.

pairs of their respective demi-plates are almost completely

outside the limits of the plate, and perforate the adjoiming

interambulacral plate.

rok

68 ON AN ABNORMAL SEA-URCHIN.

Similar irregularities, though not so well marked, occur in

plates 50 and 51 of series 6 of ambulacrum IV (text-fig. 4). In

this ambulacrum the presence of a fourth pore-pair in plate 50

Text-figure 4.

Abnormal plates in series a and 6 of ambulacrum LY.

in series @ and plate 52 in series b again suggests the inter-

polation of small and obscure elements. No. 52 im series 6

undoubtedly has an interpolated demi-plate.

P ZS, O14 Hoge ame

Cambridge University Press.

H.R.Hogg, del.

SPIDERS FROM THE MONTEBELLO ISLANDS.

PZ. 8. 1914) Hoses eae

106.

H.R.Hogg, del - Cambridge University Press.

SPIDERS FROM THE MONTEBELLO ISLANDS.

ON SPIDERS FROM THE MONTEBELLO ISLANDS. 69

6. Spiders from the Montebello Islands.

By H. R. Hoge, M.A., F.Z.S.

[Received December 22, 1913: Read March 3, 1914.]

(Plates I.—II.*)

INDEX. Page

ee Oe Aa te Te Nee Se ese AO

Nephila venosa ....... eat os AN Mena Pe debe seidzad | We

Nephila meridionalis, var. on MOV hw cern eee ee ential

AURTUODE PU EISOULIAL cisotiasilse bap oop opens poe osbiasokaneonscessaonr nscae: + UP)

AVR OPUG (NCR Oe We. bob sacormooy cacao sadece yegoseecsevsveecuncas HG)

JEGERTOUC: TROHIG IB SIDS, ogo soo sasoniv 08400 sap sssapsasebaasecasenn sone | US

ALRTGUS POUGPSUB;, BVe Wo cee sbicscenssocced seonsanbonundedcboaopanewe UE

Gasteracantha minax ...... SA Srcagt reed MID Te aan eee LD,

Gasteracantha minax, var. lenges MEAS cite hen ee age PETG

Gastenacanthormuncdc ala dSUn0g Cha eee eee ee eee eee ae

Gasteracantha minax, var. hermitis, nov. ..................... 80

RAEN OSOGR AG tS) Oe ena cee pees be concen aad ana aoe batonbenctanareateode) | Mo

SENATE TOP BG ING NS appisdedac cdaclocaciadstcoues Spee cedepbeadnageen eA

OO STCOUCGAS LET Me vate eRe eee eae eee se en ee been Om

OiioswWermitisns Ds Wey acre aero et sees ME Ee eee oeSeE EOD.

WMiontebellortenuis, seus et Spal.) --------ees see eene eee ees OO

MEY COSOICUG TG Sa RO eee ee Oe eee oe OO

OEGROPDES TATTOOS. < ecicen bieand Gericie-scc) babes Rad ste Sper cmnuceGaare) eke

IEC RGE TOCURGTUPOLOM, So Wa oodse6203002 no saadadoas -xpsancee ts)

LW GTPDOSSED TUITOS,, Oo W6- “Sac ado cnadcabsencs 405 22m oubatobaU Nea sAaRe0neS 90

The collection of Spiders hereunder described was made partly

by Mr. P. D. Montague, and partly by Mr. T. H. Haynes. By

the kindness of these gentlemen they have been placed in my

hands,

The Montebello group is situated on the N.W. coast of

Australia, off. the Port of Onslow, W. Australia, the largest,

Hermite Island, being 90 miles from the mainland, in lat.

20° 30'S., long. 145° 15’ W.

Mr. Montague visited the islands for the purpose of collecting

specimens of the fauna, and Mr. Haynes lived on Hermite

Island for about three years, until driven off by a hurricane

which destroyed his hut and a larger collection which he had

been good enough to make at my request.

Mr. Haynes informs me that the soil consists of sandstone-

rock, similar to that on the mainland, of which therefore the islands

were probably part, and the S.E. winds prevailing for the six

winter months blow off the land.

It will be noticed that by far the larger number, both of

species and specimens, are of the family Argiopide, with a small

sprinkling of Lycoside, Thomiside, Clubionide, Oxyopide, and

Attide.

* For explanation of the Plates see p. 92.

70 MR. H. R. HOGG ON SPIDERS

Notable absentees are Delena cancerides Walck., and Latro-

dectus hasseltii Thor., found generally in every part of Australia,

but no specimens have been brought from here.

Family ARGIOPID4.

: Subfamily TETRAGNATHIN#.

Group TETRAGNATHES.

Genus TerracNnatHa Latreille.

TETRAGNATHA ANGULATA, sp.n. (PI. I. fig. 1.)

A single specimen (P. D. M.).

Female. The cephalothorax is greyish yellow, with broad darker

depressions separating the cephalic from the thoracic part and

broad dark grey radial lines from the margin of the cephalo-

thorax to a transverse broad oval depression, one-third of the

distance between the end of the cephalic part and the rear end

of the thoracic, also dark grey round the margin of the thoracic

part; it is sparsely covered with short, fine, white hairs. The

eyes are yellow on black ground. Mandibles yellow tinged with

dark grey at the sides, and witha few white hairs. Fangs yellow-

brown. Sternum rather dark yellow-brown. The lower part of

the lip the same. The upper margin of the lip and the maxille

are bright yellow with light brown fringes. Legs and palpi pale

yellow with brown spines, rising from brown patches. The

abdomen above is greyish yellow, with a mottling of darker

-erey, pale yellow below with a broken median darker yellow

stripe.

The cephalothorax is twice as long as broad, convex, narrowed in

front, rounded at the sides, and slightly hollowed at the rear end.

The rear row of eyes is straight, or slightly recurved when

seen from above, of equal size, the median -23 diameters

apart and 3 diameters from their laterals, which are their

diameter away from the front laterals. The front row, more

strongly recurved, has the laterals only half the diameter of the

rear, lying on the same tubercle therewith. The front median

are rather larger than the rear, 14 times their diameter apart,

and the same from the rear, so that they form a quadrilateral,

broadest posteriorly ; they.are one-half their diameter from the

margin of the clypeus, which overhangs the insertion of the

mandibles. These are divergent, projecting forward, kneed at the

base. On the outer margin of the falx-sheath are five teeth at

equal intervals apart, the upper one situated at the anterior inner

corner is the largest, the others diminishing in order of sequence.

On the under side the upper one is similarly placed, followed by

two rather smaller, at intervals of their length, and then three

quite small close together.

The maxille are rather divergent, truncate, and broadest

FROM THE MONTEBELLO ISLANDS. 71

anteriorly, three times the height of the lip, which is broader

than long, rounded in front, straight at the sides with the

upper margin clearly protrudent. The sternum is shield-shaped,

14 times as long as broad; from its greatest breadth between

the second paix of coxee it narrows slightly anteriorly and poste-

riorly to a point between the not quite contiguous rear coxe.

The legs are long and thin, the metatarsi and tarsi tapering to a

very fine point. The abdomen is twice as long as broad, widening

from the base to halfway of its length, whence it rather suddenly

narrows and tapers to the spinnerets. The epigyne is dome-

shaped on the upper half, overhanging two oval hollows, one each

side of a median broad ridge.

This species is rather near L. Koch’s 7. gemmata from Port

Mackay, but is smaller; the first pair of legs not so long com-

pared with the second, and the mandible on the upper side is

without the large tooth near the middle of the anterior edge ;

the coloration is much lighter, and there are brown spots on

the legs.

The measurements (in millimetres) are as follows :—

Long. ‘Bread.

Cephalothorax... 2 ]

AN NCIOMME Ms cccoe0ce 4. 2 ,

Mandibles ...... 14

Pat. & Metat.

(Com, Airs es Weis ADs & tars.

La meee I = 4 5) a — eal

Me 1 3 3 3 = 94

: a 14, I a 4t

4. = 3 a 3 = 84

babes 1 1 3 hoy eee 1

IPI OVL Ee tithe ee ae omer = 5 rs} 3; = Me

Subfamily Neparmr a.

Group NepniLe.

Genus Nepuina Leach.

Neraita venosa L. Koch.

8 females (2 non-adult), P. D. Montague.

Hab. (sec. Rainbow). Polynesia, New Guinea, Torres Straits,

Queensland, N.S. Wales. Victoria and 8. Australia (H. R. #.).

In these specimens the eyes of the rear row are clearly smaller

than those of the front row, whereas in L. Koch’s description

they are of the same size. This is also the case in my specimens

from §. Australia. The longitudinal lines on the rear end

of the abdomen are also absent, but the specimens agree in other

respects; and I see no reason for making them a local variety,

which they might be said to be if only found on this particular

island.

{2 MR. H. R. HOGG ON SPIDERS

NEPHILA MERIDIONALIS Hogg, var. HERMITIS nov. (PI. I. fig. 2.)

od?

Trans. Royal Soe. of 8. Australia, vol. xxxiv. 1910, p. 59*.

4 females, T. H. 10 females (6 non-adult), P. D. M.

These correspond in almost every particular with my WV. meri-

dionalis from Kangaroo Island, 8. Australia.

They differ, however, in having the distance between the rear

row of median eyes rather greater than that between the front

median instead of the same; in the elypeus 14 times as wide as

the distance between the front median eyes instead of equal to

it; and the pale lines forming the pattern on the under side of

the abdomen much finer. There are also two small black spots

behind the eyes and two more on the margin of the cephalic part.

T have therefore made it a new variety : hermitis.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 12 le in front.

Abdomen......... 14 9

Mandibles ...... 5

Pat. & Metat.

Coxe. Tr. & fem. tib. & tars.

4&11

Wegs ites 1 24 1 15 DEES

y) 24 15 13 1D se

3 11 9 6 Ql =

4 2 14 10 4

Ball ee epee: 1 4 4 3 0 =

Tibia longer than patella.

Another specimen measures :—

Long. Broad.

Cephalothorax... 10 1 oe cia

Abdomen......... 13 8 °

Mandibles ...... Al

Pat. & Metat.

Coxe. Tr. & fem. tib. & tars.

Megs... fen. ] 24 16 15 21 =

) 24 15 13 19) =

3 2 9 7 10 =

4 24 14 10 IL: =

Palio. sn teachers 1 4 4. 4 =

bo E> OT

or SO Or

Die

Differing very slightly from the original type-specimen.

Another, with a cephalothorax 9x7mm., has the abdomen

20x 14mm., apparently full of eggs.

oO”

* The length of the cephalothorax in my original description is misprinted

14} for 103 millimetres.

FROM THE MONTEBELLO ISLANDS, 73

Subfamily ARGIOPIN»,

Group ARGIOPER.

Genus ArGiorEe Aud.

ARGIOPE TRIFASCIATA Forskal.

IS emellies, 125 1D. Wil, ah ale Isl

Widely spread over Northern Australia, the Pacific Islands,

and many other tropical and subtropical parts of the world.

ARGIOPE HAYNESI, sp.n. (PI. I. fig. 3.)

This species, of which Mr. Haynes sent 10 females, is closely

related to L. Koch’s A. protensa and A. syrmatica, though ib

differs from them both more than they do from one another, and

I have therefore named it as new.

The cephalothorax is yellowish grey, thickly covered with

smooth silvery grey hairs all over the thoracic part, and in a

median longitudinal area between the eyes and at the sides of

the thoracic part the darker under coloration shows through, but

there are no definite dark longitudinal lines as in protensa and

syrmatica.

The mandibles are dark grey at the base and on the outer

sides, rather bright yellow on the inner and anterior portions.

The fangs red-brown at base and pale red forward. ‘The lip and

maxillz are bright yellow on the upper half, brown below. The

sternum is black-brown, with fine downlying grey hairs at the

sides, and a thickly haired yellowish-white median area reaching

the whole length.

The abdomen is silvery white with fine white hairs on the

upper side; a narrow black longitudinal line, running from the

base to rather more than halfway, with two transverse lines at

the upper end, like a Russian cross. There are no longitudinal

lines at the posterior end. The base of the under side is yellow

at the sides and dark grey*in the middle. The dark area is

continued in a broad median stripe to the spinnerets, bordered on

each side by a white network-patterned area, which passes behind

the spinnerets, the dark stripe being continued to the posterior

end, asin A. protensa, but not in A. syrmatica. The sides are

dark grey with pale yellow longitudinal lines.

The legs are darkish yellow, with long brown spines and grey

hairs. The anterior half of the metatarsus and the whole of the

tarsus of all legs have short brown bristly hairs. The front of

the patella and base of the tibia is dark brown. Near the anterior

end of the tibia is a brown ring, and the anterior end of same

and base of metatarsus are likewise covered by a brown spot. The

under side of femur 1 is dark grey.

The cephalothorax, one-third longer than broad, is rounded at

the sides, the cephalic part, narrow and short, being divided by

well-marked depressions from the thoracic. It does not reach

to the median fovea, which is short and recurved. The whole

74 MR. H. R. HOGG ON SPIDERS

area iS rather flat and covered with particularly smooth long

hairs all lying longitudinally.

The front row of eyes is straight, the median pair their

diameter apart and 14 times the same distance from the laterals,

which ave only half their diameter. -These are also their dia-

meter from the rear laterals, the diameter of the latter being

half as large again, and farther away, sideways, by the bred

of the front laterals. The rear median are the same size as the

front median, two diameters therefrom and 14 diameters from

one another. Therefore they form with their laterals a strongly

procurved row.

The mandibles are short and conical, only slightly kneed at

the base and divergent at the anterior end, ‘The fangs are

slightly curved and rather long. On the upper part of the outer

margin of the falx-sheath is one long tooth between two shorter,

and on the inner in the same position three also, the lowest being

the largest.

The maxille are convex, as long as broad, obtusely arched on

the upper edge and rounded at the back. The lip is broader

than long, curved to an obtuse angle in front, where it is very

convex, with a short narrower lower stem.

The sternum is shield-shaped, scolloped at the sides, half as

long again as broad; opposite the second and third coxe the

median area is projected into high round protuberances, and at

the posterior end is a similar but still larger oval knob. The

hair is downlying, finer at the sides than in the middle streak,

and there are a few long bristles at the anterior end.

The abdomen is 23 times as long as its breadth in the middle,

whence it tapers to a narrow rounded point at each end.

On the under side the base is less constricted and more rounded

off. the sides are straighter until beyond the spinnerets, where it

is suddenly narrowed into a stunted tail. On the back are four

pairs of muscle-spots.

The epigyne is formed in the same unique fashion as in

A. protensa and A. syrmatica, a chitinous cushion-like pear-

shaped projection, broadest at the base and having a blunt ova!

fore end, standing straight up. The basal end is about one-third

of the whole breadth of the abdomen at that part.

The legs are rather long and very fine in the anterior joints,

the tibia broadened out and flattened at the anterior end, and the

trochanter of the fourth pair nearly as long as the coxe.

The femoral joint of the palpi is strongly incurved, and broadest

at the anterior end. The tibia is twice as long as the patella, and,

the metatarsal jomt is covered with long bristly hairs.

The measurements (in millimetres) are as follows :—

Long. Broad.

1) season

Cephalothorax... 4A ' e in front.

Abdomen......... 10 4

FROM THE MONTEBELLO ISLANDS. 15

Pat. & Metat.

Coxe. Tr. & fem. tib. & tars.

LCOS, 7.) eens Il 1 7 64 8 = 23

2. 1 7 6 Bo) cee

35 3 41 3 AOE eee) IDE

4 3 7 5 7 = 198

Palipw see Aaerie ees 4 2 it 1 a

The obvious difference between this species and 4. protensa

and A. syrmatica, which latter are very much alike, is the

shortened tail, the absence of the dark longitudinal markings on

the cephalothorax and of the longitudinal markings on the

posterior end of the upper side of the abdomen, almost every

other point being the same or very slightly modified in the three

species. ;

In the somewhat allied genus, Arachnura Vins., several species

have been constituted on the differences of the knobs at the end

of their tails and of their shoulder-humps. Oftentimes these

differences are seen in spiders of the same group of webs, and the

spiders are possibly all derived from the same batch of eggs. [am

not satisfied that this does not occur in the above species, the

two first named being both described from the same locality,

Port Mackay, in Queensland. The differences between them, if

any, are very trifling.

Group MancorEs.

Genus LAarinta Simon.

LARINIA MONTAGU, sp.n. (PI. I. fig. 4.)

2 females, P.D.M. 1 female, T. H.

Cephalothorax pale yellow, with a slightly darker median

longitudinal streak and thinly spread white hairs. Mandibles

the same, with pale yellow-brown fangs. Maxille and upper

half of lip pale yellow, but base of latter light brown. Sternum

pale yellow in median area, with light yellow-brown along the

sides and at the lower end. Legs and palpi similar pale yellow,

with fine white hairs, light brown spines on brown roots, and

brown bristles on tarsus and metatarsus. On femur 1 the brown

spots are much more numerous than elsewhere.

The abdomen above is pale vellow with white hairs and white

upstanding bristles, but mottled with darker brown in patches,

giving the whole a uniform dark appearance. On the under side

it is a network of paler and darker yellow. The spinnerets are

brighter yellow, and the epigyne brown with yellow in the hollows

and an oblong grey area behind it.

The cephalothorax is moderately convex, twice as long as broad

and one-half of its greatest breadth across the eye-area. The

sides of the cephalic part are straight, those of the thoracic

76 MR. H. R. HOGG ON SPIDERS

rounded and there are no depressions separating the two; a

longitudinal fovea reaches into the rear slope.

The rear row of eyes is slightly recurved and stretches quite

across the cephalic part. The median pair are less than one-

third of their diameter apart and four times their diameter from

the side eyes, which are rather more than half the same diameter

across. The front median are one- oe wider than the rear

median, twice their diameter apart, t, and 22 times the same from

the rear, thus forming a trapezium ce as wide in front as

posteriorly and slightly longer than broad. In a lateral direction

they are the same distance from the side eyes as from the rear

median. The front side eyes are the same size as the rear side,

but half their diameter away, and the front and rear are each on

a small separate tubercle. The clypeus is 14 times the width of

one of the front median eyes.

The mandibles are conical, kneed at the base, smooth and shiny.

There are two teeth on the outer falx-margin, the anterior twice

as long as the lower one; three on the inner margin.

The maxille are convex and nearly square. The lip has a

short, straight-sided base, the longer and very convex upper part

curving to an obtusely angled point.

The sternum is slightly convex, twice as long as broad, shield-

shaped, hollowed in front, almost straight at the sides, and narrows

rather suddenly to a point, which does not go between the con-

tiguous rear coxe.

“The abdomen is oval, 23 times as long as broad ; it is: sparsely

covered with fine downlying hair and has upstanding spinous

bristles on the upper side.

The legs are thin and tapering, rather profusely covered with

small brown spots, from which spring long upstanding spines

and bristles on the patellar, tibial, and metatarsal joints and

under side of femoral. Bristles only on the tarsa! joint.

The palpi have the femoral joint incurved, the patella one-half

as long as the tibia, and long spinous bristles on the latter.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 3 | : front.

Abdomen......... 5 2°

Mandibles é We

Pat. & Metat.

Coxe. Tr. &fem. tib. & tars.

Tees 2. emg 320 0s Te sea

leg 3 4 Ae IG

Be 9 Bey ws ae GE

Me B 3 31 1 = 108

Pair Sees. : bate = i 3 eS De

Of L. Koch’s two species, Z. phthisica and L. tabida, this species

more nearly resembles the former, but is two-thirds smaller and

much brighter in colouring. The rear median eyes are larger

FROM THE MONTEBELLO ISLANDS. 77

than the laterals instead of the same size, and the row recurved

from above instead of straight; the clypeus broader. The

abdomen also is broader in comparison with its length, and there

are only two teeth on the outer falx-sheath (one quite long)

instead of four small ones. The epigyne also differs from

L. Koch’s drawing.

The colouring is very close to Dr. Kulezyski’s Z. vicina, from

New Guinea, but this species is smaller: the epigyne differs ; the

rear middle eyes are farther apart; the mandibles longer, two

and three teeth respectively, instead of four, on each max gin ;

spots on first pair of legs instead of. none; no spots on “the

back instead of six pairs. The abdomen is also wider in pro-

portion to length.

L. montagui is also very close in most respects to Von

Keyserhing’s Larinia (Hpeira) talipedata, but in the latter the rear

median eyes are as large as the front median, and twice their

diameter apart, instead of close together.

This species differs in many points from M. Simon’s L. eburnei-

ventris, from 8.W. Australia.

Group ARANE#.

Genus ARANEUS Clerek.

ARANEUS REVERSUS, sp. n. (PI. I. fig. 5.)

This spider belongs to L. Koch’s and H. Simon’s first. series,

with shoulder-humps, the abdomen rounded in front and

obtusely pointed at the rear.

The cephalothorax, mandibles, and sternum are black-brown

with long coarse white hai and white spinous bristles.

The lip and maxille are black-brown with wide greyish-yellow

margins and red-brown fringes.

The coxee and femoral joints of the legs and palpi are dark

dingy yellow-brown, the patella and tibia nearly black-brown,

thickly covered with white hair and numerous spines brown

just at the base, but white from there to the point. The

metatarsus and tarsus dark dingy yellow-brown with white hair

on the basal half of the former, but brown bristles and yellow

spines on the anterior half and on the tarsus.

The upper side of the abdomen is at the base yellow mottled

with brown spots. Running from this to about halfway down

the back is a black-brown median stripe, bordered with yellow

and scolloped at the edges into three divisions, and each side

of this is a black transverse streak, separated from it by a

short longitudinal yellow streak, reaching to the shoulder-

humps. From each of these a dark brown scolloped line runs

along the side, meeting at the rear end. The latter enclose

another scolloped triangular area of brown and yellow inter-

mixed, the straight upper side touching the end of the anterior

median stripe. The front part is thickly set with upstand-

ing white spines on brown bases. On the under side a pale

78 MR. H. R. HOGG ON SPIDERS

yellow shield pattern, mottled with brown spots, reaches from

the genital fold to the spinnerets, with two dark brown longi-

tudinal spots thereon, the basal area being all brown. The

spinnerets are black-brown, the epigyne yellow-brown. There

is a pair of small brown conical protuberances, one on each

side of the breathing-slits. On the sides are vertical black-brown

stripes on a dingy yellow ground.

The cephalothorax is one-fifth longer than broad, straight in

front, where it is not quite one-half its greatest breadth, convex,

rounded at the sides, thickly covered with forward-pointing long

coarse hair and upstanding bristles, but bare on the rear slope.

The median quadrangle of eyes is on a somewhat low pro-

tuberance lying on the front slope. The rear row is straight,

the median their diameter apart, the same distance from the

front row, and six diameters from their respective laterals. ‘The

front median pair are 14 times the diameter of the rear

and that distance apart. The clypeus is the breadth of one

of them. The side eyes are equal in size on a common raised

prominence, not much smaller than the rear median, almost

touching one another, the front one lying just below and by

the distance of its diameter nearer the centre eyes than its

upper companion.

The mandibles are broad, conical, kneed at the base, some-

what divergent, and as long as the front of the cephalothorax is

broad. They are furnished with bristly hairs for one-third of

the distance from the base and on the inner edges, the remainder

being smooth. The fangs are stout and not much curved. On

the outer margin of the falx-sheath are three stout teeth near

the base. I could not see those on the inner margin.

The maxille are nearly triangular, straight on the inner side

and front edge, with rounded back and corners. They are as

high as the greatest breadth, which is at the front margin. The

lip is convex, broader than long, rounded in front, and less than

half the length of the maxille.

The sternum is convex, shield-shaped, rounded at the sides,

pointed at the rear, where it does not divide the rear coxe.

There are rounded prominences opposite the Ist, 2nd, and 5rd

coxee. It is thickly covered with long coarse hair and upstanding

bristles, particularly thick at the side edges, opposite coxe 1

and 2.

The abdomen is ovate, rounded in front, obtusely pointed at

the rear, thick at the sides. There are two shoulder-humps on

the upper side. From between these to the base it is thickly

covered with short thick upstanding spines, of which there are

also some few in other parts. It is moderately thickly covered

with downlying rather coarse hair both on the upper and under

side. From the rear end of the upper side to the spinnerets the

abdomen sinks perpendicularly a distance equal to the length of

the cephalothorax.

The epigyne is of the curious upright pillar type, a stout thick

FROM THE MONTEBELLO ISLANDS. ras)

rounded column rising from an oval muscular scape. The

anterior portion is club-shaped seen from behind, but a flattened |

wedge from the side.

The legs are moderately long and stout, the anterior end of

the tibial joints being flattened and broadened out. There are

bare streaks on the sides of these, but none above. The thick

hair is downlying and smoother than in other parts, and there

are no spines above either on these or the patelle, though several

at the sides.

The femoral joint of the palpi is curved inwards, broadest and

flattened at the anterior end. The tibial joint is twice as long

as the patellar, and numerous upstanding spines and bristles

cover the whole length.

The measurements (in millimetres) are as follows :—

Long. Broad.

it Se

Cephalothorax... 5 | i in front.

Abdomen........- 11 9

Mandibles ...... 25

Pat. Metat.

Coxe. Tr. & fem. & tib. & tars.

Wegs se psi. I, 13 6 3 oS = zd

erence ent. 6 5 = Ge

De 1s Ad 34 Sells

A. 12 54 5 a le

ingen lip tiers «asec 1 2 2 2 = A

A small non-adult male has the scolloped black stripe at the

anterior end of the abdomen and the black scollop bordering the

inner pattern at the posterior end, on a yellow-grey ground.

Group GASTERACANTHA,

Genus GASTERACANTHA Sund.

GASTERACANTHA MINAX Thor.

10 females and 1 male, P. D. M. 6 females, T. H.

This spider is found in considerable numbers from the east coast

of New South Wales, through Victoria and South Australia to the

Indian Ocean in Western Australia, its place being taken on the.

north coast by Gasteracantha vittata Thor., which appears by far

the most common species there. These specimens are from the:

most northerly point yet recorded for the species.

GASTERACANTHA MINAX Thor., var. LUGUBRIS L. Koch.

5 females; all black, no bright markings.

GASTERACANTHA MINAX Thor., var. AStRIGHRA L. K,

6 females; black, with orange spot on sternum.

80 MR. H. R. HOGG ON SPIDERS

GASTERACANTHA MINAX Thor., var. HERMITIS noy. (PI. I. fig. 6.)

4 females. Abdomen pearl-grey above, legs, cephalothorax, and

sternum bright orange.

I have previously pointed out (Proc. R. 8. Vict. vol. xi. 1900,

p. 79) that specimens of L. Koch’s species Gasteracantha astrigera

and G. lugubris were generally found wherever there was a

number of G. minax Thor., and that there was little or no

structural difference between the three. I therefore designated

the former as varieties only of the latter.

On this small island we find associated with G. minaa not only

these two varieties, but a third, emphasizing the fact that

although very different in coloration they are really only varieties,

possibly interbreeding, but all essentially the same species.

The shape of the mandibles, mouth-parts, sternum, vulva, and

ocelli markings on the back are the same in every case. The

spines, however, often vary in length and shape in the same

group of similarly coloured specimens.

Family THOMISID &.

Subfamily MisuMENINz.

Group Drerex.

Genus Dieta E. Sim.

Diera ISOLATA; sp.n. (PI. II. fig. 7.)

The cephalothorax is pale canary-yellow, except over the eye-

space, which is quite white, with a few scattered fine white hairs.

The mandibles are darker yellow for the basal half, the anterior

half bright pale yellow with pink fangs and yellow-grey fringes.

The lip and maxille pale yellow. Sternum dark grey at the sides

with yellow in the middle and pale yellow-grey hair. ‘The legs

and palpi are bright yellow, with yellowish-grey spines and a few

whitish hairs. The claw-tufts dark grey on the legs and white

on the palpi.

The cephalothorax is straight in front and at the sides as far

as the back of the eye-space, whence it is almost round, being

very slightly longer than broad (; mm.}. It is slightly convex

at the sides, but quite flat in the middle and a little higher

before the rear slope, whence it slopes gradually to the front.

On the thoracie part are faint broad shallow striations and a

similar longitudinal fovea.

The pedicule joining the cephalothorax and the abdomen is

inserted into a hollow in the former.

The clypeus slopes forward and is as broad as the median

quadrangle of eyes is long. Both rows of eyes are recurved ;

those of the front row are almost equidistant, the median 4 times

their diameter apart. The laterals, whose diameters are 23 times

FROM THE MONTEBELLO ISLANDS. 81

those of the median, are so placed that the line touching their

lower edges is the breadth of a median eye from the line

across the upper edges of the latter.

The median eyes of the rear row are the same size as the front

median, 24 times their diameter apart, 6 of same from the front

median and 8 from the laterals, whose diameter is 13 times that

of the former. This row is more recurved than the front, and

about 4 times the diameter of the median eyes wider than the

front row.

Hach eye is on a separate white tubercle, the side ones being

much higher than those of the median. The clypeus is as broad

as the area of the median quadrangle is long.

The mandibles are short and broad, kneed at the base, thence

divergent, the fangs being particularly short and weak.

The lip is straight at the sides, curving to a blunt point

anteriorly, longer than broad, and more than half the length of

the maxille, which are upright, the inner edges parallel and

straight ; from a rounded fore corner they slope downwards with

a straight edge, thence rounded at the back for about halfway,

where they curve in for the reception of the base of the palpi.

The sternum is shield-shaped, as broad as it is long, truncate

in front, flat in the middle, but sloping off in front and where it

narrows at the posterior end.

The abdomen is rounded in front, gradually widening to about

one-third of its length from the base whence, to halfway, the

sides are straight; from half its length it narrows to the rear

end, where it is just the breadth of the space occupied by its

spinnerets. The latter are quite terminal, of equal length, and

they havea short second joint. The superior are cylindrical,

about two-thirds the thickness of the inferior, which are conical,

flattened in front. The epigyne is of a horseshoe pattern, inside

of which is a long oval longitudinal depression flanked by two

shorter oval hollows in the upper half. The base is a transverse

semicylinder.

The femoral joint of the legs is moderately stout, but the

latter taper considerably and the tarsal joint is very fine. There

are Claw-tufts of flat bristles and a few scattered hairs on the

tarsus and metatarsus. On the under side of the tibia are four

pairs, and one odd one, of long spines, and four pairs of similar

long spines on the under side of the metatarsus; otherwise the

Jegs are smooth.

The palpi are short, the femoral joint incurved, the patella as

long as the tibia, and the distal joint, thickly covered with short

bristles, as long as the two preceding.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 12 | . Oe

Abdomen......... 44 iz

Proc. Zoou. Soc.—1914, No. VI. 6

82 MR. H. R. HOGG ON SPIDERS

Pat. Metat.

Coxe. Tr.& fem. &tib. & tars.

NGOS Stee ce nt. Il 4 2 24 2

2. I 3 3 Diy 5 aa Se

1 3 L Tees 3

| A, 4 2 1? Te es Bs

i 1 3 L lt = 7

Palpueieey Aes = = 5 aa — ai

Patella tibia:

This genus has been described from 8. Africa, Asia, and Japan,

but no species belonging thereto has previously been noted from

Australia, This single female specimen apparently conforms to

it in every point.

Family CLUBIONID4.

Subfamily Liocraninaz.

Group MirurcEs.

Genus Mrrurea Thor.

Mirurea Parva, sp.n. (PI. II. fig. 8.)

One male (T. H.). .

Cephalothorax pale yellow-brown in the middle and at the

margin, covered with pale yellowish-white hair; between these

areas on each side is a longitudinal darker yellow-brown streak,

with brown hair, reaching from the eye-space to the rear. At

the margin there are also darker spots, and a thick fillet of

yellowish-white hair projecting from the edge outwards.

The mandibles are dark yellow-brown for two-thirds of the

length from the base, paler anteriorly, with dingy yellowish-white

hair. The fangs are bright yellow-brown.

The lip, maxille, and sternum are pale yellow with similar

yellowish-white hair rather darker on the fringes,

The abdomen above is yellow-brown, with pale yeilow-brown

hair. On each side, reaching from the base to halfway, is a

brown streak and between these, two narrower, less distinct,

streaks, and brown blotches between the end of the side stripes

and the spinnerets. At the base are a number of brown bristles.

The under side is similarly coloured at the base and sides.

Beginning at the genital fovea and reaching nearly to the

spinnerets isa wedge-shaped area of black hair broadest anteriorly.

On this are two longitudinal rows of large white spots, and on

each side a clear white streak bounds the black area. The legs

and palpi are dark yellow on the basal part of the femoral joint,

getting paler towards the anterior joints, with yellowish-white

hair, nearly white scopule and claw-tufts, and yellowish-grey

spines.

FROM THE MONTEBELLO ISLANDS. 83

The cephalothorax is ovate, one-fourth longer than broad,

convex, thickly covered with coarse downlying hair. Round the

outer edge of the thoracic part is a thick fillet of hair extending

beyond the margin. ‘There is a long broad longitudinal fovea

reaching to the rear slope.

The rear row of eyes is recurved ; the median eyes one-third

of their diameter apart and their diameter from the side eyes,

which are on low tubercles and just slightly smaller.

The front row is straight viewed from in front, slightly re-

curved from above, all the eyes equal in size to the rear laterals

and their diameter distant from the rear median. The median

ave one-third their diameter apart and half that distance from

the laterals. The clypeus is not quite twice their diameter in

width.

The mandibles are strong aud convex, with long powerful

falces and two separated teeth on the inner margin of the falx-

sheath.

The lip is broader than long, straight in front, widening to the

base, and less than half the length of the maxille. The latter

are convex, straight on the inner side and at the apex, but

rounded at the corners and on the outer side.

The sternum is a broad oval, hollowed opposite the coxe,

and ending in a small point posteriorly well above the contiguous

rear coxe. It is thickly covered with coarse downlying hair in

the middle, upstanding round the margin.

The abdomen is oval, twice as long as broad. The spinnerets

ave terminal, with thick matted hair on the upper side, smoother

below. In the superior pair the conical second joint is two-

thirds as long as the basal.

The legs are thickly covered with long coarse hair, with

numerous short and some long powerful spines on the femora,

tibie, and metatarsi. There are two spines above on tibia iv.

On the under side of tibia 1.and 11. are three pairs of spines ; also

three spines on the inner side of tibia ii., two on tibiai. The

claws are short and weak. ‘There are thick scopule on all tarsi

and metatarsi.

The palpi have the femoral joint incurved, thinner than the

other joints, but broadest anteriorly. The patellar joint is

shorter and narrower than the tibial, which widens out in front

with an apophysis having a curved cusp on the outer corner, but

square on the inner.

The measurements (in millimetres) are as follows :—

Long. Broad.

6, She oe

Cephalothorax... 5 ie in front.

Abdomienienee-s 6 3

Mandibles ...... 24

84 MR. H. R. HOGG ON SPIDERS

Pat. Metat.

Coxe. Tr. & fem. & tib. & tars.

iG eSescesseor al z 54 6 5D Lee

Tee ae! 5A 6 Be ae 1 eS

Moe le At 5 pare ing

Me nes hae 7 aie Bid

TPRUN OI paeeoseee cosa 1 24 2 ake Sess ml

This differs from VM. lineata Thor., WV. gilva L. K.. M. age-

lina B. 8., M. occidentalis E. S., M. severa K.8., WM. ferina K.5.,

besides other points, in having a black area with white longi-

tudinal lines on the under side of the abdomen, and having darker

spots but no continuous line along the margin of the cephalic

part of the cephalothorax.

From WM. thorelli E. Sim., in having the front median eyes

smaller than the rear median and one cusp only instead of two

on the tibial apophysis of the male palp.

From I. maculata H. R. H., MW. whistleri KE. Sim., MW. impedita

E. Sim., and UW. catograpta E. Sim., besides numerous points

which will be gathered from the descriptions, in having three

spines on the inner side of tibia ii. and two spines on the inner

side of tibia 1.

Subfamily SPARASSIN#.

Group DELENE#.

Genus Outros Walck.

OLIOS CALLIGASTER Thor.

3 females (T. H.) non-adult.

1 female (P. D. M.) non-adult.

These specimens, despite the fact that none of them is fully

developed, are all larger than those measured by Thorell and

L. Koch from the eastern and southern parts of the Continent.

In other particulars they quite agree with the original

descriptions.

The under side of all the patelle and tibis is marked with

alternate stripes of brilliant silver-grey and brown instead of

partly yellow, possibly because they are younger.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 9 | : in front.

Abdomen......... 14 9

Mandibles ...... 43

q Pate Metat.

Coxe. Tr. &fem. & tib. & tars.

1WOGS sesuscase He 34 94 10 92 = 322

2. 34 10 104 10.23

3h 3 74 ar dies Ge

4, 3s 84 9 8i = 292

Pallpteeeee tes eee 14 4 3 Sos ile

FROM THE MONTEBELLO ISLANDS. 85

OLIos HERMITIS, sp.n. (PI. II. fig. 9.)

1 female (P. D. M.) non-adult.

2 females (T. H.) (1 ceph. only).

These specimens, none of which is quite adult, are very close

to L. Koch’s Olios (Sarotes) procerus from the east coast. They

differ, however, in the front median eyes being smaller instead

of larger than the rear median. The clypeus is as wide as the

distance between the front and rear median eyes—plus the

diameter of a front median. The lip is less than half the length

of the maxille instead of half as long, and the legs are more equal

in length, the fourth pair being equal to the front pair, and

the second only slightly longer. It is therefore worthy of being

made a new species.

The cephalothorax is yellow-brown, with a rather broken

yellow-brown marginal stripe on the thoracic part, a similar

horseshoe pattern of large brown spots nearer the centre with a

single spot between the forward pointing open ends. The eye-

space is black-brown, and there are two fainter brown spots

behind the rear row. The hair is fine and silvery white, but

brown on the spots. The mandibles are yellow with a brown

stripe on the outer edge reaching rather more than halfway from

the base, and the fangs are brown. On the inner edge of the falx-

sheath are three large teeth followed by one small one; on the

outer margin one large between two small. The lip, maxille, and

sternum are darker yellow with brown bristly hair.

The legs are orange, with three brown rings on the femur,

1 on the patella, 2 on the tibia, and 2 on the metatarsus of each

leg. The scopula on the metatarsus and tarsus are grey, and the

claw-tufts nearly black. There are two very long spines, one in

front of the other, on the under side of the tibial joint. The

tarsal claws are long, with about 10 pectinations on a straight

shaft bent at the anterior end, and the female palp-claw has

about half that number.

The measurements (in millimetres) of the largest (front pair of

legs only) and of a smaller whole one, are as follows :—

Long. Broad. _

Cephalothorax... 6 { 3 in front.

Abdomen......... 8

Mandibles ...... 3

Pat. Metat.

Coxe. Tr.&fem. & tib. & tars.

We es is... a3 is 2 94 114 Oe ea Be

J2by lovee eee cena aan 1 34 34 a eS

Long. Broad

O Sea ie,

Cephalothorax... 42 { 4 intone

Abdomen..... ... 5 34

Mandibles ...... 2

86 MR. H. R. HOGG ON SPIDERS

eats Metat.

Coxe. Tr. & fem. & tib. & tars.

IDS eeaateas 1 13 i 8 Oh ext BUDS

2 1 4 7 84 7 = 24

S11 GinG 7 Gn a0

dae pane iy 8 (ee

Pailricth Aleve ake Riise Yomi tou | ulouk cae Wanye

There are two quite young specimens haying the front row of

eyes straight and of equal diameter—the median pair being their

diameter apart, but only half that distance from the laterals.

Sternum yellow.

They do not seem to agree with any described species, but are

too young to found-one on.

Another still smaller seems to be the same, but the front

median eyes are apparently farther apart in comparison with

their diameter, and one-half that distance from their laterals.

Subfamily Micariina.

Group Micarig#.

Genus MonvTEBELLO, gen. nov.

This differs from Pecilipta Sim. in having the cephalothorax

only slightly less broad posteriorly than in its widest part between

the 2nd and 3rd coxee, whence it narrows to the front, which is

truncate. The clypeus distinctly narrower than the front median

eyes. The abdomen tapering posteriorly. The area of the median

eyes broader than long; the front median larger than the others ;

the rear row straight or slightly procurved. The 4th pair of legs

only moderately longer than the others.

M. TENUIS, sp.n. (PI. II. fig. 10.)

The cephalothorax is pale yellow-brown with a metallic sheen.

The mandibles greyish yellow on the basal half, yellower on the

anterior, fangs yellow. Lip, maxille, and sternum greyish yellow

with grey fringes on the former and pale grey hair on the latter.

The legs and palpi are pale yellow with almost white hairs, grey

spines, and dark grey claw-tufts. The abdomen is pale yellow on

the upper side with short fine dark, and some lighter grey hairs,

anda darker median longitudinal stripe on the posterior half ;

on the under side it is pale yellow all over.

The cephalothorax is truncate in front, longer than broad,

rounded at the rear, only slightly narrowing posteriorly from its

widest point between the 2nd and 3rd coxe. From this point it

also narrows anteriorly, where it is 2 of its greatest breadth. It

is convex, sloping rather steeply to the edge i in the cephalic part,

and to a flat mar ginal area at the sides of “the cephahc. There is

a short thin longitudinal fovea at the top of the rear slope.

There are no radial markings, the surface being quite smooth

and only a few hairs at the rear of the eye-space.

The rear row of eyes is slightly procurved, equal in diameter,

the median being three diameters apart and two of the same from

the laterals, and the same distance from the front median. The

FROM THE MONTEBELLO ISLANDS. 87

median-eye areais broader than long. The front eyes, 14 times °

the diameter of the rear, are the diameter of the latter apart ; they

are the same distance from the front laterals, the row being nearly

straight. The front and rear laterals are of equal diameter, 13 of

that distance apart. The rear median eyes are sessile, all the

laterals on slight protuberances and the front median still more

protuberant. The clypeus is only half as broad as a front median

eye.

The mandibles are straight on the outer side, slightly kneed at

the base, and as long at the front of the cepbalothorax as broad,

only diverging from each other anteriorly to the extent of the

slope of the falx-sheath. The fangs are moderately curved, slight

and fairly long; on the inner margin of the sheath are two jequiale

sized teeth, and on the outer, one equally long between two smaller.

The lip is broader than long, rounded anter riorly, straight at the

sides but narrowing slightly to the base, with the anterior margin

protuberant. It is not more than one-third the length of the

maxille. The latter are straight at the anterior margin, nearly

parallel at the inner and outer sides, but just rounded at the front

corners and widening near the base at the insertion of the palpi.

The sternum is ov Sh twice as long as broad, narrowing to a

point at each end. Tt is convex with deep depressions in the

margin between each pair of coxe; the rear pair of the latter are

contiguous and longer than the rest.

The abdomen is not quite twice as long as broad; it is rounded

in front, but not scutate, widest about the genital fold, whence it

narrows evenly to the spinnerets, which are quite terminal and

of equal length, the inferior pair contiguous, conical, with quite

short hemispherical second joint, the superior cylindrical with a

similar short rounded second joint. ‘There are no plumose hairs.

The legs are short and slender, the tarsal joints flat, the two claws

have 7 or 8 pectinations, a few long spines and bristly upstanding

hairs on the tibial and metatarsal joints, and claw-tufts of spatulate

bristles,

The palpi are inserted at the lower end of the maxille, the

femoral joint incurved and broadened at anterior end.

The measurements (in millimetres) are as follows :—-

Long. Broad.

j ie initromte

Cephalothorax... 24 Y

Abdomen......... 34 Do

Mandibles ...... 1

Pate Metat.

Coxe. Tr. & fem. & tib. & tars.

Te ssievisena |: b 2 Le 2 1 1 ==) OF

ae oy 1 2 as Ia — De

A, 2 2 2 Oe —v Ge

mlkan 1 3 1 eee)

Pallpigseceaas§ otis: eae 42 = 4 San ete a

There is one female specimen only (P. D. M.), and the

epigyne is not clear enough to draw.

88 MR. H. R. HOGG ON SPIDERS

Family Lycosip #.

Group LycosEa&.

Genus Lycosa Latr.

LycosA cLARA L. Koch.

A number of females, only one apparently adult, although they

are mostly larger than L. Koch’s specimens.

The epigyne is more like L. Koch’s drawing of that of Z. cris-

pipes, but it otherwise differs from the latter in too many points

to be confused with it, and agrees in all others with JZ. clara.

Moreover, the epigyne of the two forms are not very dissimilar

with the exception of the ogee-shaped anterior curve of the

latter, which 1s so unusual as to suggest that there may have

been some distortion in the specimen from which LL, Koch’s

drawing was made.

The measurements of the largest (in millimetres) are as

follows :—

Long. Broad.

Cephalothorax... 10 \ i ee

Abdomen......... 9 7 between 2 and 3 pairs of coxe.

Pat. Metat.

Coxe. Tr. &fem. &tib. & tars.

Legs ......... if 3 9 Js Oe

2 3 0) 83 y= 20)

Be 3 83 83 10 = 30

4. 3 10 10 1, = 35

Balpicste Sy ae AON 2 le 4 2-24 2 el

Family OxyYorip#.

Genus Oxyorss Latr.

OxyorrEs ? MuNDULUS L. K.

Four females, of which one only is adult. The eye-plan is the

same in all, and they have numerous flat hairs on the under side

of the abdomen, but the younger are all darker in colouring and

might be different.

They are rather close to several of L. Koch’s species, which are

not easy to distinguish. In pattern and size they seem nearest

to O. mundulus, O. amenus L. K., and O. variabilis L. K., to all

of which the epigyne might conform.

The measurements (in millimetres) are as follows :—

Long. Broad.

his

Cephalothorax... 3 { rs in front.

Abdomen.-....... 6 3

Mandibles ...... 1

FROM THE MONTEBELLO ISLANDS. 89

Pat. Metat.

Coxe. Tr. &fem. & tib. & tars.

ees ee 1 3 4 33 Ate er ieyoa

2 3 32 34 Ayes wig

3 3 22 21 OE ves BE

4 3 31 31 52 = 131

Railpr eNeee eee ra ey 1} lige

Genus Prucetra Thor.

PEUCETIA MARGARITATA, sp.n. (PI. II. fig. 11.)

The cephalothorax is pale yellow mottled with white and brown

upstanding bristles. The eye-space is black and dark yellow-

brown, with white downlying flat lanceolate hairs. Mandibles,

lip, maxilla, and sternum bright yellow, fangs of former darker

orange. The legs are bright yellow on all joints, with long grey

spines and upstanding brown bristles on brown roots. There are

short, fine, white scattered hairs on the femoral joints, short brown

hairs on the others.

The cephalothorax is longer than broad, rounded at the sides

and rear, narrowed in front. It slopes gradually from the sides

of the thoracic part, but steeply from the cephalic, with a nearly

perpendicular clypeus three-quarters the length of the eye-space.

The cephalic part is clearly separated by depressions from the

thoracic part, and there is a deep longitudinal fovea on the rear

slope. On the median line are three pairs of bristles with circular

roots.

The rear row of eyes is procurved, so that the uppermost points

of the laterals are on a line with the lowest part of the median.

They are equidistant, but the median are quite perceptibly larger

than the laterals, and their distance apart is rather more than

the diameter of the former.

The eyes of the second row (the laterals of the front row) are

the largest and most prominent of any, their diameter being equal

to that of the rear median and a front row combined. They are

this distance from the rear laterals. The front row (or front

median) are two-fifths the diameter of their laterals (2nd row),

their diameter apart, and the same distance from their laterals.

The clypeus is the length of the quadrilateral formed by the

rear median and 2nd row of eyes.

The mandibles are as long as the cephalothorax is broad in

front, conical and slightly kneed at the base, with scattered up-

standing bristles on the front and hair on the inner and outer

sides. The fangs are broad at the base, but short and weak.

The margin of the falx-sheath is smooth but with a fringe on the

outer margin.

The maxille are long and narrow, rounded anteriorly, and

parallel at the side, bending forward over the lip, which is twice

as long as it is wide halfway up, but broadens out at the base ;

it is rounded anteriorly and more than half the length of the

maxille,

90 MR. H. R. HOGG ON SPIDERS

The sternum is shield-shaped, straight in front, rounded at the

rear, where it is nearly as broad as in front, the rear coxe being

2 of their width apart.

The legs are long and fine, tapering anteriorly. They are only

sparsely furnished with short downlying hairs on the femoral

joints, and short bristly upstanding ones on the other joints.

There are five or six pectinations on the superior tarsal claws.

There are three pairs of long spines on the under side of all tibie,

two pairs underneath the metatarsi and about five shorter at the

anterior end of same.

The abdomen is broken in each specimen, but in one of them

the epigyne is intact; it is roughly similar to, but more elaborated

than that shown in L. Koch’s drawing of the only species pre-

viously recorded from Australia.

The measurements (in millimetres) are as follows :—

Long. Broad.

ho We

Cephalothorax... 3 1 x nL SE

Mandibles ...... li

Pat. Metat.

Coxe. Tr. & fem. & tib. & tars.

JL GCS boats 1 3 5 4} i lB

” 3 4 4 hye aa) 28

oth iaes alae SOT

A, 2 4 3 A

Aa nies hy ecenee eee i 1} 1 Byes Ba

Two females sent by Mr. Haynes.

These would seem to differ from P. albessens L. K. in the

lighter and brighter colouring, in having no row of dark spots on

the side slopes Toe he cephalothorax, in ‘the ereater length of the

legs compared with the cephalothorax (4 to 33), and the clypeus

not so long as the eye-space, and the more defined epigyne.

Family ATTID#.

Group MArpIssEz.

Genus Marpissa C. Koch.

MARPISSA RIDENS, sp. n. (PI. II. fig. 12.)

The cephalothorax is black-brown on the cephalic part, dark

yellow-brown on the thoracic. On the former and on a marginal

stripe are downlying white hairs interspersed with orange. The

side slopes and posterior end are bare; on the clypeus is a fringe

of coarse orange-coloured hair.

The mandibles are black-brown with a few brown bristles on

the basal half. The fangs are dark yellow-brown.

The lip and maxille yellow-brown, paler at the margins, with

dark grey fringes and brown upstanding hairs,

FROM THE MONTEBELLO ISLANDS. 91

The sternum dark yellow-brown with white hairs.

The abdomen on the upper side has a broad median area from

base to spinnerets of coarse white hair interspersed with orange

and upstanding brown bristles; on each side of this is a longi-

tudinal stripe of black hair reaching the whole length. ‘The

sides and under side are yellowish white. The spinnerets and

epigyne are dingy yellow-brown, with bright yellow inside the

chitinous ring of the latter.

The legs are pale yellow with white hair, brown spines, and

dark grey claw-tufts. The palpi are thickly covered with long

white upstanding hair.

The cephalothorax is nearly 13 times as long as it is broad,

straight in front, rounded at the rear, slightly narrowing from

about the middle to the front row of eyes. The cephalic ‘part 1s

flat above, sloping slightly to the sides, as does also the thoracic

part, which is rather more convex, but has a broad shallow

transverse depression at its anterior end,

The eye-space is broader than long, the rear row being narrower

by almost one-fourth than the cephalothorax at that point. The

small eyes of the second row are rather nearer to those of the

rear row than to the front laterals, and lie in a line between their

centres.

The front row is slightly recurved, the median eyes being close

together; the laterals, half their diameter, are clearly separated

from them and lie rather farther back. The clypeus is half the

breadth of the front median eyes.

The mandibles are short, flat and rather divergent, with

moderately long tapering fangs. There is one tooth, strong and

conical, on the inner margin of the falx-sheath, and two smaller

near together on the outer.

The maxille are upright, rounded anteriorly and at the outer

margin.

The lip, longer than broad, is more than half the length of the

maxilla, It curves inwards from near the front, but is nearly

straight at the end. ‘The front pair of coxe almost meet at their

bases. and with their trochanters cover the lower part of the lip

and maxille.

The sternum narrows to a point between the front pair of coxe,

broadens to its greatest width above coxe iii., and ends in front of

the fourth pair, which are close together.

The front two pairs of cox are parallel, pointing forwards at

an angle of 45 degrees from the median line, the rear two pairs

similarly pointing backwards at right angles to the front pairs,

the 2nd and 3rd being slightly separated. The rear cox are

longer than the others, which are all about the same length.

The front pair of legs are stouter than the others, the femur

being flat and club- shaped. The patella and tibia are longer

than the metatarsus and tarsus in all legs, the lattex joint shorter

than the metatarsus.

On the under side of tibia i. and ii. are three pairs of short

92 ON SPIDERS FROM THE MONTEBELLO ISLANDS.

spines, under metatarsus i. and ii. two similar pairs. On tibia ui.

and iv. is a single pair, and a bunch at the end of the meta-

tarsus of same. The femoral joint of the palpi is incurved and

broadest at the anterior end, the tibia longer than the patella.

The abdomen is rather longer than the cephalothorax, truncate

in front, straight at the sides for 3 of its length, whence it narrows

to an obtuse point; the epigyne is a chitinous oval ring thickest

at the posterior end, granular inside, with two small club-shaped

protuberances therein near the lower end.

The measurements (in millimetres) are as follows :—

Long. Broad.

Tole § y

Cephalothorax... 3°1 | ae “Un ant

Abdomen......... 3°5 eal

Mandibles ..... ‘ 3

Pat. Metat.

Coxe. Tr. & fem. & tib. & tars,

SSPE: lt 3 4 2 if = 53

3 3 1 ES 7

Bt Mes 1 Paige ease aes

g 3 . ‘ i} = 1

aang 16 2 ii = 5

Zola 3

AA Orcs ere, Sencar le - i 4 tf, = 13

This may be distinguished from any of the recorded Australian

species by the black lines on the back of the abdomen and the

pattern of the epigyne.

Another smaller specimen, apparently of the genus Marpissa,

has contiguous front coxe longer than the rest, weaker mandibles,

and tarsus i. as long as the metatarsus, but is too broken to

describe.

EXPLANATION OF THE PLATES.

Puate I.

Fig. 1. Tetragnatha angulata, sp.n.(X 2). a. Mandible from outer side. 6. Man-

dible from inner side. ¢. Epigyne.

. Nephila meridionalis Hogg, var. hermitis, nov. (nat. size). a. Underside

of abdomen. 6. Profile. ec. Eyes.

- Argiope haynesi, sp. n. (nat. size). a. Underside of abdomen (xX 2).

b. Epigyne. c. Eyes.

. Larinia montagui, sp. n. (X 2). a. Eyes. 6. Epigyne.

. Araneus reversus, sp. n. (nat. size). a. Male (non adult), X 2. b. Eyes of

female. ec. Epigyne. d. Epizyne from side.

. Gasteracantha minax Thor., var. hermitis, nov. a. Epigyne.

oe ist) bo

for)

Prare Ti

Fig. 7. Dieta isolata, sp. n. (X 2). a. Byes and mandibles. 6. Epigyne.

8. Miturga parva, sp.n. (nat. size). a. Eyes. 6. Underside of abdomen (X 2).

ce. Male palp.

9. Olios hermitis, sp. n. (nat. size of immature). a. Eyes. 6. Lip and

maxille.

10. Montebello tenuis, gen. et sp. n. (X 2). a. Eyes. 6. Mandibles. ec. Lip,

mavxille, and sternum.

11. Peucetia margaritata, sp.n.(X 2). a. Byes. 4. Epigyne.

12. Marpissa ridens, sp. n. (X 2). a. Byes. 6&6. Epigyne. c. Lip, maxille,

Jett mandible, and first pair of cox.

ON THE NESTS OF PSEUDOSCORPIONES. 93

7. On the Nests of Pseudoscorpiones: with historical

notes on the Spinning-Organs and observations on the

Building and Spinning of the Nests. By H. Watts

Kew, F.Z:S.

[ Received January 5, 1914: Read March 3, 1914.]

INDEX.

Ethology : Page

Pseudoscorpiones: moulting-, brood-, and winter-nests. 94,

Method of building and spinning the nests: Chelifer

ENO KOLA aN enna aaaen ation Addo sh aauee bare aeeoe pre neebegic a Rime tue a COPD

Structure :

The spinning equipment : historical account............... 99

External spinuning-organs of the chelicerx:

The False-Scorpions’ ability to spin was denied by some of the

older writers, e.g. by Frisch (1), and doubted by others, e.g. De

Théis (4), but it is now known that they construct nests in part

or wholly of silk from their own bodies. Balzan (15) named a

species Chelifer nidificator; but the nests are not peculiar to

particular species or groups. They are known for Chelifer,

Chewridium, Garypus, Garypinus, Olpiwm, Obisium, and Chtho-

mius: genera which represent both main divisions of the Order

—Panctenodactyli and Hemictenodactyli—and the four main

families—Cheliferidee, Garypide, Obisiidee, and Chthoniide; and

it is thus probable that these structures are common to all

Pseudoscorpiones *.

The purposes for which the nests are made are known—at

least satisfactory statements on this head exist—and the nests

themselves, which have often a more or less elaborate covering of

extraneous matters, have been described with more or less detail

and accuracy by several writers. When one enquires how they

are made, however, no answer is forthcoming. The manner in

which the extraneous matters are collected and arranged is

unknown; and as regards the spun-tissue, the position of the

silk-glands and external spinning-organs was long mis-stated ;

and the subject is still surrounded with uncertainty. With the

exception of a statement by Menge (6), which has proved to be

mistaken, no description of the animals’ methods is known; and

it does not appear that the drawing out of the silk has ever been

witnessed.

It is proposed here to give a brief re-statement of the subject,

and to record some direct observations.

* The construction of nests is the only use the animals are known to make of

their spinning. They have been said, by this means, “to envelop the eggs in a

cocoon ” (17). But no False-Scorpion covers its eggs with silk; and since the eggs

are not laid, the egg- and brood-pouch remaining attached to the mother, the thing

is impossible.

94 MR. H. W. KEW ON THE

IVE.

As already stated, the purposes for which the nests are made

are known; but they are inadequately stated in most of the

books. The animals thus enclose themselves for moulting, for

brood-purposes, and also in some cases for hibernation. Nests

for moulting are probably universal and made by all individuals

of both sexes. They protect the animal during ecdysis and the

periods of helplessness which precede and follow it. Those for

brood-purposes, made by females, are probably universal in that

sex; but in exceptional cases they are slight and easily over-

looked. They protect the distended animal and attached egg-

and brood-pouch, and also the young, which remain for a time

enclosed with the mother*. Nests for hibernation are probably

less general. It is certainly true of some Beers) perhaps of

many, that most individuals of both sexes rest in nests during

winter. In these they are protected no doubt from cold ; ant

perhaps also from moisture, which might freeze around them

with harmful results.

aisle:

References to the nests in published papers, and my own notes

on the subject, are too numerous to be collected here. It may be

permissible, however, to pass In review the main groups and to

refer more particularly to a small number of species. Godfrey

(29, 36) is the author who more than anyone else has attended to

these structures.

Of the families of Panctenodactyli—Cheliferide, Feaellidee, and

Garypide—the last two are foreign to our tauna and their nests

ave unknown to me. Those of Feaellide do not appear to have

been observed. It is otherwise with Garypide. Lucas, in

1849 (5), wrote of Olpium pallipes Liue., that it lives under or

in the crevices of stones where it constructs “une petite coque

sins issue, d’un tissu soyeux, serré, revétue a l’extérieur de

grains de sable et de parcelles de terre.” Garypinus capensis Ell.

is stated by Godfrey (36) to make nests, between flakes of bark

of trees, ‘‘of silk only without any covering of dust or specks of

wood.” According to Becker (10) the nest of the large Garypus

e the Mediterranean—Garypus litoralis L. Koch of this author—

s ‘une coque de soie hermétiquement fermée.” He refers also

i similar silken nests of Garypus minor L. Koch. These last

are more particularly described by Barrois (22), who found them

* Bouvier (21), writing of the nests of the large Garypus of the Mediterranean,

has indicated that they contained brood-pouches developing in the absence of the

mother; and some prominence has been given to this statement by With (27).

Further observations, however, convinced Bouvier that he was mistaken; the

mother had doubtless escaped, leaving the brood-pouch behind, subsequently to the

collection of the nests. It is possible, however, that some species, e.g. Cheiridium

museorum Leach, may cast off the brood-pouch and leave it in the nest after the

larvee have ceased to suck; cf: Godfrey (29). To recur to the Garypus, Becker (10)

tells us that he found “les deux sexes réunis dans une coque de soie’”’; an observation

which stands alone and deserves investigation.

NESTS OF PSEUDOSCORPIONES. 95

on the shore at Villefranche attached to the under-surface of

stones: ‘‘ Chaque loge se compose dune espéce de petite capsule

de forme arrondie tapissée intérieurement par un feutrage blanc

auquel adhérent extérieurement de menues parcelles de terre ou

de débris végétaux.” He adds that they were completely closed,

the animal apparently remaining imprisoned all the winter *.

Tn Cheliferide the nests have frequently been mentioned, and I

have had opportunities of examining those of several species.

The earliest reference for ‘“ Chelifer” is that of Hermann,

1804 (3), who mentions the occurrence of one of these animals

“dans un follicule soyeux, enduit de poussiére et attaché a une

paroi par un de ses cdtés.” Of the four sub-genera of Chelifer,

there is a mention of nests of Withiws by Godfrey (86), who

refers without particulars to those of C. simoni Balz. In

Atemnus, the same author found C. feae Hil. “in a very roomy

nest on a gum-tree”; and to this sub-genus belongs Balzan’s

O. nidificator (15), found under flakes of a tree-trunk, in “ nidi,

in forma di cellule contigue, di una sostanza cerosa, biancastra,

simile alla seta di certi ragni.” In Chelifer s. s. and Chernes, I

will refer to two species of the former C. cancroides Lin. and

C. latreillii Leach, and to two of the latter C. eyrneus LL. Koch

and ©. cimicoides Fabr.t. C. cancroides Lin., which is well-

established but doubtfully indigenous with us, lives in stables

and such-like places. Godfrey (29) found its nests in a loft in

Glasgow in narrow crevices in old harness. He describes them

as ‘closed elliptical rings of dust particles, within which was the

inner silken lining appressed throughout the extent of both upper

and lower surfaces against the leather.” They belonged evidently

to immature individuals, and some of them contained cast-skins.

A few years ago I examined a colony of this animal in stables at

Grays. Between the double boards of the stall-partitions were

spaces separated into compartments by the uprights and com-

pletely closed in; and on some of these boards being removed

many nests were found on their inner surfaces. These spaces of

course were much too wide to admit of the nests being attached

both above and below; and, as is usual in such cases, the tissue

of the floor only was attached to the wood and was continued

above as a free convex roof. They bore externally a partial

covering of extraneous fragments. Most of them were in-

habited; and in at least two were found females with the

brood-pouch attached. C. latreillii Leach is a maritime species

with us, occurring along our eastern shores. On those of Fife-

shire and East Lothian it is found in rock-crevices; and

Godfrey (29) states that its nests, used for moulting and also,

he believed, for hibernation, are large and made generally

* The quotation is from Dr. Barrois’ well-known “Mémoire sur le développe-

ment des Chelifer,” the subject of which was in reality Garypus minor L. Koch.

Iam indebted to the author for specimens of the animals and to M. Simon for

obligingly examining them.

+ For the nomenclature of British and Ivish species referred to in this paper,

cf. Kew (33).

96 MR. H. W. KEW ON THE

between two closely placed pieces of rock; they are formed,

he says, of earth with a silk lining and with a silken floor

and roof attached to the rock. In several English counties

the animal inhabits the sand-dunes of the coast; and under

these conditions I have often observed it, especially in Lincoln-

shire, where it is abundant on the fine range of dunes which

extends from the Humber to the Wash. In such places it

makes its nests for the most part in old sheathing-bases of

marram-grass (Ammophila arenaria) and sometimes under bark

of maimed stumps of Hippophaé rhamnoides and Sambucus nigra.

Those made for brood-purposes are frequently slight in texture,

the animal being distinctly visible within. Of nests for hiberna-

tion I am not able to say anything not having observed the

animal in winter. The moulting-nests, however, are of the

usual dense tissue: and when in relatively wide spaces they

have usually a free roof. In such cases and whenever there is

any considerable expanse of free tissue there is usually a regular

covering of grains of sand. C. cyrneuws L. Koch and C. cimi-

coides Fabr. are forest species found under rather close-fitting

bark of dead or partly dead standing trees. I have already given

notes of the nests of the former ohscmrat f in Sherwood Forest (26),

and more recently have had many opportunities of observing

them in Richmond Park*. The brood- and winter-nests are

surprisingly large, even for this Jarge species, having frequently

a diameter of 10 mm. They are built usually in narrow crevices,

and thus they do not as a rule exhibit a free roof. This feature

is commonly seen, however, in the smaller moulting-nests. The

free membrane has usually a covering of woody fragments , which

is often dense and beautifully regular. The nests of C. cimicoides

—smaller in accordance with the smaller size of the animal—

have been observed by me in many places, and particularly in

Epping Forest, where they are readily found under the bark of

the old pollards. The brood-nests are apt to be slight and in-

formal in this species; but the moulting- and winter-nests are

ordinarily stout and often exhibit a free roof; and they are

remarkable for the density and regularity of the almost invari-

ably complete covering of woody fragments. To the same family

belongs Cheiridium, whose nests are unusually small and flat.

Cheiridium museorum Leach occurs, ¢.g., behind the boards of

old barns where the nests are often crowded together in great

numbers. They have a diameter of about 2mm.; and the roof,

which is very slightly convex, has a close covering of minute

miscellaneous objects. The nests of Cheiridium ferwm Sim. are

still smaller and scarcely, if at all, convex ; they are pearly white

and quite destitute of attached particles. This at least was the

condition of a number sent to me by Godfrey, who found them

under flakes of pine-bark in Brittany. Subsequently (36) he

* Tam indebted to His Majesty’s Office of Works, and to Mr. 8. Pullman, the

Superintendent of the Park, for the permission and facilities necessary for observing

and collecting in this place.

NESTS OF PSEUDOSCORPIONES. Si

found the same animal in South Africa with many nests under

bark of gum-trees. The structures were similar to those above

mentioned, of white silk only, and very conspicuous.

In Hemictenodactyli—families Obisiide and Chthoniide—the

nests of both Obistwm and Chthonius ave known. ‘The first note

for Obistwm is by Menge, 1855 (6), who found a brood of young

“‘ mit der alten Mutter in einem halbrunden Gespinnst zwischen

zwei Blattern.” Low (7) records a female with brood-pouch in a

snail-shell; of which the mouth “mit einem dichten, weissen,

homogenen Spinnengewebe vollig verschlossen war.” According

to Simon (9), O. gugorum L. Koch in the frozen regions of

the Alps constructs a nest ‘ presque arrondie et sans ouvel-

ture, dont le tissu agglutinant se recouvre de terre et de

brindilles végétales.” The common O. muscorum Leach was

observed in detail in Scotland by Godfrey (29). The favourite

site for its nest is the under-surface of a stone, but other situ-

ations such as the face of a rock covered with herbage or a bed

of moss on a tree-stump may be selected. The structure is

usually strongly domed. It consists externally of earth, earth

and sand, or earth and rotten wood, and internally has a close

firm lining of silk, which is continuous, as usual, not only over

the interior of the built parts but also over the enclosed surface

of the object to which the nest is attached. In the south-west

of Ireland there is a larger species, O. carpenteri Kew (82),

whose nests [I found abundantly under loose fakes of ragged

outer-bark of Arbutus wnedo. They are larger than those of

O. muscorum, and differ in having no regular covering of earthy

or woody fragments. The dense white spun-tissue of the nest

is in fact usually quite free from extraneous particles. Both

moulting- and brood-nests occurred on the trees; and in

addition, many of the former were found in rock crevices.

O. maritimum Leach, another large species, lives below high

water-mark on the sea-shore, where it occurs in deep-seated

rock-crevices and under large embedded stones. Its nests—

first mentioned by Ferronniére (24)—have been observed by

Godfrey in Scotland (29), by Jackson in the Isle of Man (28),

and by me in Devon and Cornwall and in Iveland (82, 34). They

occur in the situations just indicated and, as in the case of

O. carpenteri, the dense white spun-tissue has no coating of

extraneous matters. Spun amidst surroundings constantly moist,

they are stout and tough; and, contrary to what is usual in

these structures, they are not always intimately attached to the

surfaces on which they rest. Thus the nests, together with

the enclosed animal, can be removed without injury. When

thus removed they have the character of a complete bag of silk,

and are sufficiently impervious to retain the enclosed air after

long submergence in spirit. Finally, for Chthonius we are in-

debted to Godfrey (29) for notes on Chth. rayi L. Koch and

Chth. tetrachelatus Preys. He describes the nests as built up

usually of particles of earth and other chance objects, aud lined

Proc. Zoou. Soc.—1914, No. VII. 7

98 MR. H. W. KEW ON THE

with silk, or less commonly formed in holes in stones, ete., the

cavity in that case being lined with silk and the entrance closed

in with a film of the same material *.

LiVic

The nests throughout the Order, and for whichever purpose

they are made, are less divergent in character than might be

supposed from the foregoing details. Their essential features

are everywhere much the same. ‘he completed nest is a closed

cell of silk with or without a complete or partial covering of

extraneous matters externally. From this the animal has to

cut or break its way out when it wishes to emerge. Viewed

from above, the structure is roughly circular; but in other

respects its form varies with the build of the animal and with

the nature of the habitat. In the case of the more flattened

animals which inhabit crevices the structure is often compressed.

The silk in these circumstances is often spread over and attached

to solid surfaces both above and below; and the films thus

attached, which form the floor and roof of the nest, are con-

tinuous with and connected together by a circular wall. At

other times, in spaces relatively wider, the film of the floor only

is thus attached, and this film is continued above as a free, more

or less convex, roof. In the case of less flattened animals living

among stones, in vegetable débris, etc., the floor may be similarly

attached and be surmounted by a pronounced dome; or the nests

may be made in roundish cavities, or more frequently attached

here and there to surrounding objects, and their form is then

roughly globular. In all cases, however, there is variation from

individual to individual, the animals enclosing themselves in con-

venient spots of varying character. The creature is always rather

closely surrounded, though with ample room for free movement,

and thus the differences in the size of the nests follow rather

closely those of the makers. In the work of adult individuals

there is a range of diameter from about 2 mm. to 10 mm. or

more. ‘The external covering of extraneous matters when it

exists consists for the most part of earthy or vegetable frag-

ments. It may be complete or partial, dense or sparse, and

occur over the unattached parts of nests of whatever form.

Some such covering is characteristic of the nests of many

species; but in certain others it is invariably absent. The

fragments are never over-spun, that is to say, bound on by

threads passing over them, but they are always firmly attached

to and form part of the structure. They never affect the interior,

which is always free from foreign substances and smooth. The

spun-material has the character of a thin dense whitish tissue,

presumably largely impervious to moisture, and opaque, or

* Godfrey mentions that some of the nests of Ohth. tetrachelatus had within

them a second “ silk cocoon of exquisite texture and quite separate from the first

lining.” This might result, perhaps, from the re-utilization of old nests. But

Godfrey saw so many of these “ double linings ” that one can scarcely dismiss the

subject with this suggestion.

NESTS OF PSEUDOSCORPIONES, 99

nearly so. This tissue, unlike that of many spiders, does not tear

with a distinct floss. Its consistency is comparable with that of

thin silk-paper; and one would not suppose, even on exam-

ination with a strong lens, that it was composed of separately

spun threads. With more ample magnification the construction

becomes more or less obvious; but it is difficult to say what the

arrangement is. Threads in extraordinary number and closeness,

and most of them extremely fine, are crossed and re-crossed and

turned about in irregular confusion. They appear to have been .

brushed on by long-continued effort, and no doubt in a viscid

condition since they have coalesced to a large extent. The com-

pleted tissue is entirely without interspaces. When incomplete,

threads of varying strength, in a more or less open, irregular

meshwork, ave observable *.

From what part of the animal does the material for this tissue

proceed ¢ It was supposed that the glands were in the abdomen

and that their ducts opened near the genital aperture in numerous

separate spinnerets. This view was started by Menge, 1855 (6);

and it remained unquestioned for more than thirty years. Even

after Croneberg had shown it to be wrong, restatements of it

continued to appear, not only in text-books, but in memoirs

dealing specially with this Order: ef. Cambridge, 1892 (17). It

was in 1887-8 that Croneberg (12, 14) showed that the glands

indicated were really accessories of the genital system, nail that

the supposed spinnerets were merely part of the bristle-armature

of the external genital-area. At the same time Croneberg found

that in Chelifer (Chernes) there were glands in the cephalothorax

with ducts opening in the chelicere ; and Bertkau (18), working

during the same years and independently, made similar discoveries

in Obisium. Both authors concluded that these were the real

producers of the silk; and all that was required was confirmation

from observations on living animals. Such observations, however,

were not made; and in the meantime various considerations have

confused the subject ; so that, as stated recently by Godfrey (29),

the spinning in this Order is still surrounded with uncertainty.

Menge was well known to have supported his anatomical investi-

gations with detailed descriptions and figures; and apart from

these he had had living animals under observation. He had

even described the movements of an individual which made part

of a nest under his eyes. Croneberg and Bertkau, on the other

hand, had arrived at their conclusions on morphological grounds

alone. On such grounds, moreover, Supino (23) had dismissed

these conclusions, regarding the organs in question as a poison -

apparatus, and falling back as regar ds spInning-organs on a view

* The only previous note on this tissue known to me is by With (25). He

examined nests of Chelifer sculpturatus Wewis, and was surprised to find that the

threads were not independent but fused, so that a complicated system of thinner

and thicker threads was formed. He adds that the structure. was difficult to

explain ; and that the newly formed threads had perhaps fused before drying.

100 MR. H. W. KEW ON THE

almost identical with that of Menge. Further, Bernard (18) had

suggested that certain structures, which he believed himself to

have discovered on the ventral face of abdominal somites V. to

XI., were possibly the openings of spinning-glands. We have

seen, however, that Menge was certainly wrong. With (27) has

already stated that Supino was mistaken ; and the same remark

applies to Bernard, whose misconception was ridiculed at the

time by Hansen (20) and soon abandoned by the author himself.

But there were other difficulties. Ducts had certainly been seen

in the chelicere, but 16 was not clear that their openings had

been fully made out; and Hansen had examined the supposed

place of disemboguement in Obisiwm muscorwm Leach with

results which he was unable to regard as satisfactory. He may

perhaps have had before him an adult male in which the spinning-

function had degenerated *. However this may be, he seems to

have regarded Croneberg and Bertkau’s conclusions as justifiable ; :

“but it ought to be sane! if the animals actually do spin

with these organs.” So also With (27) has concluded that

spinning is at least one of the functions of the chelicere ; but

proof of it was still wished for.

The required confirmation of the correctness of Croneberg and

Bertkau’s conclusions is afforded by observations now recorded.

The spinning is done by the chelicerze. ‘These appendages are of

two segments: a hand prolonged into a fixed finger, and a

movable finger articulated to the hand; and they are provided

with special structures, of which one is entirely and another

partially or entirely comb-like. The whole appendage is rela-

tively small in Panctenodactyli, and relatively large in Hemi-

ctenodactyli; and several differences in the special structures are

characteristic of these main divisions. In all Panctenodactyli

the movable finger is provided, on its outer margin just before the

apex, with a small, almost transparent, more or less flexible, pro-

jecting structure known as the galea. In Hemictenodactyli this

structure is present or absent ; imal in its absence the hard chitin

of the finger, in exactly the same position, is raised to form a

small, more or less convex, laterally compressed tubercle. Crone-

berg’s researches in Chelifer (Chernes)—repvesenting the first of

these main divisions—showed that the ducts from the cephalo-

thoracic glands passed into the hand of the chelicera, four or five

into each, and thence into the movable finger, which they tra-

versed to near the apex, where they entered the galea. Within

the galea they distributed themselves into the small branches of

that structure, and at the terminations of these branches they

opened. Bertkau found in Obisiwm—representing the second

main divis:ion—that the ducts similarly traversed the chelicere

* Bertkau states of Obisiwm that the glands, which were well developed in all the

females he examined, were absent in some of the males, though the ducts remained

in the chelicere.

NESTS OF PSEUDOSCORPIONES. 101

to a point identical with that reached by those of Chelifer

(Chernes). There was here no galea; but the ducts entered

the hard tubercle replacing that structure, and on or near the

margin of this tubercle they opened. The number of ducts in

each chelicera was about ten—or, according to one of the later

papers of Bernard (19), about seven; and one may note that on

the tubercle of O. muscorwm Leach, examined by Hansen, six

minute openings appeared to exist. It is thus established that

the galea and the tubercle which may replace it are correctly

regarded as the external spmning-organs in this Order*. These

structures bear no resemblance to one another; and no inter-

mediate condition is known. The galea is obviously a sheath

within which the ducts are carried forward beyond the apex of

the finger. Its presence or absence does not appear to be asso-

ciated with differences in the spinning or resulting tissue.

Assuming Panctenodactyli to be the older group, it seems that

the increasing size of the cheliceree in Hemictenodactyli may

have rendered this projecting organ dispensable. The structures

ave always well developed, whichever one is present, in the

young of both sexes and in adult females; and this no doubt is

in relation to the spinning of moulting- and brood-nests. In

adult males they may be fully developed or degenerate. In the

former case it will be found, I believe, that regular winter-nests

ave spun; while in the latter case presumably the male spins but

little or not at all after arriving at maturity. The character of

both structures differs throughout the Order from species to

species. Among the forms assumed by the galea the most com-

pleated is that in which it is branched from a short stout base,

the branches being re-branched, so that the shape of the organ

recalls that of the antler of a stag. More usually it has a long,

rather stout shaft, with about six small, simple branches distally.

But it may be trifid or merely styliform. The tubercle may be

rather high with a convex outline, or lower or longer and more

flattened.

- The other structures of the cheliceree, the combs, etc., have

nothing to do with the spinning f.

* Thorell, 1883 (11), gave prominence to spinning (and chelate appendages) in the

name “ Chelonethi,” with which he proposed to replace “ Pseudoscorpiones.” ‘The

spinning-openings were then supposed to be in the abdomen ; but afterwards (1890),

with references to Croneberg and Bertkau, he remarked on the increased propriety of

the name, observing that the creatures not only possess chelee and spin but spin with

the chele (of the cheliceree). At the same time he proposed the name “ procursus

textorius”” for the galea (16); and we find that Ewing (85) has already written

** spinneret ’ for both galea and tubercle.

+ It has been stated repeatedly that these combs—the serrula and lamina interior

—manipulate the silk; but im reality they are not concerned in any way with the

spinning-work, never coming into contact with the threads or with the spun-tissue.

The mistake is attributable to Bernard (19), who converted into a direct statement

an Ingenious but incorrect suggestion of Croneberg’s. To these structures another

supposed silk-combing organ, said to be like the antenna of a Lamellicorn beetle,

has been added by Shipley (80) and Warburton (81). This last is one of the inven-

tions of Stecker (8; cf. Hansen 20); nothing like it exists in nature. The so-

called “flagellum,” which occurs in the position indicated, is merely a row or tuft of

peculiar bristles; it is of unknown use and certainly not connected with spinning.

102 MR. H. W. KEW ON THE

Vir

There remains the enquiry how are the nests built and spun;

that is to say, how do the animals collect and fix the extraneous

matters of the exterior, and how do they fabricate the spun-

tissue ?

Godfrey’s field-work showed that the external coating was the

first part of the work, the silk lining being produced afterwards

and necessarily from within. This was evident from inspections

of nests in various conditions of incompleteness; but the animals

were not seen at work. Tio make the required investigations in

the open was scarcely possible. It was necessary obviously to

have captive individuals under observation ; and this in con-

ditions favouring the undisturbed performance of their functions

and at the same time permitting prolonged watching. Several

naturalists, from Résel 1755 (2) onwards, have kept the animals

in captivity ; and it has usually been found that nests were con-

structed; but Menge alone makes mention of the animals’

procedure, He placed a Chelifer (Chernes) in a glass vessel and

discovered next morning that a nest had been commenced. He

found the animal still busy, by continued movements of the body

somewhat like those of the spider Clubiona, increasing the thick-

ness of the web; and he claimed from this to have seen the

spinning. That the animal was so engaged there is no doubt ;

but since Menge was mistaken as to the position of the spinning-

organs it is evident that no precise observation had been made.

The writer’s attempts to watch the animals in captivity com-

menced in 1904; and since that time one or more species have

usually been under observation. They were housed in small flat

eases of cork and glass, forming cells with an area of three square

inches and a height of from a quarter to an eighth of an inch ;

and in these they lived in health for Jong periods. Their sur-

roundings were made as natural as possible, and the nests they

constructed were in most cases quite like those made in their

natural habitats. The creatures were examined under moderate

magnifications, the cells being placed bodily on the stage of the

microscope. Such examinations were necessary from time to

time; but much could be made out with no other aid than that

afforded by a good hand-lens. The species whose nest-making

was observed in detail were two of Panctenodactyli, Chelifer

cyrneus lL, Koch and Chelifer latreillii Leach; and one of

Hemictenodactyli, Obistwm muscorum Leach.

Chelifer cyrneus LL. Koch—belonging to the subgenus Chernes,

and eyeless—was the subject of most of the observations. It is

a large heavily-built species; and the galea is long-shafted,

equally developed in both sexes, and provided distally with six

small branches *. The specimens were obtained from narrow

* These branches are less easy to count than might besupposed. For the present

species, Tomosvary gives four and Schtschelkanowzeff five; yet there are undoubtedly

SIX,

NESTS OF PSEUDOSCORPIONES., 103

spaces under the bark of partly dead oaks; and the cells in

which they were placed were provided with strips of decayed

wood from these trees. The arrangement was such that narrow

spaces were left between the strips of wood and the glass of

the cells. Colonies thus established—strengthened by fresh

importations from time to time—-were under observation for

two years; and at the time of writing, two cells contained

together eleven nests: some on the floors of the cells, with a

complete convex roof, and others of more compressed form in

the spaces between the wood and the glass.

At various seasons certain of the animals, either young or

adult, became distended—no doubt from accumulated nutritive

matiers—and it was in this condition that they enclosed them-

selves either for moulting, for brood-purposes, or for the winter.

Such an individual restlessly perambulated the cell, investigating

the corners and crevices, every now and then picking up with

the palp-fingers and removing dead insects and other scattered

objects; and occasionally, at such times, it detached fragments

from the wood, taking hold of them with the palp-fingers and

using considerable force. It was evidently prospecting for a

position for the nest. At length, having decided on one, it

was soon at work on it with the chelicere, and usually by the

following morning or evening it had arranged round itself a

complete ring of fragments of wood, ete. It was now unsafe

to abandon the post of observation, for the animal got to work

with rapidity and within about twelve hours had generally

completed the external coating or framework of the nest, the

builder being then—except in those cases in which the glass

formed the roof—entirely concealed from view.

The manner in which the animal constructed this framework

and enclosed itself within was the subject of numerous obser-

vations. But the procedure may be indicated by taking the

case of the making of a brood-nest with a complete convex

roof, the observation of which extended almost from beginning

to end. The distended builder was restlessly moving about im

the morning, and in the evening of the same day was found

to have taken up a position on the floor of the cell, where it

had already surrounded itself with a ring of fragments. These

were of wood and cork and mostly small—sawdust from the

borings of Dryocetes villosus for the most part—and they were

all definitely placed in position and seeured from within to the

floor and to each other by threads of silk. They formed already

the beginnings of the narrow circular wall of the nest. This

was at 6 P.M., at which time the animal was within the circular

space and full of business. For the continuation of its work

it would require evidently a quantity of material, much of which

would have to be procured from a distance. Coming up close to

the wall, the animal extended the palps over it and felt about for

fragments on the floor beyond. Finding some, it withdrew and

proceeded with its work; or, failing to find any, it stepped over

104 MR. H. W. KEW ON THE

the wall and explored the floor farther afield. Finding the

necessary supply without going far, it returned to the nest by

stepping backwards, going in over the wall abdomen first ; or,

more often, having to go to a distance, it returned with a forward

locomotion, re-entering the nest head first. The fragments were

found and picked up with the palp-fingers, whence they were

immediately transferred to the chelicere, the animal never

carrying material home in the palps. Except in the case of

large fragments it was not satisfied with one, but picked up

a number in succession, transferring each as found to the

cheliceree, its actions while thus engaged recalling those of a

bird accumulating nest-materia!s in its bill. The smaller frag-

ments were received between the fingers of the chelicere, the

fixed fingers going over and the movable galea-bearing fingers

under them ; and as the materials accumulated they appeared to

be attached together by silk from the galee. Larger fragments,

while always kept in contact with the chelicerz, and apparently

more or less attached with silk, were sometimes supported in

part by the hand or other segments of the palps, or even on the

dorsal shield of the head; and in this way was brought in at

least one fragment nearly as big as the animal itself. At the

conclusion of each collecting expedition the animal returned

impatiently to the nest; but once within, it turned about and

deliberated before placing the materials. At length, running

the burdened chelicerze into some part of the top of the wall, it

managed thus to release the fragments ; and it now got to work

at once, attaching them together and to those already placed

by dabbing silk on their inner surfaces and stretching threads

from one to the other. During this proceeding the palps were

brought round so that the great hands and fingers of these

appendages were close to the inner surface of the wall, the

animal evidently directing the work by the sense of touch which

the hairs and perhaps other sense-organs give to these parts.

Occasionally it adjusted the fragments by propping them with

the hands and fingers of the palps ; and sometimes, but rarely, it

held them between these fingers while the first few threads were

being applied by the chelicere. Apart from this, it constantly

felt with the palps both the inside and outside of the wall, as if

to ascertain how the work was progressing. One was parti-

cularly interested to see how it extended the palps over the

outside for this purpose, its actions bringing to mind those of a

bricklayer giving little taps to newly-placed work. There was an

air of satisfaction about its behaviour, and with reason, for it was

wonderful to see how the wall grew and began to assume the

proper curve for the roof. From time to time, as soon as each

collection of materials had been placed in position, off the animal

went for another supply; and this continued fetching of fragments

and building them in was maintained for hours. As the work

proceeded, moreover, the task became more and more laborious,

for the animal, no longer able to step over the wall, had to climb

NESTS OF PSEULOSCORPIONES. 105

up the inside and down the outside each time it went out, and

up the outside and down the inside each time it returned; and

further, there was increasing difficulty in fixing the fragments

against gravitation as the structure converged towards the top of

the roof. The animal was watched for five and a half hours,

during which it worked untiringly and with care, with brisk and

eager movements and without once resting. Its activity was

remarkable. When I left it at 11.30 p.m. only a small aperture

at the top of the roof remained to be filled in, and thus the

framework of the nest was almost complete. It must be explained

that the silk dried rapidly, the structure being firm from the first,

so that it was little if at all damaged by the continued scrambling

in and out of the animal. At the same time a certain elasticity

was evident, and thus during the final stages of this part of the

work—not seen in the present case—the animal would not be

greatly obstructed in forcing its way in and out of the small

decreasing aperture. Next day, at 7.30 a.m., the framework was

complete, the aperture having been filled in, and the builder was

thus entirely enclosed. The creature was still busy, however,

within the nest. One could see that the palp-fingers were being

applied from place to place, and the structure was temporarily

raised or pushed out first in one part and then in another. The

animal was evidently spinning, and at the same time re-adjusting

the materials to some extent; and it even detached and threw out

certain fragments, dried remains of prey ete., which had originally

been built in.

The structure thus made consisted of a vast number of irregular,

mostly small, fragments of wood and cork fastened together by

silk attached to their inner surfaces and extending from one to

the other; and the arrangement was such that the interior had

a regular approximately even surface. The silken attachments—

seen by looking down between the fragments—had the character

of an open irregularly joined-up meshwork, of which the threads

were of varying thickness, most of them relatively stout. This

meshwork ought perhaps to be regarded as the essential frame of

the structure ; the use of fragments is probably necessary in nests

entirely unattached above but otherwise it is not indispensable ;

for the animal is able, at least in certain positions, to make nests

ef spinning-work alone. The task of bringing the nest to the

condition now indicated had entailed much work; but what

remained to be done was more laborious and occupied a longer

time. Thisadditional work consisted of long-continued spinning,

the meshwork, the inner surfaces of the fragments, and in fact

the whole interior, including the floor, having to be covered with

silk, until at last every part was lined with the almost paper-like

tissue already described.

The making of nests such as that above considered, though

showing well the manipulation of the extraneous materials,

offered but moderate facilities for observing the spinning. This

was better seen in nests made in the spaces between the strips

106 MR. H. W. KEW ON THE

of wood and the glass. In such a nest the wood formed the

foundation for the floor and the glass that for the roof, the

built part consisting of a circular wall extending from one to

the other. Here one could see something, not only of the

making of the framework of the wall, but also of the subsequent

spinning, which iatter was obscured in nests of the former

character. The general proceedings of the animal were easily

observed; but it was less easy to make sure of the precise

character of the spinning, the task of following moving organs

under the magnification required being, even in the most favour-

able conditions, somewhat difficult. However, it was seen that

the silk proceeded from the distal part of the galee—there was

no doubt that it issued from the tips of the little branches there

situated —whence it was drawn as separate, highly viscid, very fine

threads. These threads, several from each galea, either remained

separate or coalesced into stouter ones, al] those from each galea

sometimes going to form a single thread. The spinning involved

small continuous forward and backward movements of the body,

such movements being derived from the joints of the legs without

necessary replacement of thefeet. The small forward movements

brought the galez into contact with the objects to which silk was

being applied, or with the tissue which was being augmented, and

from attachments thus made threads were drawn out during the

small backward movements, at which time also there were lateral

movements of the chelicerz, the galea-bearing fingers being swung

widely open. These small movements of the body, moreover,

brought the chelicere, not always to the same spot, but sometimes

a little above or a little below, or a little to the right or to the

left; and the cheliceree themselves, it may be explained, are

much more mobile than might be supposed, being capable of con-

siderable extension both forwards and laterally. While making

the framework the animal was seen to bring the cheliceree into

contact with the glass and by small rapid movements to brush on

minute confused attachments of separate threads. During this

part of the work the creature moved about freely, and the threads

thus attached, usually coalescing into one from each galea, were

earried from place to place, from roof to floor, or from either of

these to the inner surfaces of collected fragments, or from one

fragment to another, The threads coalesced at various distances

from the gale and not all of them at the same point, and since

they fused at once either before or after coalescing with other

threads or with whatever object they came in contact, the

irregular meshwork soon resulted. As the animal continued

to work the meshes became closer and closer, till little by little

they were filled in. At the same time the animal laboured to

cover the floor and roof with silk. At frequent intervals, often

throwing back the palps along by its sides, it settled down

to long-continued spinning; and at such times it maintained

for hours the continuous movements of the body and of the

cheliceree already mentioned. By this means silk was rapidly

NESTS OF PSEUDCSCORPIONES. 107

brushed on to the interior, first in one place and then in another.

It fused at once to the substratum, the exceedingly fine threads

now usually fusing separately without coalescing one with

the other. ‘This was well seen on the roof, where the threads

fused to the glass in more or less parallel order in series of

several running side by side. An appearance similar to that

here presented would be obtained perhaps if it were possible

to take the lines from a sheet of music, obliterate one here and

there, and turn the rest backwards and forwards in iregular

fashion one over the other. The number of threads in a series

varied, and in the present species not more than five were

counted—that being the usual number ; so that it appeared that

one or more of the branches of the galea had usually been inactive.

With the two gale the animal could thus produce at least ten

threads at a time; and it will be understood that the original

parallel order was soon lost in the general confusion of a closed

layer. Continuing thus to work at intervals for many days or

even weeks, the animal at length produced the final dense tissue

over every part of the interior of the nest. In the case of a

perhaps unusually dense moulting-nest, this part of the work

occupied the indefatigable energy of the animal during six weeks,

the completed structure thus costing incredible labour.

Chelifer latreillii Leach—belonging to the subgenus Chelifer

s. S., and provided with eyes—was the subject of similar though

less numerous observations. The galea is here shorter-shafted ;

and its distal half is provided with small branches, which in the

female—the organ is degenerate in the adult male—are six in

number. Specimens were obtained from tussocks of Ammophila

arenaria on sand-dunes; and the cells in which they were kept

were provided with leaves of that grass and with sand. Brood-

nests only were made—there were no immature individuals ‘to

make moulting-nests—and little need be said of them since the

animal’s methods were identical in al] essential matters with

those of C. cyrneus. The extraneous materials employed were

grains of sand, which the animal carried in the chelicere. Only

a little sand was used, however, the spun-tissue being slight

in texture and for the most part uncovered externally, The

silk was seen to proceed from the gale, as before, and that

deposited on the glass ran similarly in irregular more or less

parallel series; in these, in the present case, six threads some-

times occurred, presumably one from each of the six branches of

the galea.

Obisium muscorum Leach, the remaining animal observed, is

of vastly different type. It is longer-legged and less solid; and in

lace of the two dull eyes of Chelifer s. s. there are four shining

with a white lustre. The chelicerz are large and without galee.

The hard laterally compressed tubercle which replaces the galea

has, especially in the female, a bold convex margin, on or near

which, as already stated, Hansen more or less satisfactorily

detected six minute openings. The animal is common and

108 MR. H. W. KEW ON THE

easily obtained; but since it requires constantly moist con-

ditions there are difficulties in keeping it for long periods.

However, a few were maintained in health for several months;

and two females made brood-nests both attached in part to the

glass. It had been supposed that the differences in the chelicere,

more particularly the absence of galez, would correspond with

differences of method; but the animal’s proceedings appeared to

be identical with those of Chelifer. As with those animals, the

framework of the nest was rapidly constructed, the extraneous

matters being carried in the chelicere ; and no differences were

observed in the manner of spinning or in the tissue. The silk

was seen to proceed from the tubercle in several very fine threads,

which, more especially during the earlier part of the work, were

apt to coalesce, those from each tubercle often forming a single

thread. At other times, that is during the formation of the

dense tissue, the threads more often remained separate, fusing

thus to the substratum, and those deposited on the glass ran in

irregularly parallel series, usually six together, presumably one

from each of the pores of the tubercle.

Tt will have been noticed that the number of spinning-openings

appears to have been six on each chelicera, twelve in all, in all

three species, the presence or absence of the galea counting for

nothing in this respect; but this number is not necessarily

universal. The branching of the galea, for instance, is a variable

feature from group to group. For the rest, however, it may be

predicted that the general lines now indicated are those on which

all the animals of this Order, whether possessing the galea or not,

set about the construction of their nests.

NARUE

Summary.—Pseudoscorpiones make nests in part or wholly of

silk from their own bodies. They enclose themselves in them for

moulting, for brood-purposes, and in some cases for hibernation.

Such nests are closed cells of spun-tissue with or without an

external covering of extraneous matters. They are roughly

circular, but their form differs with the build and habitat of the

animal. They may be attached above and below to the solid

surfaces of narrow crevices and thus flattened, or they may be

attached below only, in which case they have a free convex roof,

or, again, they may be fixed here and there to surrounding objects

and roughly globular. The external covering, when it exists,

consists of earthy or vegetable fragments, which are not bound

on to the structure but firmly attached to it. The interior is

always free from foreign matters and smooth. The spun-tissue

is thin and dense, almost like silk-paper. It is composed of

innumerable threads crossed and re-crossed and coalesced in

irregular confusion and without interspaces. The material is

derived from glands in the cephalothorax, whose ducts traverse

the chelicer to near the apex of the movable finger, and open at

NESTS OF PSEUDOSCORPIONES. 109

the tips of the branches of the galea or on or near the margin of

a tubercle which replaces that structure in some groups. The

spinning is thus done with the chelicere, the galea or tubercle

being the structure immediately concerned ; and the presence

or absence of the galea does not appear to be associated with

differences of method or in the tissue. The combs ete. of the

cheliceree have nothing to do with the silk. All the nest-making

is done from within, the animal gradually imprisoning itself. The

construction of an external framework is the first part of the task ;

and when this has a coating of extraneous matters the animal

frequently goes out to collect materials. These it picks up in the

palp-fingers, transfers them to the chelicere, and returns thus

laden to the nest, where it attaches the materials together and to

those already placed by applying silk to their inner surfaces and

stretching threads from one to another. The silken attachments

form an open irregular meshwork, which is the essential frame

of the nest, and is constructed in some cases without the use of

extraneous matters. The silk is drawn from the galea or

tubercle in several separate viscid very fine threads, which remain

separate or coalesce, all those from each galea or tubercle some-

times forming a single thread. The spinning is associated with

continuous forward and backward movements of the body and

with lateral movements of the chelicerze. During the earlier

parts of the work, when attachments are being made from place

to place, the threads usually coalesce, and since they fuse at

once, either before or after coalescing with other threads or

with whatever object they come in contact, the irregular mesh-

work soon results. Afterwards the animal settles down to long-

continued spinning, and silk is rapidly brushed on to the interior,

first in one place and then in another. The threads now usually

fuse separately, being applied in more or less parallel series of

several side by side; and when both gale or tubercles are used

together ten or twelve threads may be deposited at atime. The

animal continues thus to work at intervals for days or even

weeks, till the final dense tissue is at last produced over every

part of the interior of the nest. The methods of three

species, representing both main divisions of the Order, were

observed in detail; they were essentially identical and probably

characteristic of all Pseudoscorpiones.

IX. List of References.

1. Friscu, J. L.— Beschreibung von allerley Insecten in

Teutsch-Land, viii. Berlin, 1730.

2. Roser, A. J.—Der monatlich-herausgegebenen Insecten-

Belustigung dritter Theil, Nurnberg, 1755.

3. Hermann, J. F.—Meémoire Aptérologique. Strasbourg, 1804.

4. De Turis, C.—Letitre addressée & M. Audouin sur quelques

Arachnides des genres Hydrachna et Chelifer. Annales

des Sciences naturelles, xxvil. pp. 57-78. Paris, 1832.

110

11.

12.

13.

14.

15.

16.

19.

20.

21.

22.

MR. H. W; KEW ON THE

Lucas, H.—Exploration scientifique de |’ Algérie. Histoire

Naturelle des Animaux Articulés, 1. Paris, 1849.

Menaz, A. —Ueber die Scheerenspinnen, Chernetide. Neu-

este Schriften d. Naturforschenden Gesellschaft, v. pp. 1—

43. Danzig, 1855.

Low, F.—Beobachtungen tiber das Hierlegen und Spinnen

der After- oder Biicherskorpione. Verhandlungen d. K.-K.

zoologisch-botanischen Gesellschaft, xxi. pp. 841-3. Wien,

1871.

. Srecker, A.— Ueber neue indische Chernetiden. Sitzungs-

berichte d. K. Akademie d. Wissenschaften, Math.-nat.

Cl., lxxii. pp. 512-26. Wien, 1875.

. Stmon, E.—Les Arachnides de France, vii. Paris, 1879.

. Becker, L.—Communications arachnologiques. Ann. Société

entomologique de Belgique, xxi. pp. exxxix-cxlu. Brux-

elles, 1880.

TuoreLL, T.—Deserizione di aleuni Aracnidi inferiovi dell’

Arcipelago Malese. Ann. Museo civico di Storia Naturale,

Xvili. pp. 21-69. Genova, 1883.

Croneserc, A.—Vorlaufige Mittheilung tiber den Bau der

Pseudoscorpione. Zoologischer Auzeiger, x. pp. 147-19].

Leipzig, 1887.

BerrKau, P.—Ueber den Bau der Chernetiden. Verhand-

lungen des naturhistorischen Vereines der preussischen

Rheinlande u. s. w., xliv. Sitz. pp. 112-117. Bonn, 1887.

CroneBerG, A.—Beitrag zur Kenntniss des Baues der

Pseudoscorpione. Bull. Soc. Imp. des Naturalistes (n. s.) 11.

pp. 416-461. Moscou, 1888.

BauzAn, L.—Revisione dei Pseudoscorpioni del bacino dei

fiumi Parana e Paraguay. Ann. Museo civico di Storia

Naturale, (2) ix. pp. 401-454. Genova, 1890.

THoreELL, T.—Aracnidi di Pinang raccolti ne] 1889 dai Sigri.

L. Loria e L. Fea. bid. x. pp. 269-383. Genova, 1890.

CampBripaGs, O. P.—-On the British Species of False-Scorpions.

Proc. Dorset Nat. Hist. ete. Field Club, xiii. pp. 199-231.

Dorchester, 1892.

Bernarp, H. M.— Additional Notes on the Origin of the

Trachez from Setiparous Glands. Ann. Mag. Nat. Hist.

(6) xi. pp. 24-28. London, 1893.

Bernarp, H. M.—Notes on the Chernetide. Journ. Linn.

Soc., Zool. xxiv. pp. 410-430. London, 1894.

Hansen, H. J.—Organs and Characters in different Orders

of Arachnids. Entomologiske Meddelelser, iv. pp. 137-251.

Copenhagen, 1894.

Bouvier, HE. L.—Sur Ja ponte et le développement d’un

Pseudoscorpionide, le Garypus saxicola Waterhouse. Bull.

Soc. entomologique de France, lxv. pp. 304-807, 342-343.

Paris, 1896.

Barros, J.—Mémoire sur le développement des Chelifer.

Revue Suisse de Zoologie, i. pp. 461-498. Geneve, 1896.

23.

24.

25.

26.

27.

28.

29.

30.

31.

32.

33.

34.

35.

36.

NESTS OF PSEUDOSCORPIONES. Wali

Supino, F.—Osservazione sopra VYanatomia degli Pseudo-

scorpioni. Atti della Reale Accademia dei Lincei, (5) vill.

pp. 604-8. Rome, 1899.

Frrronnizre, G.—Contribution a VEtude de la Faune de la

Loire-Inférieure. Bull. Soc. des Sciences naturelles de

YOuest de la France, ix. pp. 187-146. Nantes, 1899.

Wiru, C. J—On Chelonethi, chiefly from the Australian

Region, in the Collection of the British Museum. Ann.

Mag. Nat. Hist. (7) xv. pp. 94-148. London, 1905.

Kew, H. W.—Chernes cyrnews m Nottinghamshire: a recent

addition to the known False-Scorpions of Britain, Trans.

Nottingham Naturalists’ Society, 1905-6, pp. 41-46:

Nottingham, 1906.

Wirn, C. J.—The Danish Expedition to Siam, 1899-1900.

Chelonethi: an Account of the Indian False-Scorpions,

together with Studies on the Anatomy and Classification of

the Order. Kgl: Danske Videnskabernes Selskabs Skrifter,

(7) iii. pp. 1-214. Copenhagen, 1906.

Jackson, A. R.—Rare Arachnids captured during 1906.

Proc. Chester Society of Nat. Science, etc., “ils JOD Laas

Chester, 1907.

Goprrey, R.—The False-Scorpions of Scotland. Annals of

Scottish Natural History, Xvi. pp. 90-100, 155-161 ;

xviii. pp. 22-26: xix. pp. 23-33. Edinburgh, 1908-10.

Surprny, A. E.—In Sepewicn, A. : A Student’s Text-Book

of Zoology, iii. London, 1909.

Warpurton, C.—In Harmer & Suipcey: The Cambridge

Natural History, iv. London, 1909.

Kew, H. W.—A Holiday in South-Western Ireland: Notes

on some False-Scorpions and other animals observed 1n

the Counties of Kerry and Cork. Irish Naturalist, pxaibes

pp. 64-73. Dublin, 1910.

Krw, H. W.—A Synopsis of the False-Scorpions of Britain

and Ireland. Proc. Roy. Irish Academy, xxix. B, pp. 38-

64. Dublin, 1911.

Kuw, H. W.—Pseudoscorpiones. In Clare Island Survey.

Ibid. xxxi. (38) pp. 1-2. Dublin, 1911.

Ewinc, H. E.—Notes on Pseudoscorpions. Journ. New

York Entomological Society, xix. pp. 65-81. New York,

OMI

Goprrey, R.—In Exixesen, E.: The Pseudoscorpions of

South Africa, based on the Collections of the South African

Museum, Cape Town. Ann. South African Museum, x.

pp. 75-128. London, 1912.

Wh) ae

i aust teal aie

x tay reas:

ya hteek PEE

VoA gi hye ih nitledniga ns

Rely ae Hae ayy Peer

Eat

ben

Pes. 1914 Meggatiaeies

West,Newman lith.

SHESUGUMUNEIE Ol! WC RMMENaOun Zn NIV ITE CCIILIS -

P.Z.8.1914, Meggitt. Pl.Il.

Newman Itth.

West

SRC WUE Ol WCEMMECOUZE INIA PIULUCO LLL

3 P.Z.S. 1914, Meggitt. Pll.

West,Newman ith.

STRUCTURE OF ICHTHYOU47 NIA FRLICOMEs.

P.Z.S. 1914, Meggitt. Pl. IV.

West,Newman Jith.

STRUCTURE OF ICHTHYOT4INIA FILICODUIS.

ON A TAPEWORM PARASITIC IN THE STICKLEBACK. TLS:

8. The Structure and Life-History of a Tapeworm

(Ichthyotenia filicollis Rud.) Parasitic in the Stickle-

back. By F. J. Meeerrr, M.Sc. (Birm.), Board of

Agriculture and Fisheries Research Scholar, University

of Birmingham *.

[Received November 17, 1913: Read March 3, 1914. ]

(Plates I.-IV.t & Text-figures 1-5.)

InDEx.

Page

IN PRODUCTIONS fs jc. c4 sy ae aaron GS

ANATOMY.

ELIS tO Cale es met err ace sae sernseee meer ites ema TTA

Occurrence menses jst een, ee LL

Externally © harackers eee eee: eee enenes milellcy

Histology Wn edi eee oe A Rea UTS

Miusculatune meee eran papa eee ares S19

Exeretory System ............0.0.0.........0... 119

Nervous System ..............0.0..:000cc:2000-. 121

Maley Orcas yee teachin eee Mik aen OT

HemaletO1cansiy ke on eee Le)

Lirs-History.

ELIS HOT COU FF eis Fears. teeta hse peau TOLL

Infection Experiments ........ 00.00.0000... 125

Developinentiee se mcs cc ee ees LO

SUMMUMAUR Wap a5. 0, bra. cece se fear eer is cance 135

AGT AUN RE ion Pete saat ts clemiees cere nts TDS

INTRODUCTION.

Ichthyotenia filicollis Rud. (= Tenia filicollis Rud.) is a species

the life-history of which, like that of most Ichthyotenie, is prac-

tically unknown. No successful infection experiments have been

carried out upon any member of the genus, and it was therefore

thought that a connected account of the whole life-history would

be of service in establishing a complete diagnosis of the family.

In carrying out this work it was found that the only two complete

descriptions of this species, those of Kraemer (10) and Benedict

(3), are said by La Rue (12) to be descriptions of other species :

if this be correct, then there is at present no paper giving a

detailed and accurate study of this species. The paper is there-

fore divided into two parts, the first being a detailed account

of tne anatomy of the specimens obtained, compared with the

descriptions of Kraemer and Benedict, and the second an account

of the life-history as far as it could be determined.

J wish here to express my indebtedness to Professor F. W.

* Communicated by Prof. F. W. Gawsrn, F.RS., F.Z.S.

+ For explanution of the Plates see p. 137.

Proc. Zoou. Soc.—id14, No. VILL. 8

114 MR. F. J. MEGGITT ON A

Gamble, F.R.S., for the very valuable help he has given me in

the course of this work by his criticism and advice. My thanks

are also due to Professor G. S. Brady for identifying specimens

of Cyclops varius Lilljeborg submitted to him.

ANATOMY.

Historical.

This species was first described by Rudolphi (21). He found

two species of Cestodes, one in the intestine of Perca fluviatilis,

and one in that of Gasterosteus aculeatus, and called them Tenia

ocellata and T' filicollis respectively. In his original descriptions

there is very little to distinguish between the two species.

Early investigators, Bellingham (1), Dujardin (8), Diesing (7),

and Cobbold (6), confined themselves to external characters,

Zschokke (26) being the first to describe the internal anatomy.

Later, Kraemer (10) made an exhaustive study of the two species,

and concluded that there was no essential difference between them:

in the same paper he gave a number of characteristics peculiar to

fish Teenie, and suggested that they might form a special group.

Still later, Lénnberg (15) separated them under the generic name

of Ichthyotenia. Riggenbach (20), for the first time, summarised

the investigations on the genus and added several new species to

the list. In 1899, Railliet (19) showed that Tenia ambigua Duj. is

synonymous with 7’. ocellata Rud. and 7’ filicollis Rud., and that

therefore the genus Proteocephalus Wein. (25) is synonymous

with Jchthyotenia: the first bemg the older name, should be

adopted according to him. Benedict (3), who confirmed Kraemer’s

description and elucidated a few fresh points, adopted this name.

The latest paper upon the subject is that by La Rue (12). He

asserts that Tenia filicollis Rud. and 7. ocellata Rud. are two

separate species. The species which Kraemer described under these

names, La Rue asserts was not 7’. filicollis nor 7. ocellata but an

entirely new species (P. fallax La Rue). He further says that

Benedict did not describe either of Rudolphi’s species, the species

actually described being P. exigwus La Rue. Zschokke (26) also,

according to him, described P. dubius La Rue, not 7. filicollis

Rud.

Tetracotylus (Monticelli, 18) he does not consider to be

synonymous with Jchthyotenia. He adopts the generic name

Proteocephalus on account of its priority : “I cannot regard the

objections of Lithe (16) as adequate for its rejection.” This

paper is merely a preliminary note to a monograph he is about

to publish.

From the brief historical account just given, it is obvious

that the nomenclature of the group is in a state of great con-

fusion. This is chiefly due to the vague descriptions of the

early investigators, most of them being based upon characters,

such as the scolex, which ave far too variable to be utilised for

TAPEWORM PARASITIC IN THE STICKLEBACK. 115

classification. Riggenbach (20) lists 29 species, of which only 7

are satisfactorily described. At the present time, it is impossible

to say with certainty whether La Rue is right in retaining the

species Tenia filicollis Rud. and 7’. ocellata Rud. The only

correct part of Rudolphi’s description is that referring to the

scolex, and since the scolex is very variable, Rudolphi’s species

cannot be determined with any degree of certainty. The question

can only remain in abeyance until the publication of La Rue’s

monograph.

Tt cannot be said, however, that La Rue is justified in retaining

the generic name Proteocephalus. As Lithe (16) has shown, this

name was used by Blainville (4) for a family of Cestodes con-

taining Caryophylleus, and for that reason should be abandoned

in favour of tue next oldest name, /chthyotenia. In the same

paper in which La Rue rejects the name Jchthyotenia, he pro-

poses the name Monticellia for the genus Tetracotylus Monti.,

Braun (5) and Lithe (16) having shown that the latter name was

proposed by Filippi to designate a group of immature Trematodes.

He thus admits the principle upon which Lihe’s objections are

based.

Occurrence.

The specimens studied were collected from the intestines of a

number of sticklebacks (Gasterostews aculeatws Artedi) taken

from the Edgbaston Reservoir. The numbers in a single host

varied from one to twenty-five. Almost every fish in autumn was

infected with one or more of these parasites, 75 per cent. of which

were adult’: in winter, the number of infected fish was considerably

smaller, and adults were rare; while in spring, the proportion

of adults again increased. Adult specimens were found, how-

ever, all through the months September to June, but while

their proportion to young forms was 75 per cent. in the first

month, it was only 15 per cent. in March. Von Linstow failed

to find it adult at all in winter, and Zschokke only noticed it

three times.

External Characters.

The length of adult specimens varies from 24 to 33 mm., the

breadth from 1:0 to 1:2 mm. The head is almost continuous

with the neck and is only slightly globular except when violently

contracted (text-fig. 1): in Kraemer’s specimens it was spherical

and sharply marked off from the neck. Jt is furnished with

four suckers, two dorsal and two ventral, and at its apex isa fifth,

which is not very well developed although apparently functional.

The shape and size of the head and suckers are subject to

considerable variation. The whole region of the head anterior to

the suckers can be retracted (Pl. I. fig. 5): or the retracted area

may even be so large as to include the suckers themselves

(Pl. I. figs. 4, 6, 7)—in the latter case, they are hidden in a

deep vertical fissure, and can only be seen by staining, while the

116 MR. F. J. MEGGITT ON A

scolex appears to have a terminal depression similar to that

of Schistocephalus dimorphus Crepl. On the other hand, the

suckers may stand out as small cups at the four corners of the

base of a tetragonal pyramid, the apex being formed by the

fifth sucker (Pl. I. fig. 2): the scolex may also be inserted in the

neck like a cork ina bottle (Pl. I. fig. 1). It is obvious, there-

fore, that the shape of the scolex is a character to which little

importance can be attached. In life the head undergoes a

regular series of changes, passing from an extremely elongated

phase to a violently contracted and spherical one (text-fig. 1).

Text-figure 1.

Movements of scolex of Ichthyotenia filicollis. XX 142.

The diameter of the head varies from ‘166 to -2 mm., and of

the suckers from ‘04 to ‘07 mm. The neck is relatively long, and

occupies about a quarter of the whole length. The first pro-

glottis separated from it is much broader than long, its

dimensions being 192-397 mm. broad x ‘1-166 mm. long.

The total number of segments varies from 24 to 33, Kraemer

states that the first proglottis is much longer than broad ; while

Benedict figures two varieties, in one of which it is longer than

broad, and in the other is broader than long. Liihe (17) gives

it as longer than broad. The following table gives the external

measurements according to various observers (Table A). Poste-

riorly, the segments become longer in proportion to their width,

until the penultimate proglottis is 1j-2 times as long as wide.

The posterior proglottis is rounded at the tip, and at its apex is the

excretory pore. In my specimens this was often very prominent,

opening on a small triangular papilla.

The separation of the proglottides from one another is very

indistinct, being indicated in the posterior portion of the stro-

bilus only by a slight notch.

The genital openings are lateral, and alternate irregularly.

According to Benedict, there is a small genital sinus, while

BSL)

TAPEWORM PARASITIC IN THE STICKLEBACK.

"SOIJOUIT|[IU UT S}UOTMOINSvOUT [VY

L00- * 4

€00. goo. | 8z0.-co. | 990. z00- 6100. G00- Os ff {sco

ae jee E = ———— =| } eae Ps

CGO. % 100. 00. 100. S00>, seme oman qolpauag

BOO: 160. 2 DO a a ie Se 00. £00. g00.-g00. J IOWIVAST

“Tpoapnu ‘epuu | “LIAVT eae “1a AB] “OAV ; ‘apotano Jo ee

Io} Aayy AvpNoTyNd-qns eccanlar -oTOSNUL -aPOSn ut Ae TLE] AST snosieofva

JO lajauvrg | Jo lajoulvig Jo s]Ta9 : [eulpnyisuoy| arpnoarg, aoe jo ssauypoiyy, | jo aojowvig

Se ie eee ee ew Su llth me ee Se ss See ee a See, Dal es ale ae

‘dq a@1avy,

t VG OU aes Heston) ele rash Pe cie ee c eesae” Sa|feet owe ons aa

T S apim L62.-Z61.- 0--f0.- 6-991. 6-1-0.1 €&-P6G JL a00Tq

Ge OGI-0S VII. 3S OOT-0€ a ee OU ET

= bee = = |e

i | OF ‘Soy 9T.1X ge oe Pe

S$_g | aplM OG.-GL- ss

¥ 8 96-ST ao pug

= GI--OT.- FO. GI: 8.

z 91-S1 AqoLAwA 4ST

| : golpouag

a | “Buoy pe.X

ia OG1—-08 Sot Gees 91-0 80. VII S 09 ees TQWIBLY

“you 70 ‘sapiyqopoord *s19qopso.d *yoatt Jo “S1aYONS FO *pvay jo “Apoq Fo *Apoq Jo

yysuaery | Foxequinyy | Asay Yj prog TAZOUIVIC] Ioqyoweiqg, Yypvatey yysuary

“VY HTEaV

118 MR. F. J. MEGGITT ON A

Kraemer states that vagina and cirrus open directly to the

exterior. My own specimens confirm the former statement.

FTistology.

The body is covered by a two-layered cuticle (Pl. II. fig. 12).

The outermost layer is dark-staining and rough in outline, and

is split up by vertical clefts into a great number of fine hair-like

processes. The whole appearance is that of a cast cuticle. A

similar cuticle is present in Schistocephalus dimorphus Crepl.,

and this Kiessling (9) considers to be a disintegration product of

the underlying layer: it is probable that this is true of [ehthyo-

tenia jilicollis also. Under this cuticle is a feebler-staining

homogeneous layer. Acco:ding to Kraemer, there is a third layer

(‘‘ Cutis”) underneath this: im one or two cases in my specimens

it has appeared as though there were a third layer, but this is pro-

bably only an optical effect. Benedict does not mention it at all.

Following the cuticle is a layer of circular muscle-fibres, and

then one of longitudinal muscle-fibres. Internally is a sub-

cuticular cell-layer, composed of spindle-shaped cells, *02—028

x ‘007 mm., drawn out into delicate protoplasmic processes

internally, and possessing a prominent nucleus, ‘005 mm. dia.,

and nucleolvs, ‘003 mm. dia.

The structures above-mentioned appear to vary greatly in size

according to different investigators (Table B); it is apparent,

therefore, that their size is a character too unstable for classi-

fication.

Under the sub-cuticula is the parenchyma. This consists of

polygonal cells, with a varying diameter of -0046—008 mm.,

with nuclei ‘0023 mm. dia. ‘The cell corners interlace with

one another to form a loose meshwork enclosing intercellular

spaces, the meshwork being closer externally and less compact

between the various organs. The intercellular spaces are often

empty, but more usually are filled with a feebler staining granular

mass which Kraemer thinks is an BEueneny product of the

parenchymatous cells.

Forming a regular layer under the cuticle in the more

posterior part of the body are large numbers of rather small

calcareous bodies. They are oval and stratified, rather like

starch-grains. In the larval cestode they are much longer and

not so stratified, and are scattered irregularly over the body ;

they show very distinctly under the cuticle, and appear as if

actually on the body surface.

There do not appear to be any oil globules comparable with

those mentioned by La Rue (11) in P. filaroides. If the Cestode

be examined in water, numerous transparent bubbles may be

seen attached to the body and gradually growing larger ; these

may correspond to the oil globules and be due to the water

round the specimens causing the minute invisible oil droplets on

the body to coalesce into larger drops.

TAPEWORM PARASITIC IN THE STICKLEBACK. 119

Musculature.

The musculature of the body consists of circular and longi-

tudinal muscles (already described) under the cuticle, and more

powerful inner longitudinal muscles (PI. IT. fig. 12). These latter

are arranged in bundles, which run in the outer layer of the paren-

chyma and havea diameter varying from ‘008 to ‘0103 mm. (Bene-

dict, 0015 to ‘(004 mm.) ; anteriorly they diminish in size, although

in the scolex anteriorly to the suckers they occasionally have a

diameter of -0092 mm. The single fibres of which they are

composed are °0035 mm. dia. (Kraemer, :007) ; occasionally

spindle-shaped nuclei, ‘0028 x -0092 mm., with a prominent

nucleolus, ‘(0023 mm. dia., are to be seen, the long axis of the

spindle being parallel to that of the fibre. In the body, and

especially in the neck, large fibres pass dorso-ventrally between

the longitudinal muscle bundles. According to Benedict,

“a loose sheet of circular muscle-fibres weaves around the

longitudinal bundles. Large fibres pass in a transverse direction

between these muscle sheets. The divisions between the pro-

glottides are formed by the interlacing of these fibres with

similar ones which cross them at right angles, both sets being

here much more complicated than in other regions.” He does

not figure them, however, and they do not appear to be present

in my specimens.

The musculature of the scolex consists of prolongations of the

inner longitudinal muscles together with scattered dorso-ventral

and sagittal fibres. The suckers have equatorial, meridional, and

inner radial muscle-layers ; they are covered by a continuation

of the body cuticle, which is not absent, as Kraemer states, from

their cavities.

Kaucretory System.

The excretory system consists of four main longitudinal vessels,

two dorsal and two ventral, which run, internally to the longi-

tudinal muscles but externally to the yolk-glands, from the scolex

to open at the posterior end of the body. ‘They are not, as

Kraemer states, of equal diameter, but are unequal, the

two dorsal being ‘0045 x -0068 mm., and the two ventral

"002 x 015 mm. In the scolex they end in a circular com-

missure, ‘0023 mm. dia., just under the suckers (text-fig. 3).

Outside their course in the neck is a system of fine canals

forming a complicated plexus from which other fine canals lead,

either to the exterior or to the four longitudinal canals. These

“foramina secundaria” have, according to Kraemer, a swollen

portion at their opening, while the opening itself is guarded by

a wisp of hairs. I have never had the good fortune to see the

dilatation, the canals apparently having a uniform diameter,

‘0017 mm., throughout their entire course; the wisp of hairs

also I have never seen, although the outer cortex at places

presents rather a hair-like appearance. La Rue (11) states:

“There exists a wide variation in the types of structures which

120 MR. F. J. MEGGITT ON A

Text-figure 2.

Main excretory vessels of a proglottis: reconstructed from camera-drawings of

sections. J.c., /.c.’, ventral and dorsal longitudinal excretory vessels; /.p.,

lateral excretory pore.

Text-fig. 3. :

LAS.

Transverse section through the neck, showing the excretory plexus. J.c., 7.c.', ventral

and dorsal longitudinal excretory vessels; /.., lateral excretory pore ;

r.c., circular commissure. X 435.

TAPEWORM PARASITIC IN THE STICKLEBACK. LOT

are grouped under this name. The ‘foramina secundaria’

described by Kraemer for P. ocellata and P. torulosa, and by .

Riggenbach for P. fossata, P. abscissa and Corallobothriwm

lobosum, are muscular pulsatile vesicles which open at the

posterior lateral margin of each proglottid. Benedict finds no

such vesicles for P. ambloplitis or for P. ocellata, nor do any of the

secondary excretory openings seen by me come under the type

described by Kraemer and Riggenbach.” Throughout their entire

course, the longitudinal canals give off many side branches which

anastomose with one another and with the main canal; in many

cases they end blindly in parenchymatous spaces. At irregular

intervals, other lateral branches are given off; these gradually

diminish in width and finally open to the exterior. Im the neck

this anastomosis is much more evident, at times the main canals

being lost in the plexus formed.

At the posterior limit of each proglottis, the four longitudinal

canals are joined by a circular commissure whose lumen is equal

to that of the canals.

The four canals open posteriorly in a notch at the apex of the

last proglottis. There does not appear to be any such “ Endblase i

as Kraemer figures, the canals meeting at one point, from which

a common canal leads to the exterior. Instead of the notch

there is often a triangular projection at the apex of which is the

excretory pore. In young forms there is a very pronounced

“ Endblase” (Pl. IV. fig. 40) into which the excretory canals

probably open, but this disappears in older specimens. .

Nervous System.

Owing to the difficulty of staining it, I have not been able to

make out the nervous system at all satisfactorily. Anteriovly

there is a nervous mass, the “brain,” lying between the four

suckers and just under the fifth; from it are given off two

longitudinal nerve-trunks which run down the lateral margins

of the strobilus externally to the excretory system. Benedict

describes in addition, four transverse trunks arising from the

anterior nerve-mass, each running straight out between the two

suckers; about halfway to the margin of the scolex each

branches into two secondary trunks, which run at right angles

to their previous course as far as the corresponding sucker.

Male Organs.

The testes are spherical bodies, ‘055 mm. dia., about 40 in

number, scattered throughout the entire space between the

ovaries and the anterior edge of the proglottis; there is no

central layer from which they are absent. ach is tightly

invested with an exceedingly delicate tunica propria, not, as

Benedict figures, in a loose membrane. In transverse sections

they appear hollow, with a cavity divided by septa into a number

of small compartments filled with granular matter. Minute

vasa efferentia lead from them to a common vas deferens,

122 MR. F. J. MEGGITT ON A

0184 mm. dia, In the centre of the proglottis, this is coiled into

a spherical ball, in the coils of which he the spermatozoa.

The walls appear structureless, but have occasional nuclei,

0046 x ‘0011 mm., scattered along them. Benedict states that

the coils are bound together by parenchymal] strands, but these

I have not been able to see. From this coiled portion, a short

duct leads to the cirrus-sac, but before entering it, decreases in

diameter to ‘008-009 mm.

The cirrus-sac itself (Pl. I. fig. 9) is oval, slightly constricted

in the middle, and stretches about 3—-}+ across the proglottis,

extending some distance beyond the vitellaria. Its small size

in my specimens is surprising, since both Kraemer and Benedict

figure it as reaching to the middle line of the proglottis, and its

comparative size is one of the characters used for specific

distinctions. Its walls (Pl. I. figs. 9 & 10) consist of outer longi-

tudinal muscles, ‘0023 mm. thick, and inner circular muscles,

0023 mm. thick, with an exceedingly delicate cuticle surrounding

them. The basal end of the cirrus-sac is turned rather obliquely

towards the dorsal side of the proglottis. Its walls bend back

to form a small tube which becomes united with the wall of the

vas deferens. This latter, just within the cirrus-sac, has a slight

muscular coat which gradually becomes thicker and passes into

the muscular coat of the cirrus. The vas deferens is coiled once

before opening into the cirrus.

The latter is an almost straight cylindrical tube, without the

enlarged distal portion figured by Benedict. It consists of an

outer layer of longitudinal muscles ‘003 mm. thick, an inner one

of circular muscles (0023-004 mm. thick, and a cuticle bearing

fine bristle-like projections externally. The lumen here is

0046 mm. dia. The external cuticle of the body tucks in

at the opening of the cirrus-sac and lines the inner wall of

the cirrus for some distance. Scattered along the course of the

cirrus-tube and opening into it are numerous pear-shaped glands,

0034 x °0009 mm.

The space between the inner tube and the wall of the cirrus-

sac is filled with fibrous tissue containing many nuclei, 0023 mm.

dia., but I have been unable to see the definite fibres figured

by Benedict. At the posterior end of the sac are muscles

extending from the cirrus-tube into the parenchyma, and

probably serving for retraction.

Female Organs.

The aperture of the vagina is 008 mm. in diameter and is just

anterior to that of the cirrus-sac. A circular sphincter muscle

(Pl. IT. fig. 13) encloses the vagina a little within the aperture ;

it is hemispherical in section, with a diameter of 008 mm. The

vagina itself runs back to the middle line of the proglottis, tums

at right angles to its former course, and finally opens into the

oviduct at the posterior end of the proglottis (text-fig. 4). Its

TAPEWORM PARASITIC IN THE STICKLEBACK. 123

walls consist of an inner ciliated epithelium—which appears to

pass into the cuticle of the body—a muscular layer, and an outer

epithelium ; altogether the wall is ‘0034 mm. thick, and the lumen

of the tube -003 mm. dia. Along its course, but particularly near

its opening to the exterior, are numerous glands, 004 mm. dia.

at their widest part x ‘007 mm. long, with distinct nuclei,

-0011 mm. dia., and having fine ducts opening into the lumen of

the vagina. Just before it opens into the oviduct, the walls have

a slightly different character. They diminish in width to

0029 mm., the muscular layer nearly disappears, and the main

thickness of the tube is due to a layer of cubical epithelium,

similar to that lining the cavity of the oviduct. The opening of

the vagina into the oviduct is very small.

The ovary is two-lobed. The lobes are elliptical, and are

joined anteriorly by a small common portion. Kach ovary is

surrounded by a delicate membrane, which, according to

Kraemer, passes into the sheath of the oviduct. The ovary is

usually so full of eggs that their shape is changed by compression

from spherical to polygonal. They will be deseribed later in

connection with the life-history.

Text figure 4.

o.d.

O.C.

Diagram of the female genital ducts. 0.c., odclapt; o.d., oviduct; o0.¢., odtype ;

0.v., ovary; v., vagina; 2.d., vitelline duet; v.d.’, common vitelline duct;

u., uterus.

Leading from the connecting-bridge of the ovaries is the

odclapt or “Schluckapparat ” (PI. Il. fig. 14). This is a nearly

spherical muscular organ, -002 mm. broad x ‘0022 mm. long,

with a lumen of 008 mm. dia. Its cavity is lined by cubical

epithelium according to Benedict, but with simple cuticle

according to Kraemer ; the latter is the case with my specimens.

The muscular band surrounding it is hemispherical in transvere

section and constitutes the greater part of its walls. An irregular

ring of nuclei surrounds it externally.

Into the odclapt opens the oviduct. This is a short tube

124 MR. F. J. MEGGITT ON A

going to the posterior margin of the proglottis, where it is joined

by the vagina, and then, bending at an angle of 30°, it runs

anteriorly to open into the obtype just under the odclapt. Its

wall consists of an outer epithelium, a muscular layer, and a

thick layer of ciliated cubical epithelium, the wall being

‘0034 mm. in thickness, surrounding a lumen ‘009 mm. dia.

At its opening into the ooty pe the lumen of the oviduct

decreases to ‘003 mm. dia.

Benedict states that gland-cells are scattered along the external

epithelium, but I have been unable to see them.

The odtype (Pl. I. fig. 11) is a rounded body, 014 mm. dia.,

with walls ‘0029 mm. thick, composed of an inner non-ciliated

epithelium, a circular machen layer, and an outer epithelium ;

round it are scattered nuclei, ‘0011 mm. dia., not arranged in

definite rows as Kraemer states. Surrounding the odtype and

opening into it by fine ducts are shell-glands of irregular shape

and with definite nuclei. Benedict represents the odtype as

an elongated tube between the opening of the common vitelline

duct and the uterus: in the position mentioned there is certainly

a duct which appears, as he says, like a specialised portion of

the oviduct, but this is not the odtype.

The vitellaria consist of small follicles scattered along the

whole length of the lateral margins of the proglottis. The

follicles are ‘0092 x 0069 mm. in transverse section, with a

distinct nucleus, (0034 mm. dia., and are grouped together in

fours or fives, each group being enclosed in a membranous sac

‘08 mm. dia. Two longitudinal ducts, one on each side, run

down the proglottis, receiving numerous fine ducts from the sacs,

to its hinder end, where they turn inward and join to form a

common duct. This duct runs longitudinally for a short distance

to open into the odtype. It has a lumen -0029 mm. dia., with

walls ‘0023 mm. thick, consisting of inner and outer epithelia

surrounding a well-developed circular muscular coat : connective

tissue nuclei form a definite layer round it.

The uterus passes as a narrow tube from the odtype under the

connecting portion of the ovaries, and expands as a large sheet

with 6-10 lateral outgrowths along the ventral surface of the

proglottis (Pl. I. fig. 8). In older segments, the uterus fills

the entire segment to the exclusion of all other organs. The

ova escape through a cleft in the mid-ventral line.

Lirr- History.

Historical.

The life-history of this species is unknown. Von Siebold (24)

states ‘‘le Zenia longicoltis et le ZL. ocellata qu'on rencontre

enkystés, hors du canal intestinal, dans la foie de divers

Salmones et Perciides:” the fish are swallowed ly pike or

perch, in the intestine of which the parasite becomes adut. In

TAPEWORM PARASITIC IN THE STICKLEBACK. 125

another work (28) he announced definitely that the larval forms

of 7’. longicollis and 7’. ocellata occur encysted, with non-mature

unsegmented bodies, in the livers of salmon and perch.

Zschokke (26) found the unsegmented larva of 7’. longicollis in

Salmo umbla.

Similarly, von Linstow (14) states. ‘‘ Die Larva findet sich

encystiert in der Leber desselben Fische, welche die erwachsene

Tanie in ihrem Darm beherbergen.”

Liihe (17) also inclines to this view. ‘ Als zugehérige Larve

ist eine in der Leber von Perciden encystiert gefundene Cestoden-

Larva betrachtet werden (¢).”

“ Leuckart (13) reports a plerocercus found by Gruber in

Cyclops serratulus which he believed to be the larva of Proteo-

cephalus torulosa Batsch. From a comparison of his figure,

which is drawn to scale, with measurements of P. torulosa, it

seems that there is some foundation for this view” (La Rue (12)).

To this larval host Lithe (17) adds C. strenwus.

It may be seen from the above summary that no accurate

experiments have been made with this genus. Most of the

investigators incline to the view that the larve occur in

the same host as the adult form, basing their opinion upon

comparisons of the size and general shape of both forms. The

connection 1s thus very loose. The infection would have to be

by means of eggs. As there is no swimming-mantle, these do

not float, but sink to the bottom; and since the sticklebacks in

Edgbaston Reservoir feed chiefly upon Entomostraca, the com-

parative rarity with which they would swallow eges could not

account for the heavy infection observed. Moreover, the oncho-

sphere would have to bore its way through the walls of the

alimentary canal to the liver, encyst there, and then return to

the alimentary canal again; for there could be no other way for

it to infect both the liver and the intestine of the same host.

This hypothesis 1s not very plausible and is not very well

supported. At different times I have examined the livers of

sticklebacks but without finding any trace of cysts, although

nearly all the stages have been found in the alimentary canal.

My own researches point to the conclusion that an inter-

mediate host, Cyclops varius Lillj., 1s necessary for the full

development of the egg, and it is through the final host

swallowing the intermediate host, that the parasite obtains

entrance to the alimentary canal of the former. This view is

supported by the discovery of the plerocercoid of P. torulosa,

previously mentioned*,

Infection Haperiments.

In studying the life-history of this parasite, I first began with

the eggs. If the Cestode be adult or nearly so, directly it is

removed from the intestine of its host, it begins to discharge the

egys In a continuous stream from a slit in the ventral body-wall.

* See Note ou pp. 1387-138.

126 MR. F. J. MEGGITT ON A

These were pipetted into a small watch-glass, water (pond or tap)

was added, and they were examined every day for about two

months. It was assumed that water would be the natural

medium for their development, since under ordinary circum-

stances they would be expelled into the pond and wait there

until swallowed by their first host. Altogether 20 cultures

were thus made and kept from the middle of October until the

end of January, but with no results. The most advanced stage

was that shown in Pl. ILI. fig. 23, but this was also observed in

eggs taken freshly from a living specimen. Cultures were also

made in salt solution, peptone, albumen, and fecal matter,

different strengths being used, but with no result.

While these cultures were going on, examples of the most

common invertebrates in the reservoir (various species of Cyclops,

Cypris, Nais, and Tubifex, together with some unidentified Ento-

mostraca and numbers of aquatic insect larvee) were collected. They

were placed in small dishes, each species being kept separately

and in definite numbers. A certain amount of Diatoms, Huglene,

Algee, etc., was placed in each dish for food, together with a large

quantity of eggs. No results were obtained until Jan. 7th, when,

after examining a Cyclops, the cover-glass accidently crushed it ;

on re-examining it, several tapeworm eggs were seen in the

crushed-out body-mass. All the cultures were then, and for

some days afterwards, minutely re-examined, but eggs were only

seen in the Cyclops.

A fresh series of cultures was made with Cyclops from three

localities to test this result, with a control to each culture. As

was expected, the controls only gave negative results. The other

cultures turned out better than was expected (see Table C): as

many as 17 out of 20 Cyclops in one case were infected, while

tliere were some infected Cyclops in every culture. The figures

do not represent the whole truth, however. It is extremely

difficult to keep the Cyclops alive under these artificial con-

ditions; in one culture 4 out of 15 died in one week, in

another 2 out of 6, in another 3 out of 7, ete.

Schneider (22) found that in his infection experiments, feeding

Gammarus locusta with eggs of a species of Proteocephalus, the

Gammarus died in large numbers owing to too heavy infection:

this may account for the death of some of the Cyclops, since as

many as six larvee in one specimen have often been seen. As

it is impossible to distinguish onchospheres in dead Cyclops,

those dead cannot be counted. (In parenthesis, it may be said

that the eggs appear to secrete a toxin which acts injuriously on

other animals in the small culture dishes, but which, in the large

volume of water in the reservoir, would be diluted until it would

become negligible. In one instance a culture was started in a

dish containing 20 Cypris, some Huglene for food, and some

of the Cestode eggs; a control of the same number of Cypris

was placed under identically the same conditions, except that

the eggs were absent. Im 18 days all the Cypris in the

first culture were dead; in the control I counted 220 living

TAPEWORM PARASITIC IN THE STICKLEBACK.

*pay.ivjs=g "QINJVUL JOU SBSI APOIO »

"JP poapoejul 957 UId.1Eg

“paqoayuy

poppe

e I

OL 4 v

wo ae

*MIVVALQO of

O00 000 6 ary |

L i S

A =

| ¢ S

= PR s

cs @ I 5 |

ae 3 os é 3

Cae eo giane es rd See P S

| |

Aa = | a 5 A

a | ee | a | ge | ee | oe | ex or | oe | o

Bo Se SS ee ee

“AIBN

‘IUIT} TOBI PazPOIJUL] 19d UN NT

°C) wavy,

“WOJLONT S SULT

“UOysNgspiy

“UOJION, S.SULST

s WOZAUIL Y.TAATO AA

“UOqsRqsSpy

“AQ ROOTT,

VEL

*sdopotp

JOMON

O0ZG

DAE

OXE

OXG

OA Gx

DAT

OXI

‘aan

128 MR. F. J. MEGGITT ON A

Cypris. In all cases the animals in the control lived longer

than those in the experimental dishes.) Moreover, no attempt

was made to discriminate between the different species of

Cyclops, since the species infected was not known. Under the

most favourable conditions it is extremely difficult to see the

eges in the host, so that, owing to the opacity of the body

and the development of the dense black ovaries, many probable

cases of infection have been overlooked. The results were there-

fore as good as might be expected.

The intermediate host is therefore a species of Cyclops, which

Professor G. 8. Brady has kindly identified for meas C. varius

Lilljeborg.

In this connection it may be said that the younger Cyclops

appear to be more easily and more heavily infected than the

adults; the nauplii, however, are never infected.

In addition to these culture experiments, large numbers of

Entomostraca, Oligocheta, and aquatic insect larve from the

reservoir have been examined at various times. In the winter

and early spring no traces of infection were found, although it is

quite possible that they may have been overlooked. In June,

however, out of 117 Cyclops taken at random, 8 were found to

be infected, and 3 out of the 8 contained larve ready to be

transferred to the stickleback. This agrees with Kraemer’s (10)

statement that the life-history takes place in the summer months,

and with the fact that the percentage of adult /chthyotenie is

erveatest at that season. The small percentage of infected Cyclops

found can be easily explained by the fact that the 117 specimens

examined contained examples of many species, but out of these

only O. varius could be infected. By examining the contents of

the alimentary canal of the stickleback, it was ascertained that

in summer they fed chiefly upon Entomostraca, so that a very

small percentage of infected Cyclops would be sufficient to ensure

a heavy infection of sticklebacks. It would thus be quite easy to

overlook this occasional infection unless one were very familiar

with the appearance of the larve.

In order to complete the life-eycle of the parasite, an experi-

ment was started to infect the stickleback. This was a failure.

A number of sticklebacks were obtained from the same pond, and

about 20 were dissected: not one of them showed any trace of

the parasite. Ten of them were then isolated and fed occasion-

ally with chopped-up earthworms and infected Cyclops ; a control

experiment was started under the same conditions, except that

only earthworms were used for food. The experiment lasted

from April 25th to May 16th. From time to time some fish

died, but no trace of the Cestode was found in either those from

the experimental jar or those from the control. On May 7th,

for the first time, a small unsegmented Jchthyotenia was tound

in the intestine of a dead fish; it was obviously in an exceedingly

young stage. On May 16th the rest of the fish were dissected,

but without finding any Cestodes.

TAPEWORM PARASITIC IN THE STICKLEBACK, 129

The failure of this experiment was probably due to the small

number of Cyclops with which the fish were fed (owing to various

causes it was difficult to get infected Cyclops at the time), the

short time between the feeding of the fish and their dissection,

and the small size of the larvee.

The experiment was then repeated, particular attention being

paid to the above points, and this time there was success. It

began on May 27th, when 11 sticklebacks from the same pond

were fed with about 20 infected Cyclops. The following results

were obtained :—

Number Larvee in Number

Date. Dead.* each. | ITntected.

etre We ieee eee cease 4, 0, 4, 7,1 3

rere ee Oth sere erie es, 3 0, 0, 4 1

a AL i ace teh gtd Lag il 2 1

Site GOL geapcecedeose reseee 1 1 i

July Ist 2 8,1 2

11 28 8

A control experiment of five sticklebacks was started at the

same time and gave a negative result.

In order to test the possibility of direct development, ‘an

experiment was started on June 12th. Nine sticklebacks were

isolated and fed with large quantities of adult proglottides of the

Cestode ; in addition, a large number of eggs was put into

the water. The experiment lasted until July 16th, when the

surviving sticklebacks were dissected. None of the nine, how-

ever, showed the slightest trace of infection, either in the liver

or in the intestine.

As a further test, thirteen sticklebacks from the reservoir were

isolated and kept for about six weeks. ‘They were then dissected,

but the only trace of infection found was a nearly adult cestode

in the intestine of one of them. It is practically certain, con-

sidering the heavy percentage of intection, that at least half these

were infected at first; the worms must therefore have been

ejected and have shed their eggs into the water. If direct

infection occur, then the sticklebacks should have been re-

infected, but since no infection was found, the conclusion is that

direct infection does not take place. La Rue (11), in attempting

to show direct development in P. filaroides from the salamander,

failed to infect the salamander itself with eggs of the Cestode,

although in his case the conditions were certainly not very

favourable.

* Both in this, and in the following experiments, a fungus (Saprolegnia ferox)

obtained entrance to the experimental jar and killed the fish, so that the adult

Ce:tode could not be bred.

Proc. Zoou. Soc.—1914, No. IX. 9

130 MR. F. J. MEGGITT ON A

Development.

The development of the egg has not been fully worked out, but

the results so far show a close similarity to those of van Beneden

(2), who used Tenia serrata.

Both fresh and stained material have been used; the results

obtained by the use of the former give a connected series, but

thers are gaps between the stages observed in stained material.

Kraemer (10) appears to be the only investigator who has

described the several stages in the development of the egg, but

my own material does not agree at all with his description.

The following results were obtained from fresh unstained

material.

When the eges pass into the uterus, they appear to consist of

a transparent ‘colourless membrane, ‘021 mm. dia., containing a

roughly spherical mass of uniform refractive grey yolk-granules ;

this inner mass lies freely within the membrane, which only

touches it at one or two points. Both the membrane and the

inner mass increase ia size, but the former more rapidly than the

latter, so that later on, the impression given is that of a grey

ball, 025 mm. dia., within a large bubble ‘047 mm. dia, (Pl. I.

fig. 15). From this point the membrane always preserves the

same relative distance between it and the inner mass, and finally

becomes the outer membrane of the onchosphere (m.). The yolk-

follicles now begin to disappear, clustering together at one pole,

the remainder of the inner mass being filled up by a fine grey

granular substance, from which the agree ea hooks appear to

originate.

Meanwhile the inner mass has become aifercnttatd into a

central mass, °023 mm. dia.—the future onehosphere—within an

outer granular coat, *028 mm. dia. and ‘002 mm. thick (Pl. II.

fig. 17), which at first touches it at every point, but later shrinks

away to leave the onchosphere entirely free. By the time the

yolk-follieles have disappeared the egg has become ready for

discharging (Pl. II. fig. 19). It then consists of the oncho-

sphere, ‘023 mm. dia., provided with six hooks, and closely sur-

rounded by a delicate membrane. This is surrounded by the

granular second membrane (m’.),°0385 mm. dia. and -002 mm.

thick, the whole free within a transparent membrane (m.),

°058 mm. dia. The hooks are :011 mm. long, ‘001 mm. at their

broadest part, and the curved part is ‘(004 mm. long. Treatment

with a solution of methyl green in | per cent. acetic acid shows

that the second membrane has a circular poe: (0057 mm. dia.,

in it.

Sections of the cestode killed with ete anle eels solution

and stained with iron hematoxylin give the best detailed

results.

The ovarian eggs (Pl. alate fig. 24) are ‘017 mm. dia., very

granular, and are surrounded by : a distinct membrane. A clear

nucleus, ‘0092-0103 mm. dia., is present, containing a dark-

TAPEWORM PARASITIC IN THE STICKLEBACK. WBA

staining nucleolus, °0057—-0069 mm. dia. ; often granular strands

connect the nucleolus to the nuclear membrane. In addition,

there is a dark-staining “* Nebenkorper” or “ corps lenticulaire ”

of irregular shape and indefinite size. It seems to disappear

before the first division and is of unknown significance. Von

Linstow (14) has figured a similar body in the eggs of Tenia

longicollis Rud.

The first division of the fertilised egg (Pl. III. fig. 25) i

into a vitelline cell crowded with yolk-granules and a “ (Ciel

zelle,” with nucleus and nucleolus. Kraemer for 7’ jilicollis, and

von Linstow for 7’. longicollis, figure the vitelline globules as

isolated, not as being contained in a single cell.

The d Keimzelle ” next divides into si, and each of these into

two again, four cells of equal size being formed (PI. ILI. fig. 29),

°012 mm. dia., with nuclei ‘(005 mm. dia, In van Beneéden’s

account, these cells are of unequal sizes, two large macromeres

forming the ‘‘ couche albumineuse,” and two small micromeres

forming the onchosphere with its membranes and hooks. In this

case there is certainly a “couche albumineuse” formed (PI. III.

fig. 31), although as the four cells are identical, it is impossible to

distinguish those from which it arises. The remaining cells

divide repeatedly, forming a cellular mass, the nuclei of which

are of two sizes :—

(1) nucleus :0057-:0069 mm. dia., nucleolus -0023—-0034 mm. dia.

Qin. "004-0046 ,, ,, As ‘0011 mm, dia,

The cells containing the larger nuclei surround the others and a

split appears between the two (Pl. IIT. fig. 30). This gradually

widens until the first cells form a definite coat—the second oncho-

spheric membrane—vound the smaller ones (Pl. IIT. fig. 32). It

is thus probable that the former correspond to van Beneden’s

“Chitinogenzellen.” The origin of the hooks and the third

onchospheric membrane I have not been able to discover.

The vitelline cell, produced by the division of the ovarian egg,

does not exist long as a distinct cell, and soon degenerates into a

mass of yolk-follicles. It is these follicles which scatter when

the onchosphere breaks through the second membrane (PI. IIT.

At the time of discharge, the onchosphere is formed of

number of spherical cells, ‘0057 mm. dia., with distinct nuclei,

"0034 mm. dia,, the cells at the posterior end being slightly larger

than those at the anterior.

When the contents of the majority of the proglottides are

ready for discharging, the Cestode slightly migrates through the

intestine, probably by the contractions of the latter, and “hangs

out of the anus. The egos are then expelled from the uterus

through a slit caused by a rupture of the ventral body-wall.

When the contents of the exposed proglottides have been dis-

charged, the body hangs further out, exposing more segments,

until all the ripe eggs have been extruded. If stickle-

32 MR. F. J. MEGGITT ON A

backs infected by this parasite be kept in an aquarium, it is no

uncommon sight to see them with °5'—-2" of cestode hanging out

of the anus. The proglottides first emptied degenerate, and all

traces of segmentation become obliterated, until by the time the

last eggs are discharged, the distal end has become an unrecog-

nisable shapeless mass. The whole cestode is then discharged

with the feces, and this is often followed by the death of the

fish.

Some time after entering the water, the onchosphere escapes

through the hole in the second membrane to lie freely inside the

outer one. Since the diameter of the hole is less than that of the

onchosphere, the latter is considerably contracted in its passage :

the third membrane is often dragged out with it, but it may

he ruptured and left behind, The cause of this escape may be

possibly due to osmosis, but it is more probable that it is due to

the onchospheric movements about to be described. The poste-

rior portion of the onchosphere is always under the aperture

in the second coat, so that the movements would have a ramming

effect upon the delicate third membrane, and either rupture it or

drag it out. The pressure of the coverslip is not the cause, for I

have often seen similar movements in onchospheres lying freely

in a watch-glass. . Yolk-follicles are often scattered within the

membrane at the time of the escape.

The onchosphere now exhibits curious movements (I have only

seen them in the free onchosphere, but it is fsrobable that they

occur also before the escape takes place). It slowly contracts

Text-figure 5,

Drawing to show movements of onchosphere.

antero-posteriorly, stretching out its hooks as far as possible; the

extreme anterior point often forms a small depression surrounded

by avidge. With a quick jerk it elongates itself along the same

line, at the same time striking downwards with its hooks until

they lie flat against the body. It then contracts as before. The

movements are repeated indefinitely, and are probably for the

ultimate purpose of attaching the onchosphere to the alimentary

canal of its host.

La Rue (11) reports similar movements in the onchospheres of

TAPEWORM PARASITIC IN THE STICKLEBACK. 133

P. filaroides. In this ease though, the movements of two pairs

of hooks alternate with those of a third pair, and the body

changes from a spherical to a pear-shaped form. He also saw

these movements in onchospheres of Hymenolepis nana.

At this stage the onchosphere is swallowed by a Cyclops. It

appears to pass through the stomach and first portion of the

alimentary canal without any change or delay, and although it

generally anchors itself to the wall about the junction of the

thorax and abdomen, I have seen cases where it has attached

itself to the wall of the alimentary canal just within the stomach

on the one hand, and just above the anus on the other, but these

are not common. After a certain time has elapsed, the extent of

which is very variable but is usually a week, it breaks through

the wall of the intestine into the dorsal sinus and is swept for-

ward to lie usually near the posterior end of the carapace, but

often in the head above the eye; it may also lie under the

alimentary canal, but this is rare. Here it becomes very vacuo-

lated at first, but later has a more solid appearance (PI. 1V.

fig. 33). The hooks gradually disappear, and the body becomes

partially covered with highly refractive granules, either isolated

or disposed in clusters. These granules greatly resemble those

found on young larve in the intestine of the stickleback, except

that they are smaller.

At the end of three weeks the larva is ready to be transferred

to its second host (Pl. IV. fig. 37). It isan elongated grey body

of variable size, and in situ seems to be an undeveloped part of

the ovary. It is apparently studded with the refractive granules

just mentioned, 004 mm. dia., which give it a characteristic

appearance when removed from the Cyclops. In sections it may

be seen that these really occupy spaces in the body parenchyma

just under the cuticle and correspond to the calcareous bodies of

older forms. There is no scolex and no neck, the body being

entirely without divisions. Atthe anterior end are four suckers,

‘03 mm. dia., which look like small blisters of the cuticle; they

are probably functionless at this period. In sections they are

practically level with the rest of the body. They are composed

of the same parenchyma as the rest of the body (Pl. IV. fig. 35)

only in a more compact form, and have two dark-staining types

of cell. The first type is elongated, -005--007 x ‘001 mm. It is

present only in larve in an advanced stage of development, and

is probably an embryonal muscle-cell. The other type is the same

in size and form as the cells of the parenchyma. In younger

cases the suckers appear like the rest of the parenchyma, from

which, in transverse sections, they are only separated by a thin

line. There is no trace of the fifth sucker. The scolex is not

invaginated at any time during the development; it appears as

if it were gradually differentiated from the body parenchyma

at an early stage.

The whole body is covered by a cuticle, extending also over

the suckers and excretory pore (PI. LV. fig. 36). Its outer layer

134 MR. F. J. MEGGITT ON A

is rough and ragged like that of the adult. Under it is a smooth

homogeneous layer, ‘(0018-0029 mm. thick. Owing to the small

size of the larva I have been unable to find any definite traces of

muscles. Muscles must undoubtedly be present, however, for the

larva is capable of sluggish contractions similar to those of the

adult.

The body within the cuticle is a nearly solid mass of paren-

chyma. Just inside the cuticle the parenchyma forms a fairly

compact sheath, but in proportion to its distance from the exterior

it loses this compactness and becomes vacuolated, the centre itself

being a fairly loose meshwork like that of the adult. The paren-

chyma between the suckers is firmer than that elsewhere. The

parenchyma itself is a faintly staining granular mass, forming a

more or less definite meshwork through which dark-staining cells

are scattered (Pl. IV. fig. 35). It is not probable that these are

nuclei owing to their comparatively large size, °(003--005 mm. dia, ;

inside them are often several darker staining bodies. In addition

to these are smaller bodies like the former only smaller, ‘0016-—

-0025 mm.; they are also present in young suckers, and may

possibly be nuclei. Occasionally circular granular bodies,

‘004 mm. dia., not so deeply staining as the others, are to be

seen. These I take to be the refractive granules (calcareous

bodies) apparently present on the surface of the larva. Running

in all directions through the parenchyma are delicate fibres,

which may possibly be the rudiments of muscles. It is ex-

ceedingly ditticult to state the histological significance of all

these different structures owing to their small size and general

indefiniteness.

At the posterior end of the larva is a small depression running

-016 mm. into the body, its apex expanding into an oval chamber,

with its long axis parallel to that of the larva (PI. IV. fig. 36).

On all sides but one, this invagination is enclosed by the body,

but on the fourth side it appears to be open to the exterior; if

seen from this side, it shows the entire depression—in shape like

the gullet and buccal groove of Parameaciwm—which is not

visible from the other side (Pl. IV. figs. 39, 40). The cuticle

covers its exterior, so that it is a true invagination. Presumably

this corresponds to the heart-shaped bubble in which, according

to Kraemer, the longitudinal excretory canals open. I have been

unable to see any signs of excretory organs, however.

There are no traces of reproductive organs in such a young

stage.

The presence of the larva is fatal to the Cyclops, apparently

causing its starvation. The orange globules usually present in

the head of the Cyclops vanish, and the ovary degenerates and

dwindles to half its usual size. The activity of the Cyclops

also suffers ; it does not swim so rapidly and takes much more

frequent and longer rests than its fellows. Ultimately it dies.

This sluggishness and death would be favourable to the larva, for

TAPEWORM PARASITIC IN THE STICKLEBACK. 135

it would result in the easier capture of the Cyclops by the stickle-

back, and thus ensure the larva a better chance of attaining

maturity.

The Cyclops is swallowed by the stickleback, and in the intestine

of the latter the larva is set free to grow into the normal adult.

SUMMARY. -

1. The paper contains a description of a tapeworm parasitic in

Gasterosteus aculeatus Artedi. I regard this tapeworm as Jchthyo-

tenia filicollis Rud., though the absence of an adequate

description of the species and the confusion into which the

nomenclature has fallen, make the identification a matter of

considerable difficulty. The synonym, Proteocephalus Wein.,

used by La Rue (12), does not seem to me justifiable.

2. An account is given of the anatomy and histology of the

species. With the exception of one or two details, this account

agrees closely with that given by Benedict (3) and by Kraemer (10).

3. Though many conjectures have been made with regard to

the life-history of /. filicollis, no reliable information has hitherto

been published. The evidence given in this paper shows that

Cyclops varius Lillj.is the intermediate host. This conclusion

is based upon successful experiments to infect the Cyclops hy the

tapeworm egg, and to infect the stickleback by the larva In the

Cyclops. Direct infection of the stickleback proved impossible.

4. A short account is given of the segmentation of the egg

and the development of the larva. Though incomplete in some

respects, it was found that it followed the general course described

by van Beneden (2).

LITERATURE.

(1) Bertincuam.—Annals of Nat. Hist. vol. xiv., 1844.

(2) Benepen, E. van.—‘‘ Recherches sur le développement

embryonnaire de quelques Ténias.” Arch. de Biol.

vol. ii. 1881; Journ. R. Micr. Soe. vol. 1.

(3) Benepicr, H. M.—* On the Structure of Two Fish Tape-

worms from the genus Proteocephalus Weinland, 1858.”

Journ. Morph. 1900, vol. xvi. No. 2.

(4) Buarnvitie.—Dict. des Sci. Nat., t. Ivii. p. 552.

(5) Braun, M.—Vermes. Abt. 1 6. Cestodes, in Drs H. G.

Bronn’s “Klassen und Ordnungen des Tierreichs.”

Leipzig, 1900.

(6) Copsotp.—Transact. Linn. Soc. xxu., 1856.

(7) Drestne, C. M.—‘‘ Systema Helminthum.” Vienna, 18950,

vol. 1.

(8) Dusarpiy, F.—‘ Histoire Naturelle des Helminths ou Vers

Intestinaux.” Paris, 1845,

136 MR. F. J. MEGGITT ON A

(9) Iresstinc, Fr.—‘* Ueber den Bau von Schistocephalus

dimorphus Crepl. und Ligula simplicissima Rud.” Arch.

f. Naturgesch. 48. Jahrg. 1 Bd., 1882.

(10; Krammer, A.—‘ Beitrige zur Anatomie und Histologie der

Cestoden der Siisswasserfische.” Zeitschr. f. wissen. Zool.

Bd. 53, pp. 647-722.

(11) La Rus, G. R.—* On the Morphology and Development of

a new Cestode of the genus Proteocephalus Weinland.”

Studies from the Zool. Lab. of the Univ. of Nebraska,

ING, HH}, OS).

(12) La Ruz, G. R.—“ A Revision of the Cestode family

Proteocephalide.” Zool. Anz. Bd. xxxvii., Nov. 1911.

(13) Leuckarr, R.—“< Die Parasiten des Menschens,” 2 Aufl.

Leipzig u. Heidelberg, 1879-86.

(14) Linsrow, O. von.—‘* Ueber den Bau und die Entwicklung

von Tenia longicollis Rud.” Jena. Zeitschr. 1891,

Bd. xxv.

(15) Lonnperc, E.—“ Ueber eine neuer Tetrabothriumspecies

und die Verwandschaftsverhiltnisse der Ichthyotenien.”

Central. f. Bakt. u. Parasit. 1894, vol. xv. No. 21.

(16) Linz, Mi—* Zur Kenntnis einiger Distowen.” Zool. Anz.

(17) Linz, M.—Cestodes, Heft 18, in Dr. Brauer’s “Die

Siisswasserfauna Deutschlands.” Jena, 1910.

(18) Monviceut, F. §.—‘ Notizie su di alcune specie di Tenia.”

Boll. Soc. di Natural. in Napoli, Ser. I., Ann. 5, vol. v.

1891, fase. 2.

(19) Rartuer.—Centralb. f. Bakt. u. Paras. Bd. xxvi., 1899.

(20) Riccenpacn, E.—“Das Genus Jchthyotenia.” Inaugural-

Dissertation, Geneva; Revue Suisse de Zool. 1896,

Bad. iv.

(21) Rupotrn1, E. A.—“Entozoorum sive Vermium In-

testinalium Historia Naturalis,” vol. ii. Amsterdam,

1810.

(22) ScunerpER, G.—“ Beitriige zur Kenntniss der Helminthen-

fauna des Finnischen Meerbusens.” Acta Soc. pro Fauna

et Flora Fenn. xxvi. No. 3, 1904. ;

(23) v. SteBoup.—“ Ueber die Band- und Blasenwiirmer.” Leipzig,

1854.

(24) v. Srspotp.—“ Mémoire sur les Vers rubannés et vésiculaires

de homme et des animaux.” Annal. des Sci. Nat. 1855,

t. lV.

(25) Werntanp, D. F.— Human Cestoides. An essay on the

Tapeworms of Man, etc.” Metcalf & Co., Cambridge,

Mass., 1858.

(26) Zscuoxke, F.—“ Recherches sur lorganisation et la distri-

bution zoologique des Vers parasites des poissons d’eau

douce.” Arch. de Biol. t. v., 1884.

TAPEWORM PARASITIC IN THE STICKLEBACK, Navy

EXPLANATION OF THE PLATES.

Puate I.

Figs. 1-7. Series of scoleces, showing variation in shape and size. Fig. 3 is the

normal form. XX 125.

Fig. 8. Optical section of a proglottis. c.s., cirrus - sac; g-8.. genital sinus;

lc, longitudinal excretory canal; ov., ovary; v., vagina; vas., vas

deferens; vit., vitellaria; w., uterus. X 185.

9. Longitudinal section of cirrus-sac. ¢.m., circular muscles ; e.w., outer, and

i.w., mer wall of cirrus-sac ; U.m., longitudinal muscles; 7.m., retractor

muscles; vas., vas deferens. XX 590.

10. Transverse section of cirrus-sac, lettering as above. e.g. gland. X 870.

11. Longitudinal section of odtype. o.d., oviduct; O.T., odtype ; Sh.G., shell

gland; w.’, specialised part of uterus. > 870.

Prats II.

Fig. 12. Longitudinal vertical section through cuticle. ¢., ¢.’, outer and inner

layers of cuticle; c.c.m., circular, c.l.m., longitudinal cuticular muscles ;

i.l.m., inner longitudinal muscles; i.s., intercellular space; p.c., cell of

parenchyma; s.c., cell of sub-cuticula. XX 600.

13. Longitudinal section through vaginal aperture. s.m., sphincter muscle ;

v.é., vaginal epithelium; v.g., vaginal gland; w.m., vaginal muscle.

x 870.

14, Longitudinal section through odclapt. c.e., cubical epithelium of oviduct ;

0.é.5 outer oviducal epithelium; o.m., oviducal muscle; ov., ovary ;

ov.b., connecting part of ovaries; s.a., muscle of odclapt. x 870.

Figs. 15-19. Stages in the development of theegg. _m., first, m.’, second onchospheric

membrane; 0., onchosphere; 0.h., onchospheric hooks ; #., aperture in

secohd egg-membrane; y., yolk-follicles. < 870.

Prate III.

Figs. 20-22. Stages in the rupture of the second onchospheric membrane. X 870.

Fig. 23. Onchosphere after rupture of second membrane. m., m.’, m.’’, first, second

and third onchospheric membranes. XX 870.

24, Hggs from ovary. e.g, Egg membrane; m., nucleus; m., nucleolus ;

mk:, “ Nebenkérper.” 870.

25. First division of egg. k.m., “Keimzelie”; mm, first onchospheric

membrane; 7., nucleus; ~.’, nucleolus; v.c., vitelline cell. XX 970,

26. Later stage, showing division of “ Keimzelle” nucleus. X 979,

27. Stained section of fig. 26. 870.

28. Section of three-cell stage. 870.

29. Four-cell stage. Only three of the four products of “Keimzelle” division

are Shown. v.c., vitelline cell. 870.

30. Section of older egg. ch., “Chitinogenzellen”; 0., onchosphere ; sp., split

between the two; v.c., vitelline follicles. x 870.

31. Older ege. ¢.a., “couche albumineuse.”

32. Section of uterine egg. X 870.

)

Prate IV.

Fig. 33. Cyclops varius, containing young larve. J., larve. X 50.

34. Transverse section of the Cyclops. X 142.

35. Longitudinal section through head of a larva from Cyclops. ¢., ¢.’, outer

and inner layers of cuticle; m., m.’, the two types of parenchymatous

cell ; 0.9., space containing calcareous body: s., sucker. X 870.

36. Longitudinal section through posterior end of larva. é.p., excretory pore.

X 830.

37. Larva from Cyclops varius. s., sucker; c.b., calcareous body. > 155.

38. Larva from an artificially infected stickleback. * 96.

39. Young larva from a stickleback. X 95.

40. Older larva. xX 95.

[ Norz.—Since sending in this paper, I have been able to obtain

a copy of Barbieri’s memoir (Central. f. Bakt. u. Pavas., Abt. i.

138 ON A TAPEWORM PARASITIC IN THE STICKLEBACK,

1909, Bd. xlix.), in which he records a cyst of I. agonis from the

mandible of Bythotrephes; Fuhrmann also records from Planaria

lactea a plerocercoid which he regards as that of a species of

Ichthyotenia. These cases further strengthen the assumption

that an intermediate host is necessary for the development of all

species of [chthyotema. |

PZ.S 1914.NICOLL.PL.L

= SFP gre

=) 2.

London Stereoscopic Co. imp.

M. Rhodes, del.

TREMATODE PARASITES.

PZ.5.1914.NICOLL.PL.IL.

WRN i ae RS

M. Rhodes and G. Roberts, del.

TREMATODE PARASITES.

PZ.S.1914.NICOLL.PL.I.

Nk:

pee

)

& 4

, :

Se A ae te ae

Paes i

RE OW arg

Ja ARAt St 2.3

M. Khodes and G. Roberts, del.

TREMATODE PARASITES.

PZ.5.1914. NICOLL.PL.IV.

Syst

oes e ws tenth fe? i

Wiig ee

| “Misseeg see Kad ©

& oe bs «

oye ty 228 owe * &) ; “=

ong SipgferreaRe Litas HPC we >

NN UOT Meee wget tiren, op ttt

oy ber a TE pegecd et =F

T Toenail Rs SS

ee oe ee

Be :

1S)

peo Ange sunt 2 dere.

ane .Y . “

OP ete teehee *, Coa cau y

ras ta ler)

=a =

es aN

mm LRN II Oa GY

~ ott

a : Ses pees SS aie iO =

as ee me Se — =

LO * ey ae

erties te gpesas see :

\ MNS GRRE Steins ecg Mo abate bt a f=)

aa) oo ox]

1, x

-M. Rhodes and G. Roberts, del.

pascopic Co, imp

London Stere

TREMATODE PARASITES.

ON TREMATODE PARASITES FROM THE SOCIETY'S GARDENS. 139

9. Trematode Parasites from Animals dying in the

Zoological Society’s Gardens during 1911-1912. By

Wiiiam Nicott, M.A., D.Sec., M.D., F.Z.8.

[Received November 3, 1913: Read March 3, 1914. |

(Plates I.-I'V.*)

> heal

ETHOLOGY : ASIDE Be

‘Trematode parasites from animals in London Zvological Gardens... 189-154

GEOGRAPHICAL ZOOLOGY :

North America: Striped Snake (Tropidonotus ordinatus) ; new trematode

from intestine ...... . 140

North Africa: Spiny -tailed “Mastigure (Uromastix lacanthinurus) + 1 new

trematode from intestine (7th Nov. MOURN ered tar chentet ea aot beaten 141

South America: Schott’s Snake (Philodryas schotti); new trematode

frome Waves (AON IE WOM) nsocoscov saa csccusassbegcesanpeassesnessecceon LAH

Europe: Asp Viper (Viper aspis); new trematode from intestine

(7th June, 1912) ..... Nis oN) Roma lel. WAIN eet ed 1233

India: Indian River-Snake (Tropidonotus piscator) 5 new trematode from

intestine (4th December, 1911) ......... 144

Moluccas: Purple-capped Lory (Lorius domicelta) 5 ; new y trematode ‘from

liver (22nd November, 1911)... : 146

Guiana: Crested Curassow (Craw ‘alector) 3 new ‘trematode from ‘liver

(26th Feb. 1912) .. 4 147

North America: Summer Snake ‘(Contia ‘estiva); new trematode from

intestine (7th July, 1911) .. : 147

North America: Ruddy Flamingo (Phenicopterus ruber); ; new ; trematode

from intestine (6th December, 1911) . a5 148

West Africa: Smyth’s Water- Snake (Gr ayia emythii) 5» new trematode

from intestine (May, 1912) .... 150

Egypt: Egyptian Eagle-Owl Bubo ‘asealaphus) 3 new - trematode trom

intestine (24th October, Ho) eevee coins hg Ra earner a Deno

SYSTEMATIC:

Family LEPODERMATID&.

Mediorima propria, gen. et sp. n., from intestine of Striped Snake ......... 140

Ommatobrephus singular is, (oe et sp. n., from intestine of Spiny-

tailed Mastigure .......... 141

Opisthogenes interrogativus, gen. “et sp. , from: intestine of Schott’s

Snake ....... Peron ae,

Opisthioglyphe adulescens, sp. mes ., from intestine of Asp Vi iper.. vee, 148

Styphlodora persimilis, sp. n., from intestine of Indian River-Snake ...... 144

Family Dicroc@1Liipz.

Lyperosomum scitulum, sp. n., from liver of Purple-capped Lory.. wae 146

Lyperosomum direptum, sp. n., from liver of Crested Curassow ............ 147

Brachycelium obesum, sp. 0., from intestine of Summer Snake

Family EcuinostomMip2.

Echinostomum aliud, sp. n., from: intestine of Ruddy Flamingo ............ 148

Family Crinostom1pD#.

Harmotrema infecundum, gen. et sp. n., from intestine of Smyth’s

LWWiaterssialccl ere nes tinny Sore etene neta ea task cau mids ohatacacte nce meee LO O

Super-family HonostomaTa.

Hemistomum canaliculatum, sp. n., from intestine of Egyptian Eagle-Owl. 151

TUR GH? TB Loss) gee eos donde scan SAP oesandato poi cenolgL canAacuies (oben cHembnnactsacnntceconar Way”

During the course of 1911 and 1912 opportunity was afforded

me of making a fairly complete examination of the viscera of

many of the animals dying in the Gardens. For this I have

* For explanation of the Plates see pp. 153-154.

140 DR. W: NICOLL ON TREMATODE

to thank the courtesy of the Secretary and the officials at the

Prosectorium. The animals were almost exclusively birds and

reptiles or batrachians; no fishes, and only a few mammals were

examined, The collection yielded between twenty and thirty

species, a considerable proportion of which are new. Some of

these have already been described in previous communications to

the Society (Nicoll, 1911, 1912, a, 6).

The reptilian parasites were the most interesting of those

obtained, as several of them represented new generic types. They

include forms from the Striped Snake, the Spiny-tailed Mastigure,

Schott’s Tree-Snake, and Smyth’s Water-Snake.

Usually there was an interval of twenty-four hours, some-

times longer, between the death of the animal and the time of

examination, but in spite of that the parasites in most cases were

in a good state of preservation. In some instances, however, they

were so macerated as to be almost useless. As might have been

expected, parasites were more frequent in those animals which

had been in the Gardens for the shortest period. Except in the

few instances reported by Dr. Leiper (1912), there appears to be

little probability of infection being acquired in the Gardens. In

most cases the infections were not heavy, there being usually

only a few worms present. Gross infections, however, were met

with in the case of some Striped Snakes which were heavily in-

fected with two larval Trematodes (Nicoll, 1912), and a Marsh

Harrier which contained a large number of liver-flukes.

The first form I shall describe is an interesting new species

from the Striped Snake (Zropidonotus ordinatus).

Family LEPODERMATID&,.

1. MeprIorIMA PRopRIA, gen. et sp. n. (PI. I. fig. 1.)

This species is a typical member of the family Lepodermatide,

to which a large proportion of reptilian trematodes belong. One

specimen was obtained from the intestine of a Striped Snake

(Z-opidonotus ordinatus).

It is an elongated, slightly flattened Trematode, both ends of

which are rounded. The cuticle is beset with small spmes, which

extend throughout the whole length of the body. The length is

about 6 mm., the greatest breadth, near the middle of the body, is

1-2 mm.

The globular oral sucker has a diameter of 5 mm.; the ventral

sucker, which is somewhat oval, measures ‘7 xX°8 mm., and is

situated 1-7 mm. from the anterior end.

There is a very short prepharynx, and a pharynx measuring

‘2x17 mm. The esophagus is about the same length as the

pharynx. The intestinal diverticula are short, not extending very

far (-4 mm.) beyond the ventral sucker. Their ends are obscured

by the enormous mass cf ova.

The genital aperture is median and is situated just behind the

PARASITES FROM THE SOCIETY'S GARDENS, 141

intestinal bifurcation. The cirrus-pouch is short and _ stout,

measuring *8x°34 mm. It is almost oval in outline. Within it

there is a small convoluted vesicula seminalis, with a moderately

long pars prostatica and a short ductus ejaculatorius. The vagina

is somewhat shorter than the cirrus-pouch.

The small globular ovary is situated a short distance behind the

ventral sucker on the right side and is largely concealed by the

uterus. The anterior testis lies on the left about 1 mm. behind

the ventral sucker, while the posterior testis les a little further

back on the right side. Both are almost entirely concealed by

the uterus, so that their size and shape could not be determined.

They are, however, apparently elongated oval in outline. The

yolk-glands are scanty, consisting on each side of a little more

than a single row of follicles external to the intestinal diverticula,

and extending from midway between the genital aperture and

the anterior edge of the ventral sucker to the level of the anterior

testis.

The uterus fills up the greater part of the posterior two-thirds

of the body. It consists of a narrow descending limb and a very

greatly distended ascending limb, which is twisted into slight

folds. The extremely numerous ova are dark brown in colour

and measure -039 x ‘018-020 mm.

This species bears a very close resemblance to the genus

Lechriorchis, but is distinguished from it by the position of the

genital aperture and the shape of the cirrus-pouch.

2. OMMATOBREPHUS SINGULARIS, gen. et sp.n. (Pl. I. fig. 2.)

A single specimen of this species was obtained from the

intestine of the Spiny-tailed Mastigure (Uromastia acanthinurus).

It presents several features of interest.

The length of the specimen, slightly pressed, is 2°73 mm., and

the maximum breadth across the ventral sucker is°8 mm. The

anterior part of the body is considerably attenuated, the posterior

part is more rounded but terminates in a somewhat acute point.

There are no cuticular spines.

The subterminal oral sucker has a diameter of °18 mm. and

the ventral sucker measures 53 mm. ‘The latter is situated ata

distance of 1:06 mm. from the anterior end, Contiguous with

the oral sucker is a large pharynx, measuring ‘12°14 mm.

This is followed by a long cesophagus, °3] mm. in length. The

intestinal bifurcation takes place a little in front of the ventral

sucker, and the short diverticula terminate about midway between

the ventral sucker and the testes.

The excretory vesicle is Y-shaped with a short sinuous stem,

bifurcating a little in front of the testes, and limbs extending

nearly to the level of the pharynx.

The genital aperture is median, just over the intestinal bifur-

cation. The cirrus-pouch is stout and almost globular, lying

almost entirely in front of the ventral sucker. Its length and

diameter are about *2 mm, It is thin-walled and contains a thick

142 DR. W. NICOLL ON TREMATODE

highly-convoluted vesicula seminalis with a short prostate and

ductus. The testes lie alongside one another, almost at the

posterior end of the body. ‘They are slightly asymmetrical, the

left being a little in advance of the right, and they are separated

by a narrow fold of the uterus. They are elongated oval in

outline and measure *35 x22 mm.

A little in front of the testes les a small round ovary, some-

what on the right side of the middle line. The shell-gland

complex is entirely obscured by the uterus. The yolk-glands are

of restricted extent, being confined to the extreme edges of the

body and extending only from the ovary to some distance behind

the ventral sucker. The uterus fills up the whole of the middle

of the body and sends a small loop down between the testes. The

eges are thin-walled and very transparent, and the majority of

them contain a fully developed miracidium, the K-shaped eye-

spots of which are very conspicuous. The eggs measure ‘095 x

"056 mm.

This form evidently belongs to the family Lepodermatide, but

it is somewhat aberrant. Its most distinctive features are the

unusual position of the genital glands and the precocious develop-

ment of the miracidia, together with the large size of the eggs.

3. OPISTHOGENES INrTERROGATIVUS, gen. et sp.n. (PI. II. fig. 4.)

Five specimens of this parasite were obtained from the intestine

of Schott’s Snake (Philodryas schotti). In length they vary from

4-5 mm. to 6°2 mm. (pressed specimens); the average is 5°-4 mm.

The greatest breadth across the ventral sucker is 17 mm. The

outline is almost fusiform, the posterior half being more attenuated

than the anterior.

Both suckers are nearly globular, the oral measuring ‘64 mm.

and the ventral ‘56 mm. In both cases the longitudinal diameter

is usually slightly greater than the transverse. The ventral

sucker is situated 1:74 mm. from the anterior end. There isa

short prepharynx followed by a pharynx measuring °21 x-2 mm.

The cesophagus is slightly longer than the pharynx, °27 mm.

The intestinal diverticula terminate some distance (about *6 mm.)

from the posterior end of the body, but their ends are obscured

by the dense mass of the uterus.

The excretory vesicle is long and wide. The median stem runs

forward to near the ovary before dividing into the two short

lateral branches. ‘The vesicle, as well as the main collecting

tubules, are clearly mapped out owing to the pigmented nature

of their contents. A large number of fine branches are given

off from the vesicle, and these form an anastomosing network

througbout the whole body.

The cirrus-pouch has the shape of an interrogation mark,

curving over the left posterior quadrant of the ventral sucker.

Its proximal end lies a short distance behind the sucker, while

the male genital aperture is 1 mm. behind the sucker and a little

to the left of the middle line. The female aperture is separated

PARASITES FROM THE SOCIETY'S GARDENS. 143

from the male but immediately behind it. The cirrus-pouch

contains a highly convoluted vesicula seminalis, a short pars pro-

Statica, and a long straight ductus and cirrus. The musculature

of the pouch and of the cirrus is extremely well developed.

The anterior testis lies on the left side, near the genital

aperture. ‘The second testis lies on the other side of the body,

and is separated from the first by the uterus. The anterior border

of the one is about ‘2 mm. in front of that of the other. Both

are somewhat elongated oval in outline, their long diameter

measuring *5—65 mm. ‘They are moderately thick and are not

much overlapped by the uterus.

The ovary lies not far behind the ventral sucker, alongside the

proximal end of the cirrus-pouch. It is globular and considerably

smaller than the testes. It is usually difficult to see, owing to

its being obscured by the uterus. Laurer’s canal is present, but

there is no receptaculum seminis. he initial part of the uterus

is filled with sperms. The yolk-glands havean unusual disposition.

Instead of being lateral they are entirely dorsal. Situated in

the posterior half of the body, they extend from the level of the

genital aperture to a short distance (‘4 mm.) behind the testes.

They consist of fairly large follicles which le between the ex-

eretory vesicle and the cuticle, and which do not spread to the

outer side of the intestinal diverticula.

Starting from the ovary, the uterus forms several small dorsal

convolutions on its way towards the posterior end of the body.

On turning forward it becomes greatly dilated. At first it runs

almost straight forwards, then bends towards the left testis.

Passing between the testes it forms a fairly large convolution in

front of the right testis. It then passes across the ventral sucker

and finally runs down along the left side of the cirrus-pouch. It

overlaps the intestinal diverticula to a considerable extent. The

vagina is a wide muscular structure about half the length of the

cirrus-pouch. The ova are very numerous, dark brown in colour,

and measure *020—:023 x :013—014 mm.

This species is most closely allied to Opisthogonimus philo-

dryochus West, but it appears to present features of sufticient

importance to warrant its being regarded as the type of a distinct

genus. The chief distinctive features are the position of the

genital aperture, the shape and structure of the cirrus-pouch, and

the position of the yolk-glands.

4, OPISTHIOGLYPHE ADULESCENS, sp.n. (PI. I. fig. 3.)

A few specimens of this form were taken from the intestine of

an Asp Viper (Vipera aspis). They were all obviously immature,

and on that account the following can only be regarded as a

provisional description. For the same reason it is impossible to

be absolutely certain that this form is distinct from the already

known species of the genus, but one or two distinctive features

seem to point to the fact that it is a separate species.

144 DR. W. NICOLL ON TREMATODE

In shape it is elongated oval with somewhat pointed ends and

almost parallel sides. The length is about 1:3 mm. and the

greatest breadth -46 mm. ‘The cuticle is beset for at least three-

quarters of the length of the body with minute spines. The oral

sucker is subterminal and has a diameter of 17 mm. The round

ventral sucker is much smaller, measuring only -10 mm,, and

it is situated at a distance of -55 mm. from the anterior end of

the body.

There is a short prepharynx, a pharynx measuring ‘06 x '08 mm,,

and a long cesophagus, twice the length of the pharynx. The

intestinal diverticula are rather wide, and do not extend more

than halfway between the ventral sucker and the posterior end

of the body, The excretory vesicle is dilated and the median

stem divides a short distance behind the ovary. The paired

limbs only extend to the ventral sucker.

The genital aperture lies a short distance behind the intestinal

bifurcation. The cirrus-pouch is short and plump, reaching only

a short distance behind the anterior border of the ventral sucker.

It contains a highly convoluted vesicula seminalis, a short pars

prostatica, and a short ductus ejaculatorius. The testes lie at the

level of the end of the intestines. They are tandem or slightly

oblique, and almost contiguous. They are of very irregular shape,

and are always broader than long, Their dimensions are about

"06 x13 mm. The post-testicular space is almost exactly one-

quarter of the body-length.

The ovary is contiguous with the ventral sucker on its right

posterior border. It also touches the right intestinal diverticulum,

Tts size is ‘08-06 mm, The yolk-glands are extremely scattered

and imperfectly formed. They consist of small, rather isolated

follicles extending along the sides of the body from the middle of

the esophagus to the middle of the post-testicular space. In the

latter they extend inwards but do not meet in the middle line.

They overlap the intestinal diverticula only to a very slight

extent. The uterus is confined between the testes and the ventral

sucker, and does not overlap the intestinal diverticula, It con-

tains about two dozen eggs, measuring ‘042 x -021 mm.

It does not appear likely that this can be the young stage of

any of the already known species of Opisthioglyphe, for it is

differentiated from O. rane by the length of the intestinal

diverticula and the ratio of the suckers. The configuration of the

excretory vesicle is also different. From O. hystrix it differs in

the position of the ventral sucker, the length of the intestinal

diverticula and of the cirrus-pouch, and the extent of the yolk-

glands. It also differs in several important particulars from the

more recently described O. locellus Kossack. I am unable to

compare it with O. siredonis, as Poirier’s paper is not available.

5, SYYPHLODORA PERSIMILIS, sp.n. (PI. IL. fig. 5.)

About thirty specimens of this parasite were removed from

the ureters of an Indian River-Snake (Zropidonotus piscator)

PARASITES FROM THE SOCIETY'S GARDENS. 145

which died in the Gardens on December 4th, 1911. Owing to

its extreme resemblance to S. serrata Lss. and S. horrida (Leidy),

I have had considerable hesitation in deciding to regard it as

a new species. Coming, however, from such different hosts as

the Egyptian Monitor (Varanus niloticus), Boa constrictor, and

the present host, and from such widely separated parts of the

world as Egypt, North America, and India, the probability is

that they represent three distinct species. It also bears some

resemblance to S. najw Nicoll (1912) from the Indian Cobra,

though the length of the intestinal diverticula and the size of

the suckers appear to be characters sufficient to separate it from

that species.

The length of mature specimens is 3-4 mm. and the maximum

breadth -9-1-0 mm. just behind the ventral sucker. The cuticle

is beset with fairly prominent spines throughout practically its

whole extent.

The oral sucker measures *24--28 mm. in diameter and the

ventral sucker -28--34 mm. The latter is almost invariably

transversely elongated, its average diameter being -29 x -°33 mm.

The oral sucker is sometimes elongated, sometimes transverse,

the latter being usually the case, and its average dimensions are

°25x°26 mm. ‘The ratio of the mean diameters of the suckers is

therefore about 5:6; it is always greater than 4:5 and never

greater than 6:7. In S. najw and S. serrata the suckers are

much more nearly equal. The ventral sucker is situated at a

distance of -6--8 mm. from the anterior end of the body. The

neck thus comprises only about one-fifth of the body-length,

but it may be remarked that the specimens were somewhat

contracted.

There is no prepharynx, and the pharynx measures *17 x -13 mm.

The esophagus is about] mm. long, and the intestinal bifur-

‘ation takes place close in front of the ventral sucker. The

intestinal diverticula are practically equal in length, and extend

to a distance of *65-"9 mm. from the posterior end. The average

in about a dozen specimens was °76 mm., which is almost exactly

two-ninths of the body-length. They are therefore practically

about the same length as those in S. serrata.

The median genital aperture is immediately in front of the

ventral sucker. ‘The cirrus-pouch just reaches the posterior

border of the ventral sucker, and is frequently contiguous with

the ovary. The latter is globular, with a diameter of :2 mm.

Behind it lies the receptaculum seminis. The anterior testis,

on the left, lies about *25 mm. from the posterior border of the

ventral sucker, though the distance varies from -18 mm. to

°38 mm., according to the state of contraction. The posterior

testis, on the right, is separated from the other by a distance of

"17 mm., though this again varies. The anterior border of the

right testis 1s usually a little in advance of the posterior border

of the left.

The yolk-glands have the usual situation, almost entirely

Proc. Zoon. Soc.—1914, No. X, 10

146 DR, W. NICOLL ON TREMATODE

external to the intestinal diverticula, and extend from the

posterior border of the ventral sucker to the middle of the

anterior testis. The uterus, which is voluminous, bears a close

resemblance to that of S. serrata, and the ova measure ‘039-047 x

°019--020 mm.

Family DicRoc@LIIpD&#,

6. LyPEROSOMUM SCITULUM, sp.n. (PI. ITI. fig. 6.)

Four specimens of this species were taken from the liver of a

Purple-capped Lory (Lorius domicella). It is a greatly elongated

form, measuring 6°8-7°2 mm. in length. The breadth is fairly

uniform, but attains its maximum (‘8 mm.) across the ventral

sucker. There are no cuticular spines.

The oral sucker is almost terminal, and has a diameter of

"45 mm.; the ventral sucker is slightly smaller, °42 mm., and is

situated 1°33 mm. from the anterior end. The pharynx is con-

tiguous with the oral sucker, and has a diameter of *14 mm. ;

the cesophagus is about the same length. The narrow tortuous

intestinal diverticula extend down the sides of the body to a

distance of about 2 mm. from the tip of the tail. They are of

unequal length, sometimes the right, sometimes the left being

the longer.

The anterior testis is separated from the ventral sucker by a

space of 33 mm. The posterior testis lies about the same length

behind the first, the latter being close against the right intestinal

diverticulum and the former against the left diverticulum.

They are oval bodies measuring about 25-33 mm. The space

between the second testis and the ovary is equal to that between

the two testes. The ovary is a transversely oval body situated in

the middle line and measuring ‘21 x ‘24mm. The genital aperture

lies over the intestinal bifurcation. The cirrus-pouch is fairly

stout, and extends to the anterior border of the ventral sucker.

It contains a highly convoluted, voluminous vesicula seminalis

and a moderately long prostate and ductus.

The yolk-glands are limited to an area on each side of the

body about the level of the ovary. They are variable in extent,

but as a rule their anterior limit is on a level with the second

testis, while their posterior limit is about half as far again on the

other side of the ovary. They are usually more extensive on one

side of the body than on the other. They almost completely

overlap the intestinal diverticula. The uterus is very highly

convoluted, the loops being mostly transverse. It fills up almost

all the posterior part of the body not occupied by the genital

glands, which it does not overlap. It does, however, overlap the

intestinal diverticula to a very considerable extent. The eggs

-are extremely numerous, and measure °029x-019 mm. The

older ones contain the characteristic miracidium larva.

PARASITES FROM THE SOCIETY’S GARDENS. 147

7. LyPEROSOMUM DIREPTUM, sp.n. (PI. III. fig. 7.)

Three fragments of this species were obtained from the liver

' of a Crested Curassow (Crax alector). From these it was possible

to get a fairly accurate idea of the whole animal. It is about

9 mm. long and about ‘9 mm. broad at its broadest part, which 1s

just behind the ventral sucker. The neck narrows rather

abruptly, but the posterior part of the body gradually narrows to

an acutely pointed tail. Ata distance of | mm. from the tip of

the tail the breadth is less than half what it is behind the ventral

sucker. There are no cuticular spines.

The globular oral sucker measures °35 x °39 mm., the ventral

sucker °46 x °42 mm., and the latter is 1-03 mm. from the anterior

end. In one fragment belonging to a specimen apparently well

over 10 mm. in length, the ventral sucker had a diameter of

54 mm.and the maximnm breadth of the body was 1:1 mm.

The pharynx is contiguous with the oral sucker, and measures

-17x:14mm. There is practically no cesophagus, the intestinal

diverticula branching out directly from the pharynx. The diver-

ticula terminate a considerable distance from the tip of the tail,

but the ends are obscured by the uterus, and could not be made

out.

The genital aperture lies immediately behind the pharynx.

The small slender cirrus-pouch, *3 mm. in length, is slightly

curved, and does not reach the ventral sucker. The testes are

close behind the ventral sucker, the left being in advance, but

only by the distance of its own diameter, so that the posterior

border of one is on the same level as the anterior border of the

other. They are almost globular, and their diameter varies from

"24 mm. to °31 mm.

The globular ovary is separated from the posterior testis by

a space of 6 mm. It lies somewhat to the right side, and is

rather larger than the testis, having a diameter of -34—41 mm.

Behind it lies a small globular receptaculum seminis. The yolk-

glands are much more extensive than in the preceding species.

They reach from the level of the posterior testis to halfway

between the ovary and the tip of the tail. They are more

extensive on one side than on the other. The uterus is very

voluminous and highly convoluted, but the convolutions are not

quite so regularly disposed as in Z. seitulum. The very numerous

eggs measure *025—028 x ‘019-021 mm.

8. BRACHYCG@LIUM OBESUM, sp. n. (PI. III. fig. 8.)

A few specimens of this species were obtained from the intestine

of a Summer Snake (Contia estiva), which died in the Gardens

on July 2nd, 1911.. They were all, unfortunately, in a very

contracted state, and it was only with some difficulty that their

internal anatomy could be made out. At first sight they appeared

to be identical with B. salamandre, but their exceedingly small

size-and the fact that even the smallest was.fully, mature, raised

ets io

148 DR. W. NICOLL ON TREMATODE

suspicion that this could not be the case. It was difficult, how-

ever, to obtain other grounds for regarding them as a distinct

species. These relate only to slight differences in the size of the

suckers, the cirrus-pouch, and the ova.

The body in its contracted state is plump and equally rounded

at both ends. In some specimens each end is curled ventrally.

The length of the smallest specimen was °75 mm. and of the

largest 1-4 mm., though the latter had been subjected to a certain

amount of pressure. In the natural state the length is probably

1-15 mm. The greatest breadth, in the middle of the body, is

‘4-65 mm. ‘The cuticular spines are rather minute.

The suckers are more unequal than in B. salamandre. The

diameter of the oral sucker is -17—25 mm., the average being

22mm. The ventral sucker only measures -11—15 mm., with

an average of -135 mm. ‘The ratio is therefore very nearly 5:3

instead of 5:4, as in B. salamandre. The ventral sucker 1s

also nearer the middle of the body, being about two-fifths of the

total length from the anterior end.

The pharynx measures ‘075 x-07 mm.; the csophagus is

short, and the dilated intestinal diverticula reach the level of the

middle of the ventral sucker.

The genital aperture has the same position as in B. salamandre,

but the cirrus-pouch is much longer and more slender, and it

extends right across the ventral sucker to its posterior border.

Tt contains a very large bipartite vesicula seminalis, of which the

posterior portion is the larger. The transversely oval ovary lies

on the left side caomnerlietielly behind and overlapping the ventral

sucker. Just behind it, and a little externai, lies the left testis,

while on the other side, on the same level, lies the right testis.

The ovary measures -13x‘07 mm.; the testes are somewhat

larger. The yolk-glands extend from about the middle of the

oral sucker to the level of the ovary. The uterus is very volu-

minous, and fills up the whole of the posterior part of the body.

The arrangement of the convolutions cannot be made out, as

they are so closely packed together, but it is noticeable that

most of the young eggs are near the tail, while the older eggs

occupy a more forward position. The eggs are thick-shelled and

somewhat larger than those of B. salamandre, measuring

*050—052 x 034-036 mm.

This species differs from B. hospitale Stafford (1903) in having

relatively larger and more unequal suckers. The intestinal

diverticula are somewhat longer, and the ova are considerably

larger. It is altogether a much smaller species.

Family ECHINOSTOMIDAZ.

9. EXCHINOSTOMUM ALIUD, sp. n. (PI. IV. figs. 9, 9 a.)

Four specimens of this parasite were obtained from the intestine

of a Ruddy Flamingo (Phenicopterus ruber). It is an elongated,

PARASITES FROM THE SOCIETY'S GARDENS. 149

somewhat flat species, with a pronounced cervical concavity. As

in many other species of the genus, the head has a tendency to

be bent sharply on to the ventral surface of the body.

The specimens measured 2°9-6°3 mm. in length. One, 4°6 mm.

in length, was immature, but another, of 5:6 mm. in length, was

full of ova, so that the maturity len ath is probably about 5 mm.

The areatest breadth, across the nee sucker, is about one-fifth

of the length. Behind the ventral sucker the breadth is fairly

uniform, but the tail is distinctly pointed. The breadth across

the head is about one-tenth of the body-length.

There are 35 cephalic spines, arranged in the typical fashion in

two uninterrupted rows, with a group of five terminal spines at

each end. In the anterior row the spines are slightly shorter than

in the posterior row, the average size being ‘(099 mm. and:103 mm.

respectively. The terminal spines are slightly smaller, measuring

"093 mm. The shortest spine on either side is the superficial

spine third from the inner end of the terminal group. In the

only specimen in which its dimensions could be accurately

determined, it measured ‘091 mm. on the right side and only

078 mm. on the left. The cephalic spines present two peculi-

arities which have not apparently been observed in any hitherto

described species of Hchinostomum. In the first place their shape

is unusual. Instead of the common symmetrical peg-shape, their

distal end is inflated in the form of a somewhat triangular knob,

which is slightly sculptured. ‘The knob is situated on the upper

surface of the spine, and comprises about one-third of its total

length. An idea of its shape will best be gathered from reference

to fig. 9a. There is the further peculiarity that each spine is

connected to the edge of the cephalic dise by a thin web-like

membrane which is joined to the spine along pretty nearly its

whole length.

The whole of the neck, both dorsally and ventrally, is covered

with stout cuticular spines, but these do not extend very far

beyond the ventral sucker.

In a 5-6 mm. specimen the large oral sucker measures *22 mm.

in diameter, and the ventral sucker °72 mm. The latter is usually

globular, but in one case it was somewhat deepened. It lies

at a distance of 1°43 mm. from the anterior end. The neck is

thus a little more than one-fourth of the body-length. There

is a short prepharynx about -06 mm. in length, followed by

a pharvnx measuring °19x-13 mm. The cesophagus is about

*46 mm. in length, and the bifurcation occurs just in front of

the ventral sucker. The diverticula are somewhat crenated, and

extend right to the posterior end of the body.

The genital aperture lies in the middle line just over the

intestinal bifurcation. The cirrus-pouch is small and stout, and

overlaps the ventral sucker only to a slight extent. Its dimensions

are 45 x °18 mm., and it encloses a convoluted vesicula seminalis,

a short pars prostatica, and a twisted ductus and cirrus. The

testes are rather small and of irregular contour, there being

150 DR. W. NICOLL ON TREMATODE

usually a slight indentation on either side about the middle of

their length. Their dimensions are -34x°25mm. The posterior

testis is a trifle larger than the anterior one. They are very

close together, but are not quite contiguous. In front they are

separated from the ovary by a space of -15 mm.; behind them

the post-testicular space is 1:68 mm. in length.

The small transversely oval ovary is situated 1:02 mm.

behind the ventral sucker. It is almost median, and measures

"15x:21 mm. The yolk-glands are rather restricted in extent.

On each side they reach the posterior border of the ventral

sucker ; posteriorly they cease at some distance from the tip of

the tail. They are for the most part confined to the outer side of

the intestinal diverticula, and overlap them only to a very slight

extent. The transverse yolk-ducts cross between the ovary and

anterior testis. The uterus does not overlap the intestinal

diverticula, and is confined between the ovary and the ventral

sucker. The ova are very large, measuring *114—-122 x -069-

074 mm.

Family CLINOSTOMIDA.

10. HARMOTREMA INFECUNDUM, gen. et sp.n. (PI. IV. fig. 10.)

A considerable number of specimens of this species was ob-

tained from the intestine of Smyth’s Water-Snake (Grayia

smythii). It is one of the most remarkable forms in the present

collection.

It is a small, rather flat species, white in colour and with the

edges of the body thrown into irregular wrinkles. The cuticle

is unarmed.

Tt is about 2 mm. in length, with a fairly uniform breadth of

-6 mm. Both ends are slightly pointed. The suckers are small

and feeble, the oral having a diameter of -13 mm. and the

ventral 08 x ‘10 mm. The latter lies °8 mm. from the anterior

end. : ‘Ei

There is no prepharynx; the pharynx measures ‘06 x ‘04 mm.,

and the csophagus is about the same length as the pharynx.

The intestinal diverticula are wide and sinuous, and reach the

posterior end of the body. Behind the ventral sucker they

usually bend inwards before passing out again to make room for

the genital glands. At their termination they again turn in

towards the middle line.

The genital aperture is situated on the left side of the body,

internal to the intestinal diverticulum and almost midway

between the ventral sucker and the posterior end of the body.

The curious thin-walled cirrus-pouch stretches forwards from

the genital aperture in a zigzag or sinuous fashion, and its

proximal end lies against the right intestinal diverticulum. It

contains a bipartite vesicula seminalis, of which the proximal

part is oval and the distal elongated. There is a moderately

PARASITES FROM THE SOCTETY’S GARDENS, 151

long fusiform pars prostatica and a long curved ductus ejacu-

Jatorius.

The testes are two irregularly lobed bodies lying behind the

cirrus-pouch. The anterior is just behind the genital aperture

but on the right side of the body. The posterior hes between

the ends of the intestinal diverticula, and is separated from the

end of the body by a space equal to its own diameter.

Between the testes les the much smaller ovary. It is also

irregularly lobed, but not so distinctly as the testes. It is

separated from the posterior testes by a large yolk receptacle.

The yolk-glands extend a considerable part of the length of the

intestinal diverticula and le both internally and externally to

them, but not to any great extent dorsally or ventrally. Their

anterior limit is midway between the intestinal bifurcation and

the ventral sucker. Posteriorly they cease at the ends of the

intestines. The uterus is extremely short and does not usually

contain more than two eggs. The latter are comparatively huge

but very variable in size. The normal size appears to be about

"15-18 x:08-11 mm. They lhe alongside the ovary and testes.

This species evidently bears a close resemblance to the genus

Harmostomum, but differs from it particularly in the position of

the uterus and the size of the eggs.

HoLoSstTOMATA.

11. HemistomuM CANALICULATUM, sp.n. (PI. IV. fig. 11.)

A few specimens of this parasite were obtained from the

intestine of an Egyptian Eagle-Owl (Bubo ascalaphus). It is

of moderate size, 1:6-2°9 mm. in length. The anterior part of

the body is considerably longer than the posterior part, in the

vatio of 5: 3 on an average. When expanded the anterior part

cf the body is almost oval in outline, with a pointed tip. The

lateral glandular pits are not very well marked. When con-

tracted the body is almost uniformly cylindrical. The long axis

of the two parts are almost in the same straight line; so that

there is little dorsal flexure. The posterior part of the body

may attain a breadth of ‘7 mm.

The small oral sucker measures ‘08-09 mm. The ventral

sucker is slightly larger (‘1 mm.), and is situated at a distance

of -6 mm. from the anterior end in a specimen 2°4 mm. long.

The pharynx is contiguous with the oral sucker and somewhat

larger than it, measuring “08-1 x -06--08 mm. The short

esophagus measures only -03—C4 mm. The diverticula have

the usual configuration.

The most characteristic feature of the species is the clinging-

plug, which is situated immediately behind the ventral sucker.

It is an oval structure about four times as long as the ventral

sucker and two or three times as broad. It does not conceal

152 DR. W. NICOLL ON TREMATODE

the ventral sucker, but in the older specimens it overlaps the

posterior border of the sucker. It is not much raised above

the surface of the body, and is traversed by a deep median

longitudinal furrow which extends along its whole length. In

well-expanded specimens the furrow is much shallower. A

similar furrow exists in H. pileatwm (Rud.).

The genital glands do not display any striking peculiarities,

except that the testes are reniform in transverse section and the

yolk-glands extend forward only to about ‘5 mm, from the

anterior end. Their auterior edge forms a semi-circle with the

convexity directed forward.

The ova measure “105—1] x -065- 07 mm.

This species bears most resemblance to H. spathula (Creplin),

but differs from it in several important details. In the latter

the two parts of the body are nearly equal; the oral sucker and

the pharynx are only half as large, while the clinging-plug is

much larger and does not display the longitudinal furrow. The

yolk-glands, too, are probably more extensive.

The following is a list of the hosts and of the Trematode

parasites obtained from them :—

MAMMALS.

ANOA DEPRESSICORNIS.

Fasciola gigantea (Cobbold) ..................... Laver.

BIRDS.

CIRCUS HRUGINOSUS. Marsh Harrier.

Metorchis crassiusculus (Rud.)................... Liver and Gall-bladder.

BuBo ASCALAPHUS. Kgyptian Eagle-Owl.

Hemistomum canaliculatum Nicoll ............ Intestine.

LaRuUS RIDIBUNDUS. Black-headed Gull.

Gigantobilharzia acotylea Odhner ............ Mesenteric veins.

LARUS FUSCUS. Black-backed Gull.

Tocotrema lingua (Creplin) ..... ........-.....- Intestine.

LARUS ATRICILLA. Laughing Gull.

Tocutrema lingua (Creplin) ...... .............. Intestine.

ALCA TORDA. Razorbill.

Hemistomum pileatum (Rud.) ................-. Intestine.

Metorchis xanthosomus (Creplin) ............... Gall-bladder.

(EDEMIA NIGRA. Black Scoter.

Catatropis verrucosa (Froelich) ............... Ceea.

Paramonostomum alveatum (Mehlis) ......... Intestine.

Lorivs DOMICELLA. Purple-capped Lory.

Lyperosomum seitulum Nicoll ............... Liver.

PHENICOPTERUS RUBER. Ruddy Flamingo

Echinostomum aliud Nicoll ..................... Intestine.

CRAX ALECTOR. Crested Curassow.

Lyperosomum direptwm Nicoll ............0:-. Liver,

PARASITES FROM THE SOCIETY'S GARDENS. 153

REPTILES.

TROPIDONOTUS ORDINATUS. Striped Snake.

Cercaria ordinata Nicoll .............s.s02 see eee Mesentery.

Diplostomum sirtale Nicoll ............-:2+0++ Mesentery.

Mediorima propria Nicoll .............:.....0+ + Intestine.

TROPIDONOTUS PISCATOR. Indian River-Snake.

Styphlodora persimilis Nicoll ..................Ureters.

TROPICONOTUS NATRIX. Common Snake.

Cercorchis nematoides (Muhling) ............... Intestine.

NaAJA TRIPUDIANS. Indian Cobra.

Xenopharyna solus Nicoll ......0....2000..... Gall-bladder.

Styphlodora naj@ Nicoll .......................... Ureters.

PHILODRYAS SCHOTTI. Schott’s Snake.

Opisthogenes interrogativus Nicoll ............ Intestine.

GRAYIA SMYTHII. Smyth’s Water-Snake.

Harmotrema infecundum Nicoll ...... ...... Intestine.

VIPERA ASPIS. Asp Viper.

Opisthioglyphe adulescens Nicoll ..............._ Intestine.

Cercaria ordinata Nicoll ..................+:.-.. Mesentery.

ContTIs HSTIVA. Summer Snake.

Brachyceelium obesum Nicoll ....................._ Intestine.

URoMASTIX ACANTHINURUS. Spiny-tailed Mastigure.

Ommatobrephus singularis Nicoll ............ Intestine.

References.

Kossack, W. 1910.—Neue Distomen. Centralbl. fiir Bak-

teriologie, &c., 1 Abt. Orig. lvi. pp. 114-120.

Lerper, R. T. 1911.—(Demonstration of Nematode Parasites).

Proc. Zool. Soc. 1911, pp. 620-621.

Nicott, W. 1911.—On three new ‘Trematodes from Reptiles.

Proc. Zool. Soc. 1911, pp. 677-686.

Nicott, W. 1912a@—On two new larval Trematodes from the

Striped Snake (Zropidonotus ordinatus sirtalis). Proc.

Zool. Soc. 1912, pp. 767-770.

Nicort, W. 19126.—On two new Trematode parasites from

the Indian Cobra. Proce. Zool. Soc. 1912, pp. 851-856.

SrarrorpD, J. 1903.—'wo Distomes from Canadian Urodela.

Centralbl. f. Bakt., &c., 1 Abt. xxxiv. pp. 822-830.

West, G. 8S. 1896.—On a new species of Listomum. Journ.

Linn. Soe. Lond., Zool. xxv. pp. 322-324.

EXPLANATION OF THE PLATES,

Prate [.

Fig.1. Mediorima propria. X 20.

2. Ommatobrephus singularis. X 40.

3. Opisthivglyphe adulescens. X 7.

154 ON TREMATODE PARASITES FROM THE SOCIETY’S GARDENS,

Prats II.

.A. Opisthogenes interrogativus. X 25.

5. Styphlodora persimilis. X 40.

Puate III.

6. Lyperosomum scitulum. X 23.

7. Lyperosomum direptum. X 18.

8. Brachycelium obesum. X 100.

Prats LV.

Fig.9. Hchinostomum aliud. X 20.

9a. i - Cephalic spine. XX 250.

10. Harmotrema infecundum. X 60.

11. Hemistomum canaliculatum. ‘Transverse section through

chnging-plug. X 65.

Figures 1-10 are drawn from the ventral surface.

C.B. Cirrus-pouch. K.Sé. Ovary.

Hx. Excretory vesicle. T., T), Tz. Testes.

ON THE SKULL OF A PARIASAURIAN REPTILE. 155

10. On the Skull of a Pariasaurian Reptile, and on the

Relationship of that Type. By D. M.S. Watson, M.Sce.,

F.Z.8., Lecturer on Vertebrate Paleontology in Uni-

versity College, London.

[Received December 20, 1913: Read March 3, 1914]

(Text-figures 1-7.)

INDEX. pave

Sfatementiob matenialaused meee meat eee eaten econ ates LOO

Maseniotiom wir wie SHewIl bo: ase ssdoanoe s00005000000000900008 Gusnic| 298000 156

IBTAIZCASCD A OPER aca eee En ae Sane COR Lee oO O

TAL ahem eee Se eee rn) ha OAIREE aA So eathc nt peste

TRG(0) Basar ba enon bduaen hoa HeLnbb Budeaoeee coaane acommaone ncsnen creado 162

Relations of bones of cranial roof ...............s0000022-02 168

“Move STrwulll OF AUPHAORIC, 265 cos'be5008000 90 205 494 100 b690000e doe p9FOR0 CON 165

Dsorssion OF Hoa Guill ..sccccasccecss bao snanvacnavooqeovademooasasaced IGG

The Sphenethmoid in early Tetrapods .....................+.. 166

Nave IS apnemracoerall "ab ead debdoud bends nanadansokae seaeeticus eoubseaeoaa 10%)

Systematic position and Relationship of Pariasaurus ......... 170

Diagnostic characters of the Cotylosauria .................... --. 170

* Comparison of Pariasaurus with other Cotylosaurs ............ 172

General characters of Deimocephalia ...............00208 sce eee eee ees 175

iss Fri ANTOTAOMOIMHES caecatensccoocd0n coopeocesaceene LG

3 45 “ Therocephalia”’ and “Cynodontia”’... 176

3 ex IDROTIARAMIND, scoocavcascgoave0 adasonasoedo “6c 177

Diagnostic characters of the Therapsida .................--:.220-+ 178

Comparison with the Pelycosauria.........22..:.2:0ss:see sere reese 178

Comparison of Pariasaurus with the Therapsids ............... 179

AGIAO WTOC RSTNETILID copccsocs asa econo ado osoobeonsosocboucosobscbccasoonen IED

In 1838 Andew Geddes Bain discovered, on the Blinkwater

Commonage, in Cape Colony, the skull and a good deal of the

skeleton of a large reptile, which was subsequently described by

Owen as Pariasaurus serridens. Our knowledge of the structure

of the animal remained very slight until H. G. Seeley, in 1888,

deseribed a skull and axial skeleton referred to Pariasaurus

bombidens *. Some years later he obtained in South Africa a fine

skeleton, and the imperfect skull and axial skeleton of another, of

which he gave a good description.

In 1893 E.'T. Newton gave an excellent account of the skull of

Elginia mirabilis, a closely allied reptile from the Upper Permian

Culties Hillock Sandstone of Elgin.

A.S. Woodward, in 1898, published a diagram of the palate of

Pariasaurus, which he correctly interpreted. Broom, in several

papers, has added to our knowledge of Pariasawrus and allied

types; and J. Versluys has corrected his account of the palate.

_. * Tt is almost certain that P. bombidens is not congeneric with P. serridens. As

this paper is purely morphological I have postponed all discussion of nomenclatural

“difficulties to a more fitting occasion.

156 MR. D. M. S. WATSON ON THE

Despite this large amount of work, practically nothing is known

of the detailed structure of the skull, and I was therefore very

pleased to find that a fossil I collected on the farm Hottentots

Rivier, Gouph, Beaufort West, Cape Colony, would enable me to

give a nearly complete description of the skull. With this

description Pariasaurus becomes, on the whole, the best known

Permian vertebrate.

The material I have used consists of :—

J. A skull from Hottentots Rivier. This apparently belongs

to the same species as the skeleton in the South African Museum

described by Broom as P. serridens, a species to which it certainly

does not belong. It differs from P. baini in many features of the

skull and skeleton, and is probably quite worthy of generic rank.

This specimen consists of the entire brain-case, which has been

very completely cleaned inside and out, the posterior part being

divided by a sagittal cut, and the anterior part sliced through

horizontally. The whole of the right side of the skull-roof is very

perfectly preserved, and the fragment extends so far across the

middle line as to give the shape of the skull and the complete

structure with certainty. The sutures are shown only on the

inner surface. ‘The palate is missing.

IT. No. 49426, British Museum (Natural History), is the skull

of the specimen collected by T. Bain at Palmiet Fontein, Dist.

Beaufort West, described by Seeley as P. bombidens. His

lithographic figures give a good idea of the specimen, which,

however, has been very much more developed, the palate having

been completely freed from the stone on both palatal and dor sal

surfaces and the lower jaw disarticulated.

Many, but not all, Prof. Seeley’s sutures are correctly deter-

mined, and it is certain that the structure is essentially similar to

that of specimen I.

III. R. 1970 is the Tamboer Fontein specimen of P. bombidens

collected by Prof. Seeley ; it shows the entire structure of the

palate well-preserved and perfectly prepared, the sutures being

visible on the dorsal surface.

Description oF SKULL, drawn mainly from the Hottentots

Ltivier specimen; others are indicated when used.

The bones of the brain-case are fused into a single mass of

bone in which sutures are not distinguishable. ‘The various

regions are, however, readily identified, and are used in the

following account.

Basioccipital. —The basioccipital 1 is a large bone whose posterior

end forms the condyle. This is almost exactly circular, but its

border is slightly depressed below the foramen magnum. ‘The

outer part of the condyle is rounded, but the centre is depressed

into a deep conical notochordal pit exactly similar to that which

occurs in every vertebral centrum of the animal. ‘The condyle is

separated off from the body of the bone by a neck formed by a

SKULL OF A PARIASAURIAN REPTILE. WH

groove running round the bone, which particularly impresses the

lower surface. In section a disturbance of the cancellar tissue

suggests the point of separation of the basisphenoid. ‘The upper

surface of the basioccipital supports the very massive exoccipital,

in front of which its upper surface is excavated on each side of

the middle line by a deep pit, continued outwards by a groove

which forms the lower half of the enormous foramen jugulare.

The lower part of the basioccipital is almost entirely concealed

by the overlapping basisphenoid, but it is produced in low yet

massive tubera basisphenoidales.

Text-figure 1.

The cranium of “ Pariasaurus,” the Hottentots Rivier specimen, viewed

from the side.(X 4%).

Boc., Basioccipital ; B.Sp., Basisphenoid; Car., Canal for internal carotid artery;

Fen.Ov., Fenestra ovalis ; Parv.Sp., Parasphenoid; Pin., Pineal foramen ;

Pit.Foss., Pituitary fossa; Sph.Hth., Sphenethmoid. The figures -XIT

refer to the exits of the cranial nerves.

Basisphenoid.--The basisphenoid covers the lower surface of

the anterior part of the basioccipital, sending back processes over

those of the basioccipital to form the tubera. They are continued

forward as strong rounded ridges on each side of the bone which

pass into the great basipterygoid processes, very well shown on

R. 1870. These project downwards and outwards, leaving

between them a deep channel on the under surface of the bone.

They are of irregular shape, and the pterygoids articulate with a

158 " MR. D. M. §. WATSON ON THE

very large area of the outer surface. In advance of them a

narrow rostrum, which is probably to some extent parasphenoid,

projects forward and articulates with a special facet on the

dorsal surface of the pterygoids so as to divide completely the

interpterygoid vacuity.

Although the lower surface of the basisphenoid is very long its

upper surface is short, owing to the great anterior production of

the basioccipital. The lateral border of the upper surface and a

good deal of the side of this bone are covered by the prootic, In

advance of this region the bone suddenly narrows, its sides being

in contact with the sphenethmoid. The front of the bone ter-

minates in a smoothly rounded edge, which is the back of the

opening to the pituitary fossa. This runs backwards with sides

widely open above the basipterygoid processes; its base is closed by

the upper surface of the rostrum, and in front it passes into the

cavity of the sphenethmoid. The internal carotids enter the

pituitary fossa by a single foramen, which leads into a canal

passing down in the body of the basisphenoid for about a centi-

metre and then splitting into two, which pass out on the sides of

the bone above the basipterygoid processes. In front of the

pituitary fossa the rostrum 1s in contact with the under surface

of the sphenethmoid.

Otic bones.—The side walls and roof of the posterior part of the

brain-case are formed by the fused otic bones and exoccipitals.

The exoccipital part of the mass forms a massive pillar pierced

near its base by a single small foramen for the XIIth nerve; its

anterior border forms the back of the huge jugular foramen,

which is about 1°5 em. in diameter; it must form part of the very

powerful paroccipital process, and it is not improbable that the

two exoccipitals met above the foramen magnum.

The front border of the jugular foramen must be formed by

the opisthotic, which is so fused with the prootic and supra-

occipital that it is hopeless to separate them ; the three bones

with the exoccipital form a very massive paroccipital process,

whose outer end is firmly united with the tabular and squamosal

by an irregular suture seen only in fractures.

The foramen jugulare issues below and behind the process, and

on its lower front surface, not very far out, is the fenestra ovale,

a large hole of irregular shape.

In section, on the left side, the horizontal semicircular canal

and part of the vestibule are seen filled with matrix.

The foramen through which the VIIIth nerve gains entrance to

the ear lies far out on the front wall of the bony canal, which

forms the foramen jugulare. The foramen is a small one, the

vestibule being separated from the brain-cavity by bone and not

merely a membrane, as in most early reptiles.

The aqueductus fallopii for the VIIth nerve has an exit on the

outer surface of the prootic in advance of and slightly above the

fenestra ovale; its opening to the brain-cavity is not quite

certain, 6

SKULL OF A PARIASAURIAN REPTILE, 159

The front border of the prootic forms a smooth curve with a

notch at the bottom for the exit of the Vth and VIth nerves.

The supraoceipital part of the brain-case is produced upward

as a very massive process of roughly quadrangular section

channelled and grooved vertically and fused with the postparietals

over a large area. This separates the large posttemporal fossie.

Text-figure 2.

B.S.

The cranium of “ Pariasaurus,” the Hottentots Rivier specimen, in

sagittal section, X 4.

Reference-letters as before.

The Sphenethmoid.—The anterior part of the brain is surrounded

by a single bone, which I can only compare with the sphenethmoid

of a frog.

Posteriorly, on either side of the pineal foramen, this bone

forms a flat thin plate which thickens and turns inwards as it

passes downwards until it unites with the side of the basisphenoid

above the pituitary fossa. In this region the brain-cavity is

roofed solely by the parietals, but farther forward the cavity of

the sphenethmoid contracts very much, and the bone becomes

quite continuous over the brain: in this region the bone is

massive although of very loose texture. Farther forward the

cavity suddenly widens.

The thick floor is directly continuous with the parasphenoidal

rostrum, from which it is not visibly separated by suture (49426).

160 MR. D. M. S. WATSON ON THE

The bone is apparently all one, there being no visible sutures,

and it is very clearly distinct from the membrane bones of the

skull roof.

Viewed from within its floor is pierced by two pairs of foramina :

the posterior are large rounded openings, for the IInd and IIIrd

nerves; the anterior are much smailer foramina lying somewhat

outside the others, and possibly transmitted the IVth cranial

nerves. These two foramina have a common opening on the

outside.

The olfactory nerves passed out by the widely open front of

the bone.

Text-figure 3.

The palate of Pariasaurus bombidens, X }.

Viewed from above, the top of the skull being cut off about 5 em. above its

dorsal surface.

Reference-letters as before, with :—Mz., Maxilla; Pal., Palatine; P.V., Prevomer ;

P.T'., Pterygoid; Trans., Transpalatine. :

The Palate.—The general form of the palate has long been

known, and the distribution of the teeth with which it is armed

was fully described by Seeley (1892).

R. 1870, which shows the sutures between the bones on the

dorsal surface, 49426, which illustrates very well its general form

and relation, and another specimen, collected by myself at

Hottentots Rivier, enable me to complete our knowledge.

Pterygoid.—The pterygoid is a large triradiate bone. It

articulates by an immovable and very powerful junction with the

basipterygoid process of the basisphenoid.

SKULL OF A PARIASAURIAN REPTILE. 16]

The anterior ramus runs forward at a lower level than the

basipterygoid process, in contact with its fellow, nearly to the

extreme front of the skull, where the bone ends in an overlap

on the palatal process of the premaxilla. The two taken

together form a powerful ridge along the dorsal surface cf the

palate, which is particularly high j in the region of the prevomers,

and further back is thickened to articulate “with the anterior end

of the parasphenoidal rostrum.

The front part of the pterygoid is underlain by the prevomer,

as is well seen in a section of the second Hottentots Rivier

specimen.

The lateral border’ of the anterior ramus is in contact with the

palatine, and the external ramus supports the ectopterygoid and

with it forms the small pterygoid flange which faces the inner

side of the lower jaw.

The posterior ramus runs outwards and backwards from the

basipterygoid process ; it forms a deep plate very thiek and solid

below, nearly the whole of whose anterior face is covered by the

quadrate. This flange reaches nearly to the roof of the skull,

and at its upper edge apparently touches, in 49426, a special

process of the squamosal which will be described later.

EL pipterygoid.—No. 49426 on the right side shows appearances

suggesting the presence of a small epipterygoid resting on the

upper surface of the pterygoid. I have seen a similar suggestion

in another specimen, but am not prepared definitely to affirm the

presence of the bone.

Prevomer.—The prevomer is a large thick bone which meets its

fellow in the middle line, whose dorsal surface is partly covered

by the pterygoid and whose lower surface supports two irregular

rows of teeth: these are quite large, recurved, and sharply pointed;

in section they are seen to be set in distinct sockets.

Palatine.—The extension of the palatine on the buccal surface

of the palate cannot be determined. An interesting feature of

its dorsal surface is the presence of a process which is directed

upwards and articulates with a corresponding descending process

of the prefrontal. The outer edge of the palatine articulates

wholly with the maxilla.

Lctopterygoid.—An ectopterygoid is undoubtedly present. It

unites with the external ramus of the pterygoid and passes out-

wards behind the suborbital vacuity to articulate with the maxilla,

lachrymal, and jugal.

A very curious feature of the palate is the presence of a large

round opening between the palatal processes of the premaxille,

which probably housed an intermaxillary gland.

Quadrate. —The quadrate is a large bone standing nearly

vertical in the skull. Its lower border forms the condyle for the

lower jaw, which is placed nearly transversely.

The bone rises above this as a plate whose outer border is

notched at one place by the very small quadrate foramen, but is

Proc. Zoou. Soc.—1914, No. XJ. ll

162 MR. D. M. S. WATSON ON THE

otherwise in contact with the quadratojue

a al and s

throughout its whole height. : oe es

The body of the bone passes gradually into a powerful pterygoid

ramus, which covers nearly the whole of the posterior ramus of

the pterygoid. 7

The upper border of the bone is covered by a special process of

the squamosal.

Text-figure 4.

The skull of “ Pariasaurus,’ Hottentots Rivier specimen, X %. Dorsal aspect.

The sutures have only been seen on the inner surface and are drawn here on the

assumption (apparently justified by the specimen) that they pass through

at right angles to the surface. The part to the left of the thick dotted line

is not preserved in the specimen.

Fy., Frontal; Ju., Jugal; Lac., Lachrymal; MMyv., Maxilla; Nae., Nasal; P.Par.,

Postparietal ; Po., Postorbital: P¢.Fr., Posttrontal; P7.Fr., Prefrontal ;

Par., Parietal; Qu., Quadrate; Qu.J., Quadratojugal; Sqg., Squamosal ;

Tab., Vabulare.

Roof of the Skull—The general structure of the roof of the

skull will be more readily understood from text-figs. 4 and 5 than

from any description. It must, however, be remembered that all

the sutures shown in these figures have been seen only on the

inside of the skull, their positions on the outside being drawn on

the assumption that the sutures pass straight through at right

angles to the surface; observations on fractures show that this is

really the case. It is impossible to give an intelligible figure of

SKULL OF A PARIASAURIAN REPTILE. 163

the inside of the skull, and no errors of any morphological

importance can possibly be introduced by the method of figuring

I have employed here.

Text-figure 5.

i PLFr

The skull of “ Pariasawrus,’ Hottentots Rivier specimen, X 3. Side view.

Reference-letters as pefore.

The most striking features of the skull pattern are :-—

1. The very large size of the parietals, frontals, and nasals.

2. The very reduced postfrontal wedged in between the

frontal, prefrontal, parietal, and postorbital.

3. The very great downward extension of the prefrontal, so

that it nearly reaches the jugal.

4, The fact that, as is usually the case in primitive reptiles,

the lachrymal enters into both the nostril and the orbit.

5. The shallowness of the maxilla.

6. The fact that the quadratojugal meets the maxilla.

7. The large size of the postparietal and tabulare, which are

placed immediately behind and in the same plane as the parietal

and postorbital.

8. The reduction of the bones in the temporal region to one

on each side.

Description of the Attachment of Bones of the Cranial Roof

to Underlying Bones.

Postparietal.—The two postparietals fuse in the middle line,

no suture being visible. Their posterior edge projects freely at

the back of the skull, and the top of the supraoccipital is

tightly fused on to their lower surface near the front.

Parietal.—The parietals completely surround the pineal fora-

men, which is of moderate size. Near their posterior border they

are in contact with the supraoccipital, and further forward with

the sphenethmoid.

11*

164 MR. D. M. S. WATSON ON THE

frontal.—The frontals are in contact with the anterior part of

the sphenethmoid.

Tabulare.—The tabulare is a large bone which has a large and

very firm attachment to the‘end of the paroccipital process.

Squamosal.—Vhe squamosal is a large bone on the side of the

skull which has an articulation with the outer edge of the

quadrate, and in addition sends a special delicate process back

over the top of that bone. In the Hottentots Rivier specimen

these two bones are not quite in contact, but in 49426 they are

for a long distance, and it is very probable that the end of the

squamosal process is in contact with the pterygoid.

The squamosal is produced into a plate on the outer surface

of the skull behind the quadrate, and on the border of this, just

below the paroccipital process, is a small smooth groove, the last

remnant of the otic notch.

Text-figure 6.

The skull of “ Pariasaurus,” Hottentots Rivier specimen, restored,

from behind.

Reference-letters as before, with Ot.n., otic notch.

Quadratojugal.—The quadratojugal is avery large bone which

has an articulation with the outer side of the quadrate, and helps

to form the minute foramen quadrati. It is, however, largely

produced behind and below this bone.

Maailla.—The maxilla is a shallow bone, articulating along the

whole of its upper edge with the lachrymal and posteriorly with

the jugal and quadratojugal. It is almost certain that it really

overlaps the lachrymal so as to be rather higher than it is repre-

sented in text-fig. 5.

It bears numerous teeth, which are inserted in sockets and

very firmly held by the development of bone round their roots.

On the inner side the maxilla articulates with the ectopterygoid

and the palatine.

Prefrontal—The large prefrontal sends a process downwards

SKULL OF A PARIASAURIAN REPTILE. 165

within the lachrymal just in front of the-orbit, which reaches

and articulates with the corresponding process on the upper

surface of the palatine.

Septomawillare.— Despite the fact that this bone has been

described by Seeley and Broom, I can find no certain evidence of

its presence, and believe that if it existed, as it probably did, it

was only loosely placed in the nostril as in Sphenodon and most

Stegocephalia.

Stapes.—A. small fragment of bone lying in contact with the

fenestra ovale on the right side of the Hottentots Rivier specimen,

is probably the proximal end of a stapes; it is very imperfectly

preserved, but seems to be a plain plug about 10 mm. across

proximally, rapidly narrowing to 6 mm.

The only other types of Pariasaurian skull that are at my dis-

posal are: Ist, the type skull of Owen’s Anthodon serraius, and

2nd, the type skull of Hlginia mirabilis.

Text-figure 7.

Pin _-7

vo. 4

2 ae — ae FARNSSS

. PUREE) \| /

r Pr we )

The type-skull of Anthodon serraius Owen, X 3.

The side of the face, showing the sutures.

This sketch should be compared with Owen’s excellent lithographic drawing.

The skull of Anthodon is very badly preserved, being crushed,

weathered, and all the surface removed from what bone remains;

none the less it shows the sutures between the bones of the side

of the skull quite clearly, they being indicated by the direction

of the radiating fibres of the bones.

The structure will be most clearly understood from text-fig. 7.

166 MR. D. M. S. WATSON ON TIIE

The general plan of the structure is the same as in the larger

and older South African Pariasaurians, but the more remarkable

specializations of the latter types are absent; for example, the

quadratojugal, though it does extend forwards below the jugal,

does not reach the maxilla, and the lachrymal has a large expo-

sure in the orbital margins. The quadrate is present in the

specimen at the extreme hinder end of the part preserved.

General Discussion of the Skull.

Basis Cranii.—The occipital condyle is rather unusual in

the great development of the pit for the anterior end of the

notochord, which renders it on the whole concave; a similar

condition occurs in Diadectes and Limnoscelis amongst early

reptiles, and in a still more marked form amongst the primitive

amphibia Loxomma, Pteroplax, etc. The condition is a primitive

one.

The basisphenoid, except for its very great mass, agrees in its

structure with that of most Cotylosaurs, having very powerful

tubera and distinct basipterygoid processes.

The parasphenoid, which is as usual indistinguishably fused

with the basisphenoid, is of great length and touches the ptery-

goid in front as in Labidosaurus, Seymouria, and many other

Cotylosaurians.

Brain-case.—The more remarkable features of the posterior

part of the brain-cavity depend very largely on the fact that

Pariasaurus is a large and very massively built animal. The great

development of the supraoccipital region and its powerful fusion

with the roof of the skull, so different from the conditions in

such types as Procolophon and Labidosaurus, are probably pro-

duced in response to a mechanical necessity. The long, low, and

wide form of the brain-cavity is a very remarkable feature which

is not, so far as I know, paralleled by any other reptile.

The determination of the nerve-exits presents no difficulty,

and the only unusual feature, the entrance of the VIIIth nerve

into the ear on the anterior wall of the bony canal which forms

the foramen jugulare, is, I believe, very largely dependent on the

enormous size of the latter.

The ear, so far as can be seen, is of a very ordinary descrip-

tion, lying well up in the side wall of the brain-case.

It is apparently certain that the VIth nerve did not pierce the

bas'sphenoid but must have issued through the prootic fissure.

Sphenethmoid.—The sphenethmoid was the name given by

W. K. Parker tothe small ring-shaped bone which surrounds the

anterior part of the brain in the frog (Cuvier’s os en ceinture).

In the frog it is a ring-shaped bone, ossified in the cartilage of

the anterior part of the brain-case, and lying between the mem-

brane-bones of the roof of the skull and those of the palate,

particularly the parasphenoid, which it directly overlies. The

whole bone in this animal lies in advance of the points of exit

SKULL OF A PARIASAURIAN REPTILE. 167

of the optic nerves, but the olfactory nerves pass out in front,

of it.

A bone exactly corresponding to this in all its relations occurs

in many Stegocephalia ; it has been figured as a‘ Rhinencephalic

chamber” by Williston in Cacops and Aspidosaurus novomexicanus,

the latter specimen giving evidence from the fact that it is dis-

placed, that it is not a downgrowth of the bones of the skull

roof, By Fraas it has been figured but not determined or de-

seribed in COyclotosaurus posthumus, and it also occurs in some

specimens of “‘ Lothriceps” hualeyi.*

It is, however, much best shown in a skull in the Pretoria

Museum found at Senekal, O. F.8., which is one of the speci-

mens described by v. Hoepen as Myriodon senekalensis. In that

specimen the roofing bones have been split away leaving the

impression of their lower surface on the matrix, through an

extremely thin and transparent film of which the upper surface

of the sphenethmoid is clearly seen to be quite continuous over

the brain. None of these specimens shows the distinction of the

bone from the parasphenoid of which it might conceivably be an

outgrowth ; but a large and well-preserved skull apparently of a

form very near to Capitosaurus, which I found on the farm

Watford, Dist. Albert, Cape Colony, in the Cynognathus beds,

shows the bone clearly, and it is extremely spongy, quite different

from the hard membrane-bone of the parasphenoid.

No. 36358 in the British Museum is a fragment of a skull of

Capitosaurus nasutus, Meyer, from Bernberg. It shows the right

side of the face with the orbit, and on the back of the specimen

part of the vertical plate of pterygoid which passes backward

to the quadrate. In advance of this lies the sphenethmoid, only

the right wall of which’ is preserved, and that with its inner

surface destroyed so that the loose cancellar tissue is visible.

The posterior end of the bone has a notch which présumably

transmitted the optic nerve, and the anterior end is also notched.

This bone is extremely clearly shown to rest in the deeply grooved

upper surface of the parasphenoid.

This specimen affords conclusive evidence that the ‘ rhinen-

cephalic chamber” is a separate bone, for there can be no

doubt of the interpretation of the present fragment, as the bone

from its structure is obviously not the prootic, and also lies far

in advance of the actual position of that bone in all known

Stegocephalia. It is therefore certain that a bone surrounding the

anterior part of the brain and separating the exits of the olfactory

and optie nerves, which lies freely between the parasphenoid and

the membrane-bones of the roof of the skull, occurs in many

Temnospondylous and Stereospondylous Stegocephalia, and as it

agrees exactly in all features with the sphenethmoid of the frog

should be called by that name.

The bone which has been described above as surrounding the

* Since this was written Broom has described the sphenethmoid in Hyvyops.

168 MR. D. M. 8. WATSON ON THE

anterior part of the brain in Pariasaurus differs from the

sphenethmoid only in that its lateral walls ave produced backwards

so as to surround the exit of the optic nerve and reach the basi-

sphenoid. In my opinion such a change, depending in the end

solely on the degree to which ossification has proceeded, is not an

important one, and we are thus justified in calling the bone in

Pariasaurus also a sphenethmoid.

_ In the large Coal-measure amphibia such as Pteroplax, which

I hold to be (in the wide sense) ancestral to both the Rachitomous

and Stereospondylous Stegocephalia and the Cotylosauria, there

is a great mass of bone, sheathing the front of the brain and

passing forward as a septum nearly to the front of the head.

From such a bone the sphenethmoid of Pariasaurus, like that of

the later Stegocephalia, is easily derived by reduction. If this

view be true, it will follow that W. K. Parker was essentially

justified in identifying the pair of bones in the brain-case of

Urodeles usually called ‘ orbito-sphenoids” with the frog’s

sphenethmoid, for they also can be readily derived from the

Pieroplax ethmoidal complex.

The bone is also interesting from the light it throws on the

ethmoid of Therapsids. This bone has been carefully and excel-

Jently described by Prof. and Miss Sollas in Dieynodon and I

know it well in Hndothiodon, where it has an essentially similar

structure. In this type it consists of a short, thick median

septum which rests on the dorsal surface of the “ vomer,” which in

Anomodonts is certainly composed of a pair of fused prevomers.

This septum at the top and the back is split into two branches

which form a covering to the olfactory nerves, which issue-at the

sides of the bone at about half its length, and are in front

separated from one another by the septum reaching up to the

roof of the skull. This bone is even more like the sphenethmoid

of the frog than is that of Pariasawrus. There can be no doubt

that it is homologous with the bone I described and called

ethmoid in the skull of a Gorgonopsid.

Nor can there be any doubt in my opinion that the ‘ meseth-

moid” described in Diademodon by Dr. Broom and myself

veally represents the lower septal part of this bone, for it lies

on the dorsal surface of the palate, above the vomer in some

species and altogether in advance of it in the region of the

palatines in others.

There can, I think, from a study of some models of a foetal

skull of Perameles which Mr. R. W. Palmer was good enough

to show me, be no doubt that this bone is correctly imterpreted

as the mammalian mesethmoid.

If this series of comparisons. be justified, we shall have shown

that the mesethmoid of a mammal, the ethmoid of Anomodonts,

Gorgonopsids, and Cynognathids, the sphenethmoid of Batrachia,

and the orbitosphenoids of Urodeles are all allied bones, and

that they have all been derived from a condition resembling that

SKULL OF A PARIASAURIAN REPTILE, 169

found in Pariasaurus, and in a more primitive form in Ptero-

plax and other Embolomerous Stegocephalia.

It may at some future time be possible to bring this bone

into relation with the so-called alisphenoids of Crocodiles, which,

as is already generally recognized, have nothing to do with the

mammalian alisphenoid, but in the present state of our knowledge

of the development of the crocodile skull it is unwise to institute

such a comparison.

With regard to the identification of the nerve-exits little can

be said. The large foramen is undoubtedly for the optic nerve,

and the small foramen above it can only be for the trochlearius,

its peculiar position being somewhat paralleled by a metamor-

phosing skull of Kana temporaria figured by Gaupp, fig. 372,

Band iii. Hefte 2, of Hertwig’s ‘ Handbuch der Entwicklungs-

lehre.’

Palate-—The palate of Pariasaurus only differs from the

primitive reptilian condition in the fact that the pterygoids

extend forwards over the prevomers to reach the premaxille.

I believe this;unusual character to be an adaptive one. The

whole palate of Pariasaurus is covered with a unique armature

of small, sharply pointed teeth. The whole structure is such

as to give very great strength to the roof of the mouth. The

bone is thick, the middle line is raised into a ridge which forms

a girder along the dorsal surface; in the prevomerine region

this may be six centimetres deep; posteriorly the pterygoids are

supported by the anterior end of the parasphenoid which, with

the massive sphenethmoid above it, form a rigid connection

between the palate and the roof of the skull. Finally, the palatine

is supported about the middle of its area by the descending

process of the prefrontal. The whole forms an assemblage of

supports scarcely paralleled in any other type.

Squamosal.—The presence of only one bone in the temporal

region makes it necessary to discuss which of the three bones of

a primitive reptile has survived in Pariasaurus.

The relations of the quadrate in all Embolomerous, Rachi-

tomous, and Stereospondylous Stegocephalia are in essentials

identical, although they have seldom been accurately described,

probably because they are usually best shown in broken and

unpromising looking fragments. In all types of which I have

been able to examine satisfactory material [Pteroplux, “ Lox-

omma,” ‘* Bothriceps,” Micropholis, Capitosaurus, Trematosaurus,

Batrachosuchus, Aphaneramma, and others] the quadrate is a bone

consisting essentially of a thick lower margin, provided with

condyles for the articulation of the lower jaw, from which runs

a thin plate which inclines more or less forwards. The posterior

surface of this plate is covered to a greater or less degree by the

posterior ramus of the pterygoid, which in later types is con-

nected with a thin vertical wall rising nearly to the roof of the

skull, and by the posterior edge of the outer and lower of the

170 MR. D. M. S. WATSON ON THE

temporal bones, the squamosal. In the majority of types the

pterygoul and squamosal meet behind the quadrate, often in a

long suture.

The other two bones never in my experience haye any relation

with either the quadrate or the pterygoid,

Dr. Broili’s figures and descriptions of the type skulls of

Seymouria bayloriensis show that that remarkable reptile is iden-

tical in the structure of the temporal region and the relations of

the quadrate with such an amphibian as “ Loxomma,” a con-

clusion that I have verified by a personal examination of the

Munich material. '

‘The single temporal element in Pariasaurus is articulated with

the outer edge of the quadrate and sends a process inward along

its upper border, which in No. 49426 seems definitely to meet the

posterior ramus of the pterygoid.. Their relations are utterly

different from those held by the upper two temporal bones in

Stegocephaha and Seymouria, whilst they are very easily derived

from those of the lower temporal element of these more primitive

types by the reduction of the part of the squamosal which

formerly covered the posterior surface of the quadrate.

It thus seems that we are justified in identifying the temporal

element of Puriasaurus with the lower and outermost of the three

of primitive Cotylosaurs,

This element is, I believe, the mammalian squamosal, a thesis

which will be discussed in connection with the Deinocephalian

skull.

One curious feature of the Pariasaurus squamosal, the pro-

jection of a flange of that bone behind the quadrate, is probably

to be accounted for by supposing that the bone formerly finished

at the quadrate, leaving a very large otic notch which was sub-

sequently reduced to its actual minute dimensions by the

production of the flange.

Systematic position and Relationship of Paviasaurus.

The first author who treated of the systematic position of

Pariasaurus with the use of adequate material was H. G. Seeley

in 1888. In that paper he reached the conclusion, very novel for

its time, that Puriasaurus resembled Amphibia, Reptilia, and

Mammals in different characters, and was intermediate between

them. Although much of the evidence on which this conclusion

was based has been shown to be ineorrect, the conclusion itself

remains remarkably near the truth.

All recent authors agree in placing Pariaswurus amongst the

Cotylosanria, an attribution the meaning of which will now be

discussed,

The Group Cotylosauria was founded originally on an erroneous

interpretation of the character of Diadectes, which is the typical

form of the group. As extended and used by all modern authors

SKULL OF A PARIASAURIAN REPTILE. Wyil

it includes such types as Seymouria, Diadectes, Procolophon,

Captorhinus, Pariotichus and Pantylus. ‘These reptiles are

extraordinarily different in many features, and those in which

they agree are essentially primitive characters which they share

with Temnospondylous Stegocephalia.

The group is in fact only held together by the following

character, which separates it from other reptiles :—

The skull is completely roofed in the temporal region ; every

other feature in the skeleton can be matched in some or other

early reptile.

Cotylosaurs are distinguished from the Temnospondylous

Amphibia solely by the facts, that the intercentra are reduced and

the neural arches are expanded and thickened and the zygapo-

physial articulating faces placed horizontally, and by the presence

of only two bones in the proximal row of the tarsus (I believe this

feature does occur in some small Stegocephalia which, as shown

by the fusion of the hemal arches to the centra in the caudal

region, are quite unconnected with reptilian ancestry).

It is quite certain that the Reptilia, as a whole, must have been

derived from a form with a roofed temporal region, for all known

Carboniferous amphibia have this feature, and so also have all

Paleozoic bony fish. Therefore this character, which alone

separates and holds together the Cotylosaurs, is merely a

primitive one. Whether all reptiles are derived from a cotylo-

saurian ancestor is not so certain ; it is conceivable, though I do

not regard it as at all probable, that some of them might be

derived from amphibian types which had developed temporal

vacuities. The fact that quite a number of early reptiles shew

traces of the broad neural arches and horizontally placed zyga-

pophysial facettes, which are on the whole the most characteristic

of all the structures of the post cranial skeleton of Cotylosaurs,

suggests strongly that these types at any rate, and of course their

allies, have been derived from reptiles which, if we knew them,

would be unhesitatingly called Cotylosaurs,

Seymouria stands apart from all other Cotylosaurs in the

extraordinarily Stegocephalian appearance of its skull, its

resemblance that is to the skulls of the majority of Temno-

spondyls and Stereospondyls, for it does not resemble more

closely than those of other Cotylosaurs the skulls of the smaller

Stegocephalia, the Branchiosaurs and “ Microsaurs.”

This resemblance depends on :—

1st. The shape of the skull.

2nd. The narrow otic notch placed high up so that the

abularia are fairly near to the middle line.

3rd. The fact that the quadrate slopes backwards so that its

lower end lies far behind the upper.

Ath. The upward divection of the opisthotics.

These four characters are really all connected, the presence of

any one almost implies the others.

I do not think there can be much doubt that in these features

Nye MR. D. M. S. WATSON ON THE

Seymouria has retained the structure of the most primitive

reptiles.

If we consider later large Amphibia, we find that there isa

tendency, which is repeatedly expressed, to replace the inclined

quadrate by a vertical one; this is the case for example in Butra-

chosuchus, Brachyops, Plagiosternwm, ete.

Exactly the same change takes place in Cotylosaurs. There

are two extreme ways in which an inclined quadrate can be

converted into a vertical one, either (a) the lower end is kept

fixed and the upper end swung backwards, or (0) the reverse takes

place.

The process (a) will result in a complete obliteration of the otic

notch, and the squamosal will be brought into line with the

tabular on the extreme back of the skull; this is probably the

type of change which has produced such types as Labidosaurus

and Captorhinus. In such types the tabulare, if it be present,

being firmly fixed between the squamosal and the postparietal,

does not really require any additional support, and the outer end

of the opisthotic is free to wander down to the region of the

quadrate condyle to render support to that bone.

The process (4) results in the retention of the otic nee and in

fact in its exaggeration. ‘To it we owe types like Diadectes and

Procolophon, with an enormous otic notch overhung at the top

by the squamosal and tabulare, and with the whole of the quadrate

in advance of the basioccipital condyle. In these types the

opisthotic is far removed from the quadrate and has no possible

opportunity of supporting it.

From such a type Pariaswurus was undoubtedly derived by

the subsequent obliteration of the otic notch, by the develop-

ment of secondary plates from the squamosal and quadrato-

jugal behind the quadrate. Even in these types the opisthotics

have a tendency to rotate downwards, probably to extend the

area for the attachment of neck muscles; in doing so they take

the tabulars which are attached to their outer ends with them.

Another important type of change has been pointed out by

v. Huene. This is that the postparietals and tabulares, which in

Stegocephalia and Seymowria ave bones on the upper surface of

the skull, tend in later Cotylosaurs to be reduced to thin films

placed vertically on the back of the skull. This change is a very

important one.

We may now consider the relationship of Pariasawrus to

such other Cotylosaurs as are sufficiently well known to make a

comparison of any value.

Seymouria bayloriensis differs in the following characters, which

are primitive ones :—

(1) The shape of the skull.

(2) The narrow otic notch.

(3) The inclined quadrate.

(4) The upwardly directed opisthotiec.

SKULL OF A PARIASAURIAN REPTILE. 7S

(5) The retention of three temporal bones.

(6) The primitive palate, identical in all important features

with that of the primitive embolomerous Stegocephalia.

(7) The primitive humerus with a huge entepicondyle.

[(8) The single sacral rib. From the conditions found in

carboniferous embolomerous Stegocephalia I am inclined to

doubt if this is really primitive. |

(9) The ordinary pelvis.

(10) The expanded ribs.

And in the following specialisations :—

1i) The loss of the cleithrum.

(12) The loss of the posterior coracoidal element.

Diadectes and its allies differ considerably less from Paria-

saurus than does Seymouria in most features, whilst they possess

many advanced characters in which they differ from it more than

does the latter type.

The only primitive features in which Diadectes differs from

Pariasaurus are :

1. The retention of a supratemporal.

2. The expanded ribs.

3. The large entepicondyle of the humerus.

4. The simple pelvis.

Diadectes differs in the following advanced characters :—

5. The posttemporal vacuities are closed.

6. The postparietals and tabulares are placed more on the

back of the skull than on the dorsal surface, so that they overlap

the supraoccipital.

7. The common development of hypopophysial articulations.

The two types agree in many characters, for example :—

1. The concave basioccipital condyle.

2. The fact that the vertically placed quadrate is far forward.

3. The fact that the anterior part of the brain-case is sur-

rounded by bone, which in Pariasaurus is a single sphenethmoid

whilst in Diadectes it is said to be a paired “alisphenoid,” but

there is no doubt that the conditions are essentially similar.

These characters are all primitive ones.

There is in fact no doubt whatever that Diadectes and Paria-

saurus are not in the least closely allied, but represent two lines

differing fundamentally in the evolution of the brain-case, which

in Pariasawrus is depressed and articulates with the roof of the

skull only by a supraoccipital which is a solid narrow pillar

separating large posttemporal vacuities and articulating with the

lower surface of the parietal and postparietal ; whilst in Diadectes

the brain-cavity is high and the supraoccipital is expanded into a

wide plate which is overlapped by tl:e downturned postparietals.

It is difficult to compare Pariasaurus with Limncscelis, for no

174 MR. D. M. S. WATSON ON THE

description of the occiput has yet been published, and I am not

sure that I understand rightly the figure of that region.

Apparently, however, that type resembles Dzadectes in the

closure of its posttemporal fosse by the tabulares extending

downwards to meet the whole of the opisthotic and supraoccipital

border, and the supraoccipital is spread out into a wide plate. If

this is so, the type can have no more than the most remote

relationship to Pariasaurus.

Labidosaurus and Captorhinus resemble Pariasaurus in the loss

of the temporal bones except the squamosal, and in the preservation

of the posttemporal fosse. They differ, however, completely in

the primary loss of the otic notch, the bending down of the opis-

thotics till their outer ends are near the condyles of the

quadrates, and in the placing of the postparietals vertically on

the back of the skull. They also seem to differ in the loss of the

sphenethmoid or any ossification of that character.

In the postcranial skeleton they differ in having lost the

cleithrum, and in their curious humerus, which is quite different

from that of Pariasawrus; and also in the presence of abdominal

ribs and in the retention of an unspecialised pelvis.

Too little is known of Pantylus to make a comparison of any

value. :

I postpone any comparison with Procolophon until I describe

that form.

In the foregoing comparisons I have laid great stress on the

characters of the occiput and brain-case generally. J have done

so because a study of this region in material representing nearly

all the large groups of Reptiles and large Stegocephalia has con-

vinced me that it is really one of the most important regions of

the animal from a taxonomic standpoint.

It is in direct relation with the brain, and the general trend of

paleontological thought seems to be tending to the view that the

important part of evolution takes place in the brain, other

characters following after. In addition this region houses the

ear, and is far more removed from the action of external con-

ditions than are such features as the palate and the temporal

region. It is well known that in mammals the otic region and

the base of the skull are of great importance in determining

relationships for precisely this reason.

Since the time of Seeley most authors have felt that Paria-

saurus might have some connection with Therapsid ancestry.

This view was founded mainly on the extraordinarily mammalian

appearance of the pelvis, where, although there is no pubo-ischiac

SKULL OF A PARIASAURIAN REPTILE. 175

vacuity, the thrusting back of the acetabulum till it lies entirely

behind the sacrum and the size of the latter, make the whole very

mammalian ; another feature in which this type resembles the

Therapsids is the reduction of the phalangeal formula.

It is therefore necessary to compare the skull of Pariasaurus

with that of a Therapsid; before doing so it will be convenient to

discuss what are really the essential features of that great group.

The group Therapsida was founded by Broom to include all

the South African reptiles which are admittedly closely related,

i. e. the Anomodontia, “ Cynodontia,” ‘ Therocephalia,” Deino-

cephalia, and Dromasauria. Jt will be most convenient to see

what characters are really common to all these types, then to

discuss which of these characters are common also to other great

groups of early reptiles which are admittedly not very closely

related, and which features may be regarded merely as a primitive

inheritance, and so to discover by elimination what characters are

really diagnostic of the group. I have been able to examine

satisfactory material of all the orders, that of the Dromasauria,

which I only know through the kindness of Dr. Broom, being the

least satisfactory in details of cranial structure.

The Deinocephalia are large reptiles of very massive structure.

Skull with one temporal vacuity surrounded by the squamosal

and postorbital (or by the same two bones with the parietal in

addition ?). No temporal bone except the squamosal. Temporal

region short and pineal foramen far back. The occiput composed

of a plate with very small laterally placed posttemporal fosse,

the supraoccipital overlapped by the vertically placed interparietal

and tabulares, which are entirely on the back of the skull, and

the latter of which reach down outside the posttemporal fosse to

the ends of the opisthotics. Brain-cavity very high. [Opening

to inner ear placed low down?] Stapes in contact with the

quadrate. Quadrate large, partially overlapped behind by the

squamosal ; a quadratojugal present. Palate not well known.

Septomaxillary present on the face; lachrymal not reaching

septomaxillary.

Lower jaw with flat angular, with a notch on the lower border.

No intercentra behind the atlas ; ribs double-headed throughout

the presacral part of the column ; four sacral vertebree ¢

Two coracoidal elements, the anterior not contributing to the

glenoid cavity.

Pelvis with a short vertically placed ilum; pubis and ischia

meting each other to form a plate-like pelvis.

Humerus of an expanded and twisted type, with an entepi-

condylar foramen.

[This description is founded on the British Museum material of

Tapinocephalus and closely allied genera. |

176 MR. D. M. 8S. WATSON ON THE

AnomopontIA. (Dicynodon etc.)

The Anomodonts are small to large reptiles of semi-crawling

gait and herbivorous diet. There is one temporal vacuity sur-

rounded by the postorbital and squamosal, or the same bones with

the jugal in addition. No temporal bone except the squamosal.

Temporal region short, pineal foramen not very far back, a pre-

parietal present.

Occiput composed of a plate with small laterally placed post-

temporal vacuities. Supraoccipital overlapped by the vertically

placed interparietal [and tabulares when present.| Brain-cavity

very high. Opening to inner ear placed very low down. Stapes

in contact with the quadrate. Quadrate small, almost completely

overlapped behind by the large triradiate squamosal, Quadrato-

jugal present, but almost invariably fused with the quadrate.

Palate with large pterygoids, meeting below the basisphenoid,

then separating so as to leave a large interpterygoid vacuity,

reaching forward to the prevomers. Prevomers—fused, separating

the posterior nares, a rudimentary secondary palate formed by

maxille and palatines. Transverse present or not. Parasphenoid

a long, thin vertically placed plate forming a rostrum to the

basisphenoid and extending forward over the interpterygoid

vacuity to the prevomers (? in all types). Teeth on the maxilla

only, sometimes absent. Septomaxillary always present, some-

times in the nostril, sometimes on the face; lachrymal meeting

or not meeting the septomaxillary. Lower jaw with flat angular,

with a deep notch overluung by a reflected lamina. 25-28 pre-

sacral vertebree; no intercentra behind the atlas. Ribs double-

headed in front, single-headed behind. Sacrum of four to seven

vertebre. Tail short. In shoulder-girdle—scapula with strong

acromion, two coracoidal elements, the anterior excluded from

the glenoid cavity. Clavicles and broad flat interclavicle always,

cleithrum sometimes present. Humerus short, broad, twisted,

with an entepi- and sometimes ectepicondylar foramina. Carpus

with two centralia and five distal carpals, sometimes unossified ;

formula of phalanges 2, 3, 3, 3, 3. Pelvis with a large ilium,

small pubis and ischia, with a pubo-ischiadic vacuity, sometimes

no symphysis between the two halves. Femur long and com-

paratively slender; two proximal and four distal tarsals and one

centrale. Five digits in pes, formula 2, 3. 3, 3, 3 usually, some-

times with additional phalanges in 3rd and 4th toes.

Tt will be convenient to treat the ‘Therocephalia” and

“‘ Cynodontia” together, as the two groups in a wide sense stand

in the relation of parent and child.

Small to large reptiles of carnivorous habit, semi-crawling to

thoroughly cursorial.

Skull with a single temporal vacuity (? traces of another in

Cynognathus) bounded by the postorbital and squamosal, some-

times with the parietal and jugal in addition. Only a squamosal

of the temporal elements. Temporal region short to fairly long.

SKULL OF A PARIASAURIAN REPTILE. WA 7

‘Pineal foramen far back to far forward. Occiput composed of a

plate with small laterally placed post-temporal vacuities ; supra-

occipital overlapped by the vertically placed interparietals and

tabulares, which reach down to the opisthotics. Brain-case

very high ; opening to inner ear placed very low down. Stapes

in contact with quadrate. Quadrate small to very small, almost

completely overlapped behind by the squamosal. Quadratojugal

usually absent, but sometimes represented by a rudiment. Palate

very variable, showing the gradual development of a mammalian

type from an almost typical primitive reptilian palate. (It is

completely known in very few types.)

Septomaxillary always present on the face or in the nostril.

Lower jaw with flat angular, with a deep notch overlaid by a

reflected lamina, which becomes rudimentary in later forms.

About 28 presacral vertebree, three or four sacrals. Inter-

centra in the front part of the column. Ribs double-headed in

front, single-headed behind.

Scapula with or without an acromion. Two coracoidal

elements, the anterior excluded from the glenoid cavity, clavicles

and broad flat interclavicle always present, cleithrum present or

absent.

Humerus short and expanded to long and very slender, always

twisted ; entepi- and ectepicondylar foramina always ? present.

Carpus (only few types) with two centralia and five distalia.

Five digits; formula 2, 3, 4, 5, 4, later reduced to 2, 3, 3, 3, 3.

Pelvis without a pubo-ischiadic vacuity in early types, with one

in later forms.

Dromasauria. (Galecheirus, etc.) —

Small arboreal reptiles with long slender limbs.

Skull short, with one temporal vacuity bounded by the squamosal

and postorbital and @jugal. Pineal foramen far back. Preparietal

present (?). Occiput very badly known but apparently very

similar to that of a Deinocephalian. Kar and _ brain-cavity

unknown. Quadrate unknown. Quadratojugal probably absent.

Palate unknown. Lower jaw with large dentary and apparently

flat notched angular.

Tail very long.

Scapula without acromion, two large coracoidal elements.

Clavicies and broad flat interclavicle.

Humerus long, very slender, and twisted.

Carpus with two centralia ; digital formula 2, 3, 3, 3, 3.

Pelvis with small upright ilium and large plate-like pubis and

ischium.

Femur very long and slender. Tarsus with two proximal and

four distal tarsals and one centrale. Digital formula 2, 3, 3, 3, 3.

Proc. Zoou, Soc.—1914, No. XII. 12

178 MR. D. M. 8. WATSON ON THE

Comparison of these short descriptions will show that the only

features which are of any importance * in which the five great

groups agree are :—

1. There is one lateral temporal vacuity.

‘The material suggests that in primitive types this

was bounded by the postorbital and squamosal alone,

and is hence not homologous with either of those of

Sphenodon. |

. There is only a squamosal in the temporal region.

. The occiput is plate-like, the supraoccipital being broadened

to a wide flat plate, which separates very widely the small

posttemporal vacuities.

4, There is a single interparietal formed by a fused pair of post-

parietals, which, with the tabularia which are usually

present, is placed entirely on the back of the skull and

overlaps the supraoccipital.

5. The brain-cavity is very high.

6. The opening from the brain-cavity to the ear is very low

down.

7. The stapes articulates with the quadrate.

8. The angular is flat and notched.

9. The interclavicle is always flat and wide, not T-shaped.

ts bo

. There are two coracoidal elements.

. The anterior coracoidal element does not contribute to the

glenoid cavity,

12. There are always two centralia and five distal carpals when

the carpus is well ossified.

13. There are always two proximal and four distal tarsals and

one centrale when the tarsus is well ossified.

Of these characters, which include all common to all South-

African Therapsids that are likely to be of taxonomic importance,

Nos. 9 & 10 are merely primitive features, and so in all probability

are 12 & 13.

In fact, the characteristic features of the Therapsids, which

show the real individuality of the group, are those numbered 1-8.

Numbers 3-6 are really so connected as to be essentially one

character, and with the condition of the angular and stapes

are the only features which we could hope to recognise in a

Cotylosaurian ancestor.

It will be found that all these characters, except the exclusion

of the anterior coracoidal element from the glenoid cavity, and

the occasional presence of a vestigial supratemporal in the

temporal region, occur in the various American types which have

been included in the Pelycosauria. ,

Dimetrodon, for instance, has a single temporal vacuity, its

* They, of course, agree in such characters as the presence of parietals, prefrontals,

lachrymals, ctc.

SKULL OF A PARIASAURIAN REPTILE. 179

occiput is plate-like with a wide supraoccipital separating the

posttemporal fosse. Any bones which could be postparietals or

tabulares are on the posterior surface overlapping the supra-

occipital, the brain-cavity is very high, and the ear, as shown

extremely well by the ‘ brain-cast” figured by Case, is very low

in the skull.

The angular is flat and notched; there are two coracoidal

elements, the interclavicle is flat and wide and not T-shaped, and

the carpus and tarsus are of thoroughly Therapsid type. In fact,

every character that is found in all South-African Therapsids is

also present in Dimetrodon, which must hence be included in the

same group. [It is, however, much more primitive than any

South-African type in many features. |

With Dimetrodon, Hdaphosaurus must go to the Therapsids, for

it also has the characteristic occiput and angular.

Varanosaurus, as shown by the magnificent type-specimen of

V. acutirostris in Munich, has an occiput of the same type,

although of course it is very incompletely known. Unfortunately

the angular of this type is completely unknown; the form has,

however, the two coracoidal elements, shown extraordinarily

clearly in the Munich specimen, a flat and not T-shaped inter-~

clavicle, and, as shown by Williston’s excellent description, the

feet only differ by lack of ossification of centralia. It can, I

think, also be regarded as a Therapsid.

When one comes to Ophiacodon the problem becomes much

more difficult. The whole of the post-cranial skeleton of that

type, as described by Williston and Case, seems to be essentially

identical with Varanosaurus, but the extraordinary skull with two

temporal vacuities is apparently very different. It is exceedingly

unfortunate that no description of the occiput or lower jaw is

possible, and, in my opinion, in the absence of that knowledge,

we are not justified in discussing the position of Ophiacodon

amongst primitive Reptilia.

The foregoing discussion will, I hope, have made clear what, in

my opinion, are the really important characters of the Therapsids ;

it remains only to examine Pariasaurus in the light of them.

It shares with the Therapsids the possession of two coracoidal

elements and a single squamosal bone. The occiput is not in the

least plate-like, the postparietals and tabulares are quite on the

upper surface of the skull, the brain-cavity is long and low,

the opening from the brain-cavity to the inner ear is high on the

side-wall of the brain-cavity, the angular is boat-shaped, and the

internal mandibular vacuity, which, by its excessive enlargement,

gives rise to the flat Therapsid angular, is extremely small; the

interclavicle is T-shaped and narrow; there are no centralia on

the carpus or tarsus, and the proximal tarsals are fused. _

It is thus certain that so far from being at all closely related

to the Therapsids, Pariasawrus represents an extremely different

12*

180 ON THE SKULL OF A PARIASAURIAN REPTILE.

branch of the early reptile stock, any resemblance which it bears

to them being simply due to convergence.

I wish to express my thanks to Drs. A. Smith Woodward and

C. W. Andrews for much help during my work at the Natural

History Museum, and to Herren Prof. F. Broili in Miinchen and

Prof. F. v. Huene in Tiibingen, who allowed me to work over the

fine series of Texas ‘“‘ Permian” vertebrates in the museums of

their respective Universities. Finally, I have to thank the Percy

Sladen Trustees, who assisted me to visit South Africa, and

especially G. Gordon, Esq., the owner of Hottentots Rivier, to

whose hospitality and interest I owe the specimen of Pariasawrus

which forms the basis of this paper, and Mr. R. Hall of the

British Museum, to whose skill and care as a preparator our

detailed knowledge of Pariasawrus is very largely due.

ON DEATHS IN THE GARDENS DURING 1913. 181

11. Report on the Deaths which occurred in the Zoological

Gardens during 1913, together with a list of the Blood-

Parasites found during the Year. By H. G. PLimmer,

F.R.S., F.Z.8., Pathologist to the Society.

[Received February 2, 1914: Read February 3, 1914.]

INDEX. Page

JP EAA IYO KGyAY? aac eantco ahodacue tec tndconOLa ta tee Oo ncaC ony CoeCC eC nme cel

JERSE, JEROMOVAOE, cdasancoscandonsonnoospbuagonasonveuseccoo lusts)

arasificeNemabodesinemseee cece cee ee eset eee 187

On January Ist, 1913, there were 826 mammals, 2162 birds,

and 486 reptiles in the Zoological Gardens; and during the year

446 mammals, 1356 birds, and 683 reptiles were admitted,

making a total for the year of 1272 mammals, 3518 birds, and

1169 reptiles.

During 1913, 356 mammals, 857 birds, and 467 reptiles have

died: vhat is, a percentage of 27-9 for mammals, 24°3 for birds,

and 39:4 for reptiles.

Out of the total deaths for the year, 1680 in all, 723 occurred

in animals which had not been six months in the Gardens: that

is nearly half the total number. It has been found that after

six months’ residence in the Gardens, the death-rate falls rapidly ;

so that it is assumed that by that time the new animals have got

over their journeys, or have died from any diseases they may have

brought with them, or have got quite used to their new environ-

ment. Of these 723 animals, 141 were mammals, 277 were birds,

and 305 were reptiles; and if these be deducted from their

respective totals the death-rate will appear as 16-9 per cent. for

mammals, 16:4 per cent. for birds, and 13:8 per cent. for reptiles.

The following Tables show the facts which have been ascer-

tained, in outline. ‘Table I. summarizes the actual causes of

death in the three groups specified. Under Reptiles are included

Amphibia and a few Fishes.

Tas.E I.— Analysis of the Causes of Death.

Reference

Diseases. Mammals.|! Birds. |Reptiles.) to Notes

| following.

1. Microbie or Parasitic

Diseases. :

{Nn oerROMIORTES Sooke nccnnecboodonsseec 31 104 6

IMiv COSISI Rad canscrreecserencecee, sae 8 75 1

1 PVE OHTA VINEY” caodadbenoua seseeanse ase 34 89 138

Septiczemia...............- oeeondaee i Be

PANDSCESSE ee tec u emi naeee rer seen J

Peri cardubisinenscseas-css eee seca 1

CODD

IReritoniuisw-ssess sence soonee see osen:

TOWNS ELENA Saeed ososne cadaconeotae ee

SIIGENENTINE ccs cosoes Soatedees Sovednane We tit aL

ylephlebitistynecacie wasn ee he Qe: oe a 5

- Hydatids Berean ena hcoste. 2 sal EO ay ates ‘

Worms ] see On Labi dah tai ae Reese oa. z

182 MR. H. G. PLIMMER ON DEATHS

Tasie I,—Analysis of the Causes of Death (continued).

Reference

Disease. Mammals.) Birds. | Reptiles.| to Notes

following.

2. Diseases of Respiratory

Organs.

Wracheitisieayescey phere eee ies ei

Broncho- -pneumonia . Eco cae tain | 26 aoe

Bronchitis sees. ee | 6 defen <2) OE Fea ; 6

Congestion of lungs...............| 14 Shs

Atelectasis |... igen siceieises er 1 el eee

3. Diseases of the Heart. |

PeGicandutismmeenreeeee terre eee) ee oie Ih ian 7

Degeneration of heart-muscle . 1 1 ee

4. Diseases of the Liver. | |

Elepatitishiee: see terace ee sence | BAS ee tie Sie

Fatty gees ALTON rep eesacueeneet

Cirrhosis .. es

5. Diseases of the Alimentary |

Tract.

Gastiibisie ts acne Sececpay bacen eset |

Gastriciulceration).ss-se1e-+ 2a |

Gastro-enteritis .................005

Enteritis.. Berea tes 3

Intestinal obstruction .............

Intussusception

Strangulated intestine . :

Perforation of intestine .........

Prolapse of rectum (sloughing). | ae uu

Perforation of proventriculus ...| oe 1 nh 11

aonre

143 >

Hee oo bo Go TB oo

6. Diseases of Urinary and

Generative Organs. :

INSTR ty sock Ged esopecenoanaennass 90 135 Sao | 12

Cystichkidmeysierc. ren te eeeee FAA aH 1 13

Stone #).!.s ae uae eB 1 aS oot || 14

Inflamed oviduct . Ree eee ba 6 ae

Retained placenta .........ccscs.e- 1

7. Various. |

SARCOMA Ager reer eee ener ee| |

May elitisie ase sects samccmceumareencne | cit WE ae

Injuries disecvered post - } | ” a | 1

mortem iB) RE S|

Besides those tabulated above,

44 mammals, 127 birds, 4 reptiles, were killed by order or

by companions,

3 5) Le SS aes died from malnutrition

or starvation,

8 BOL Bs) SOmate were too decomposed

for examination,

these completing the total.

In Table I. a classification is made of those diseases which

actually caused death, but in most cases the animals were suffering

from other diseases as well. Table J]. summarizes those other

IN THE GARDENS DURING 1915.

diseases from which the animals were suffering ;

be taken in conjunction with Table I.,a much more accurate

disease in the Gardens will be

estimate of the amount of

arrived at.

183

Tasie II.—Other Diseases found in the animals tabulated

Table I.

Diseases.

ieee

Tuberculosis

| Mycosis See ocn Svice Aectomen a kat

eum Oma) easel secudaiccocsdcces

JEeRICAROUIMS, -todcecodeossovssodsonoe den ese

| Peritonitis SSA SN SEA Aero are

PANS CESS oxic tne Ai PAN Cet

Empyema

Septicae paral Masks crests eceeret css

SKOWAVONDIS|. Gcokuedonsebbaocoshesnuo née ccs

UAE NEA AIS ere ee aps Ge see aa

JO ENBiey hateoeucosestaaee amen i

| Hemogregarines .....................

{OP OATNOSOTNES) .50000 056051 sn5054sa0605008

EA RYOVPEOES 7 sdauanae sacembradtes cae Sue ecoceeene

| lalsp@leiaialsines nesses condousses sacet cose cuore

SHINGO PSUS cou coadeuvatsescce seo sonseode

JUEWOCOV WOOD, cooccp cones acoacadcooae nce

I PAWCVBE NK02E eer ind coceeteaereneeccee eeoctae are

{

Mammals.) Birds. |Reptiles.

PN rFON FE Ww

Pop:

bo:

mace

Bronchitis Rend otidadecocepan

Broncho-pneumonia ..................

Congestion of lungs ..................

Cidema of lungs .......................:

Kmphysema

Hydrothorax SEHD Ree abe CHORE

PIG DIB IG) Pe Wace son cutoroconsmenceea aE

JPERNGATRGNES csoboasoobuBeasse dosonosda nes

Hattyslearty eo meee ee ee ee:

DilatedGheart, Wace. Ges eee) .qeeclees

ANBIKESKOVGT ERY Nae tough er snaedaatos seonee ete

HERTS MAGES nice cc cteesaty Meas tamutet ereenclh

Cimmhosisioteliversnees-ee eee eee

Watty iver AA hscoaes: ese n eee cate!

Gastritis Sse pesca inant conoes

Gastric ulceration .....................

Gastro-enteritis ..........0. 0.0.0.0.

Intussusception ....................008

Intestinal obstruction Been

TRITIUre Te HG We ann 2 ee Ran eee

Neo ling! Mees biker me ane helen oie Janke

Cystic kidneys

(COVEISIUTS » MESURE Esher Meee me yntiae, Cae

| Inflamed oviduct .....................

Rickets eee ee eee ee.

Mialinutaitivonly Seana: sce: adesestoee ce

PAIS GILES ee eee SE Stns Woe aa

Tnjuries

Pobre eo

pS :

PNOENWPANrRAe-

Ow:

OUR RTs Cur con:

114

sr : a 6

HONAANNOS: :

Omdwee:

bo:

bo C

OV Go

Ss moO:

Reference

to Notes

following.

and if this Table

184 MR. H. G. PLIMMER ON DEATHS

Table IIT. shows, in still further detail, the distribution of

diseases amongst the various orders of mammals.

TaBLe III.—The Distribution of Diseases causing Death

amongst the principal Orders of Mammals.

abel acts eal

| 4) & =

| Pas |< | 2 |e iee

| Diseases. ie || e S 3 es a

| = = By a eae Zi

ae = zs Be eSe sil een

Soe owas) eae

eRulberewlosisy aes ee ee ene ln TI Geat| lite 9 1 | Pe ea

| Mycosis Ae eta ts axeeeet aun ee bes bee el me es 1 1 6 |

| JEROMNOMIE, 55. 2asscascosonenansauecoooseossal| LO 6 13 2 | at |

KaAtbecons marten ann natn kena at ha ie 1 ne |

Koylepblebitisayn ctiy ye stoce sees ve etme ee = | |

MIP VieMay ences aerate cca roan al “2 1 ala 1

eT CAUCILIS eee Seva es meee eM earner eh at a Spe 1 eo

ft JELLO UNIS) oe ee es sbeener abacadoaaaaaced| | beer aon i2 1 | 1

I@bdiy. atid Sienna ch GtnUh eminence Bh Bl ae ene aan eel a een |

Sepbicaaiiairs mics: em tee cca tae seereee| yea Mt | vt

Bronchitis s Samp sarstoaec 2 3 1 Loa ere |

Broncho-pneumonia ..................... 8 6 3 Gea betel 1

Congestion of lungs 6 | 5 2 1

| Atelectasis Ds st Fh

Mattyheanti: tare e eee au: She Be an se a 1

Gastritis ere teiewte ae waste ene alian aes ae ae he eI oA

CRINTRO-GITAUINS cencoonancrascosstacdacoced| 4 || Il 3 Sh cootlligeeas |

| Intestinal obstruction .................. Neuse 2 be aes

Perforation of intestine .................. Be ase te | |

INGastnrenal cerationeee eee eee eee 2 oe 2 |

Strangulation of intestine ............ 1 |

Sloughing intestine ............0........ | 1

| LRG OUSRESEI IOV Gosconsetanaasdencoseserebell sae 2 ie ate a 1

/e JBAO RHEE CH act hll So ueeraaniee con EMMA Ane aes 9 9 7 a ¥!

Nem matte meds acne tots on csteinccshonaee | 20 31 13 We 283 10

| Stone en a aEeS See Soh ee San ee 1 |

SANGO Tiny atten. Mans clecnuen yAeeula ects il 2 |

[aM iiy Ghibis Mie re ae chew nic Se erase |

Notes on the foregoing Tables.

1. The total incidence of infectious diseases in the Gardens is

about 7 per cent. for mammals and birds and 12 per cent. for

reptiles.

2. There has been an increase in the deaths from tubercle

amongst the mammals and birds. In the former it was due to

an epidemic which broke out in the old Insect House in 1912:

15 of the 31 cases in mammals came from this House. 13 of the

31 mammals had not been in the Gardens for six months, and

4 of them had been pet animals. The most interesting case was

that of an [Ibex under one month old, which had tubercle in one

Jang and in the thymus: there was scarcely any healthy tissue

IN THE GARDENS DURING 1913. 185.

left in the latter organ. In the case of a Bear the disease was

of human type. Most of the cases in birds would appear to have

been acquired in the Gardens, as only 22 of them had been less

than six months in the Gardens. In 61 of the birds it was

acute, and was a general infection: in 1] it was of bovine type.

4 of the reptiles, which show a considerable decrease, were

tortoises.

3. All the mould-diseases have been grouped under mycosis.

Of the 8 mammals, 6 were Kangaroos and the disease was of the

same typeas that previously described, 1 was in a Gazelle, of same

type, and the other was a mycotic disease of the intestine in a

Beaver, in which the mould was of a different variety. The

number of deaths from mycosis in birds is still very high, and

constitutes 8°7 per cent. of the deaths in birds. Some young

Pheasants died from mycosis at the age of 14,17, and 20 days,

with mycotic growth in all the organs and filling the body-cavities.

4. There has been a slight decrease in the incidence of pneu-

monia in mammals and birds, and a slight increase in reptiles.

In 5 of the latter it was due to irritation caused by worm eggs ;

the rest were pneumococcal and constitute about 30 per cent. of

the number of deaths in reptiles.

5. In umbilical veins of recently born Buffalo and Gnu.

6. There has been a general decrease of these diseases of the

respiratory organs: they are largely dependent on weather.

7. These cases of pericarditis in birds were not due to infection,

but to a deposit of crystals in the pericardium associated with

chronic kidney disease.

8. The two birds were Penguins, in which over-distension with

fish was the cause.

9. In 6 of the mammals, 14 of the birds, and 3 of the reptiles

the inflammation was caused by parasites (worms and coccidia).

In 1 of the mammals and 7 of the birds the cause was a trau-

matic one (sand, hay, ete.). In 7 of the mammals and 65 of the

birds it was hemorrhagic, and probably of bacterial origin. The

remainder of the cases were apparently due to the quantity or

quality of the food not being suitable to the animal.

10. Two of these bad intussusceptions were in Genets.

11. Ina Plover from wire.

12. There has been a slight increase in the number of cases of

nephritis. Under this are grouped the acute and chronic cases,

many of the latter being the result of old age. 14 of the cases

in mammals were acute, and 32 of the cases in birds: the rest

were of varying degrees of chronicity. Many of the mammals

and birds had associated lung lesions, which would seem to

indicate that climatic conditions and exposure may be answerable

for these cases,

13. Ina Terrapin in which both kidneys were converted into

multiple cysts.

14.-In an Antelope in which a stone impacted in the urethra

had produced a ruptured bladder.

186 MR. H. G. PLIMMER ON DEATHS

15. Two of these were lympho-sarcomata of the abdominal

glands in two sheep, mother and son, from the same house. The

others were an angiosarcoma of liver in a Coypu rat, a sarcoma of

the scalp in a Cercopitheque, and an adeno-sarcoma in a Rail.

16. The diseases grouped under the term malaria were due in

12 instances to Hemoproteus danilewskyi and in 6 instances to

Plasmodium precox.

16, 17, 18, 19, 21. See the section on blood-parasites below.

20. This was a very considerable infection of the muscles of a

Langur, which was not visible to the naked eye.

22. There has been a considerable decrease in the number of

rickety mammals.

In comparing the deaths recorded in this Report with those of

the five preceding years, there are two points which seem to be

in continual prominence, and which must therefore be of practical

importance. The one is the fact that a large number of animals

—in the large sense—nearly half of the total number, have died

within six months from their admission to the Gardens, the

majority of these dying within three months. Of these, a large

number die of microbic or parasitic diseases.

The other fact is that a very large percentage of animals have

died from inflammatory conditions of the alimentary tract which

cannot be attributed to mechanical or microbic causes, and which

are apparently due to some defect in their food, the quality or

quantity or both not being suitable to the animal.

From a consideration of these facts, it would seem possible that

the point first mentioned could be dealt with practically by a

proper and effective quarantine, which would prevent the intro-

duction into the Gardens of new infections, or of those already

existing there, in a condition of increased virulence.

As regards the cases next mentioned, a careful consideration

by experts of the feeding of the animals throughout the Gardens,

would enable reasonable and physiological alterations to be made,

and probably would reduce effectively the death-rate from

this cause.

BiLoop-PARASITES.

During the year the blood of every animal which died has

been examined, with the result that parasites have been found in

138; in 60 species for the first time.

They have been distributed as follows :—

Filarie. In 5 mammals; found in 3 species for the first time.

28 birds; in 20 species for the first time.

5 reptiles ; in | species for the first time.

Trypanosomes. In 8 birds; in 5 species for the first time.

2 reptiles.

IN THE GARDENS DURING 1913.

187

( Hemoproteusdanilewskyi. In 14 birds; in 7 species

Malaria. < Plasmodium precox.

| Heemocystidium.

for the first time.

In 6 birds; in all for the

first time.

In | reptile.

Leucocytozoa. \n 4 birds; in 2 species for the first time.

Hemogregarines. In 64 reptiles; in 14 species for the first

time.

Hexamitus type.

Ente tinal organisms of } In 1 reptile for the first time.

The following Tables show the occurrence of the blood-parasites .

in detail :—

Embryo Filarie found in the blood of Mammals.

Hasirat.

Brazil.

2 Lion Marmosets (Leontocebus rosalia).

Found in the following for the first time :

Short-tailed Wallaby (Mlacropus bra- Australia.

chyurus).

Rock Wallaby (Petrogale penicillata)... Australia.

Martin’s Cercopitheque (Cercopithecus W. Africa.

martini).

Embryo Filarice found in the blood

White-throated Jay Thrush (Garrulax India.

albigularis).

Lanceolated Jay (Garrulus lanceolatus). India.

Brazilian Hangnest (Icterus jamaicai)... Brazil.

Mexican Jay (Xanthura luxuosus) ...... Mexico.

Wood Thrush (Lurdus mustelinus) ......

Found in the following for the first time:

Albert Towhee (Pipilo alberti) ............

Rose Finch (Carpodacus erythrinus) India.

Black Bullfinch (Melanopyrrha nigra) . Cuba.

Black F'rancolin (Francolinus vulgaris). India.

Mexican Rose-Finch (Carpodacus mexi- Mexico.

canus).

Black-shouldered Tanager (Calliste Brazil.

melanonota). ;

Mahali Weaver-bird (Plocepasser ma- S. Africa.

hali).

Nuthatch (Sitta cinnamomeiventris) India.

Chestnut-bellied Nuthatch (Sitta casta- India.

neiventris).

Pileated Song Sparrow (Zonotrichia S. America.

pileata).

Cuban Amazon (Chrysotis lewcocephala). Cuba.

Red-shouldered Starling (Ageleus phe-

niceus).

Sulphury Tyrant (Pifangus sulphuratus). S. America.

N. America.

TyPrE oF FILARIA.

Long.

Long, thick, very pointed.

Long, thick.

Long.

of Birds.

Short, thick, no vacuole.

Short, pointed.

Long, no capsule.

Long, pointed.

Short, pointed,

Short, thick.

Long.

Long, pointed.

Long.

Short, thick.

Short, pointed.

Long, thick.

Long, pointed.

Short.

Short, thick, pointed.

Short, pointed.

2 kinds; one very long,

the other very short

and thick.

188

Great Grey Shrike (Lanius excubitor) ...

Swainson’s Blue Jay (Aphelocoma sor-

dida).

Red-billed Hornbill (Lophoceros erythro-

rhyncus).

3 Indian Rollers (Ooracias indica) ......

2 White-headed Starlings (Poliopsar

leucocephalus).

Larger Hill-Mynah

media).

Malaccan Parrakeet (Paleornis longi-

cauda).

(Gracula inter-

MR. H. G. PLIMMER ON DEATHS

Europe. No capsule, very striated.

Mexico. Short.

Africa. Short.

Tndia. Short.

China. Long.

India. Short, thick, and pointed.

Malacca. Long.

Embryo Filarie found in the blood of Reptiles.

Edible Frog (Rana esculenta)

2 Say’s Snakes (Pituophis sayi)

N. America.

Europe. Short, thick.

Long.

Found in the following for the first time :

2 Warty Chameleons (Chameleon verru-

cosus).

Madagascar. Short, stout, with thick

capsule.

Trypanosomes found in the blood of Birds.

2 Blue-crowned Hanging Parrakeets

(Loriculus galgulus).

Little Owl (Athene noctua) .............00-+

Malay.

Europe.

Found in the following for the first time :

Goldfinch (Carduelis elegans)

Shama (Cittocinela macrura)

Tawny Owl (Syrniwm aluco) ........

Spotted-sided Finch (Sieagematannn Git

tata).

Brazilian Hangnest (Icterus jamaicai)

N. Europe.

India.

Europe.

Australia.

Brazil.

These were all of the type of Trypanosoma avium.

Trypanosomes found in the blood of Reptiles.

Tree-Frog (Hyla arborea) (blue variety).

Edible Frog (Rana esculenta)

S. Europe.

‘S. Europe.

These were of the type of Zrypanosoma rotatorium.

Intestinal Organism found in the Blood of the following Reptile

for the first time.

Rough Terrapin (Wicoria punctularia) .

S. America. Of Hexamitus type.

Hemogregarines found in the blood of Reptiles.

3 King-Snakes (Coronella getula)

3 Rat-Snakes (Zamenis mucosus) .........

Reeves’s Terrapin (Damonia reevesi)

2 Bushmasters (Lachesis mutus)

8 Cobras: (Waia tripudians) 2... 2.00.04.

3 Dark Green rae (Zamenis gemon-

ensis).

N. America. Small, short.

India. Medium sized.

China. Small, short.

Trinidad. Large, host-cells enlarged

and dehemog lobinised.

India. Long.

S. Europe. Small, host-cells enlar aath

IN THE GARDENS DURING 1915.

Diamond Python (Python spilotes) ......

Blood-stained Terrapin (Cinosternum

cruentatum).

3 Hog-nosed Snakes (Heterodon platy-

rhinos).

8 Common Boas (Boa constrictor) ......

Eyed Lizard (Lacerta ocellata) ............

5 Rattlesnakes (Crotalus atrox)

3 Testaceous Snakes (Zamenis flagelli.

formis).

3 Indian Pythons (Python molurus) ......

Anaconda (Hunectes marinus)

4 Say’s Snakes (Pituophis sayi)

Vivaceous Snake (Larbophis fallax)

Gallot’s Lizard (Lacerta galloti)

Australia.

S. America.

N. America.

S. America.

S. Europe.

N. America.

India.

S. America.

N. America.

S. Europe.

N. Africa.

Found in the following for the first time:

8 Pigmy Rattlesnakes (Sistrurus mili-

arius).

Spinose Land-Emys ( Geoemyda spinosa).

Helen’s Snake (Coluber heleni) ..

Banded Trichogaster (Tr Schrametian ines

ciatus).

Bengal Monitor (Varanus bengalensis)...

Cape Viper (Causus rhombeatus)

Four-lined Chicken Snake (Coluber obso-

letus, var. quadrivittatus).

2 Emerald Green Tree-Snakes (G‘astzo-

pyxis smaragdina).

Graham’s Snake (Zamenis grahami)

Harlequin Elaps (Hlaps fulvius) .........

Long-nosed Crocodile {Crocodilus cata-

phractus).

Leopardine Snake (Coluber leopardinus) .

Cananina Snake (Phrynonax sulphureus).

Sooty Snake (Boodon fuliginosus) .........

N. America.

Malay.

Ceylon.

India.

S. Africa.

N. America.

Sierra Leone.

N. America.

N. Nigeria.

S. Europe.

Trinidad.

W. Africa.

189

Large.

Long.

Large, host-cells enlarged.

Large,host-cells enlarged.

Long, host-cells enlarged.

Large, host-cells enlarged.

Long, host-cells enlarged

and dehemoglobinised.

Medium, cells deformed.

Long, doubled over.

Long, host-cells enlarged.

Medium.

Of Karyolysus type.

Small.

Medium.

Small.

Short and thick; of in-

terest, as they are said

not to occur in fresh-

water fishes.

Large, of ordinary type.

Of Drepanidium type;

nearly every corpuscle

infected.

Large.

Long, doubled over; host-

cells enlarged.

Large,host-cellsenlarged.

Large and doubled over.

Some short and_ thick,

others long and doubled

over.

Small and thin.

Large, host-cells enlarged.

Large, host-cells enlarged

and dehzmoglobinised.

Hemoproteus danilewskyi fownd in the blood of Birds.

2 Blue-crowned Hanging Parrakeets (Loriculus eee

Kestrel (Tinnunculus alaudarius)

nye Owl! (een LACTATE) 000 cn nonaee acca coe pcapne cea oe 2ouAABAno cer

3 Indian Rollers (Coracias indica)

Found in the following for the first time :

Malayan Peacock-Pheasant (Polyplectrum bicalcaratum)

Shama (Cittocincla pee

Siskin (Chrysomitris spinus) ..

Tawny Owl (Syrniwm aluco) ..

Lanceolated Jay (Garrulus Uonceolabuse: ai cerrone a otan ocean

Sulphury Tyrant (Pitangus Sulpanoeu ee. es Ea IN Th

Lesser Kestrel (Tinnunculus cenchris) ...... 2.0060 ccc cccses eee een vee eee

Hasirat,

Malay.

Europe.

S. Africa.

India.

Malay.

India.

N. Europe.

Europe.

India.

S. America.

S. Europe.

190 ON DEATHS IN THE GARDENS DURING 1913.

Plasmodium precox found in the blood of Birds:

Found in the following for the first time : Hasitat.

Yellow-fronted Barbet (Cyanops flavifrons) 11. .....0ccccccceecee ees Ceylon.

Mongolian Pheasant (Phasianus mongolicus) ....10....sseee cen eee Mongolia.

Goldfinch (Carduelis elegans) ........ Aten Rate eemraneeee N. Europe.

White-crested Touracou (Tuwracus cor, CE ae Pra ae eee enacuaced S. Africa.

Swainson’s Blue Jay (Aphelocoma sordida) .............sc eee eev eee Mexico.

Wood-Thrush (Turdus mustelinus) .......00ccc cee cee see eee cee cee tenes N. America.

Leucocytozoa found in the blood of birds.

Ibi Onsils (AGiGiae TOGHE@) - coc caseceaceons eas 0tHc66c00 cos paneno DAA cod S. Europe.

Found in the following for the first time :

Chat (Oreicola ferra) .. SRE arcntere me aaonse nacacunoce India.

Tawny Owl (Syrnium Hieay" JON SOREN DIRE RRS Europe.

PB, Z, 8. 1914, JONES, Pi, we

Gensta/ tubercle

Urinary

passage

R.Wolffian

Mullerian pies

ducts

Genital tubercle

Tissue of the

floor of

urinary passage

os a

= / Mullerian ducts

INTERNAL GENITALIA OF THE MOLE, ©,

Dm, 4, S, IsiZs, JONES, Il, ul.

Genital tubercle

Penile urethra

Tissue continuous

with terminations

of mullerian ducts

Ep ithelial

ingrowth from

base of genital

tubercle

Pelvic urethra

Neck of bladder

Mullerian ducts

Genital tubercle

Urinary

passage

R.Wolffian

duct

Mullerian

ducts

UN TIIRINZNL, GIs INIOEAIL IGA, Or Wane: MOLE, 9.

12 4S, 1S, JONES, Pl. M0.

Genital tubercle

Epithelial

ingrowth fy

from base of [fa

ental mi

tubercle

Strand of tissue

in which

mullerian ducts

terminate

7 Lower ends

of mullerian

ducts

Base of genital

tubercle

Epithelial ingrowth

at base o

genital tubercle

Unilateral lumen f

in epithelial

portion of

female passages

Urinary

passage

Area of

continurty of

urethral and

utero-vaginal

Cavity of

walls

utero-vaginal

canal

NIE IRIN, CHENIN IA, Ol” Wish MOLE, 9-

ON THE REPRODUCTIVE HISTORY OF THE MOLE. 191

12. Some Phases in the Reproductive History of the Female

Mole (Talpa europea). By FrepEeric Woop-Jones,

D.Sc., F.Z.8.

[Received February 3, 1914: Read February 17, 1914. ]

(Plates I.-I1I.,* and Text-figures 1-13.)

INDEX. Page

Sinuciunerorm Morphol ogyererene: eee eeeeeene ene eemeeen PLOO

Wevelopmientinc nce csse ceen cee oe ceay Los) 200

A. Introduction.

For long it has been known to country folk that in some way

the sexual life of the Mole (Valpa europea) is peculiar.

In an indefinite manner this idea has been given concrete form

in works on Natural History, and practically every field-natur-

alist who has written of the habits of the mole has stated that

the males greatly outnumber the females. Various habits of the

mole are described which are supposed to be the outcome of this

great preponderance of the males; and most popular works

allude to the love contests, the fierce battles, and the slaughter of

the surplus males during the spring mating season. It is an

observation which has been handed down almost unaltered from

the very oldest authors, that the proportion of the sexes 1s

equalised by the killing of superfluous males in the struggles of

courtship.

I have, however, heard another account of this idea of the

animal’s sexual peculiarity from mole-catchers. I have been told

by a most observant trapper that moles are all of one sex until

their second season, and that then, and only then, do they be-

come malesand females. J do not know of any printed expression

of this particular statement of a piece of nature lore, but I have

no doubt the idea is more prevalent, among a class of men who

gain a partial livelihood by mole trapping, than the mere repe-

tition of an isolated observation might lead one to suppose.

The mole is an inconspicuous animal, not readily observed

in its natural habitat nor easily kept in confinement, and so it

has not reached the dignity attained by the spotted hyzena of

having a literature of legend concerning its sexual habits which

stretches back into. the. classics; nevertheless it has taken its

place among the creatures of fabulous habit in the nature lore of

a very limited class of country folk.

As is the case with so many of the nature beliefs current up

and down the country, there is far more than mere countryside

superstition in this idea that the mole can change its sex, or at

any rate its sexual appearances. Asa matter of fact it is now

more than eighty years ago that the question was removed from

* For explanation of the Plates see pp. 215-216.

192 DR. F. WOOD-JONES ON THE

the region of nature superstitions; but this has not yet effected a

complete elimination from works on popular natural history of

the imaginative accounts given by the older authors.

The apparent enormous preponderance of the males thrust

itself upon the attention of Etienne Geoffroy (St. Hilaire), and to

him is due the credit of placing on a firm scientific basis what

was at the time a complete mystery to the scientific man and a

subject of superstition to the ignorant (1). The work of Geoffroy

was published in 1829, and, looking back, it seems a wonderful

thing that this man, who with scalpel and forceps made clear all

the essential facts, produced, in this respect, so little impression

upon the succeeding generations of naturalists that all the old

inaccuracies and traditions were repeated regularly in successive

works on natural history.

_ Jn writings on the natural history of the Mammalia, for at any

rate another seventy years, English authors were contented with

a repetition of the popular stories concerning the numerical

inequality of the sexes, and the supposed elimination of the

superfluous males in the fierce contests of the pairing season.

Briefly, Geoftroy discovered by his investigations that although

young moles all appeared to be males when the examination was

confined to the external appearances of sex, yet, on dissection,

some of these seeming males proved to be females in possession

of well-developed internal female genitalia. Such a finding was

indeed remarkable, but still the mole was by no means an isolated

example of external sexual anomaly; for other cases of the

difficulty, or even impossibility, of distinguishing the sex of

mammals by mere external examination were already well

known. But Geoffroy went further than this, for he showed

that this peculiarity was limited to some females, and that

although some appeared outwardly to be males, others, like the

normal females of most mammals, possessed a vaginal orifice.

In some females an organ almost exactly similar to the penis of

the male alone existed, and in others there was an added orifice

situated between the penis and the anus. This vaginal orifice he

recognised as being a new formation in those females which

possessed it; he assumed that this new orifice existed only after

pairing, and he supposed it was actually produced by the efforts

of the male in the act of copulation. It would be imagined that

merely to call attention to this very unusual state of affairs would

have been sufficient to attract a large army of anatomists and

zoologists to the field of investigation, and yet but little attention

seems to have been given to the subject. Indeed when, ten

years later, Thomas Bell wrote his article on Insectivora, he opens

his account of Zalpa by saying “the female organs in the mole

offer some peculiarities which deserve more attention than they

have hitherto received” (2). Bell's account is obviously based

upon the work of Geoffroy, and it is well to quote it in full. ‘In

the first place, it appears that in this animal the urinary and

genital orifices are wholly distinct. The clitoris, which is of

REPRODUCTIVE HISTORY OF THE MOLE, 193

considerable length and.very much resembles the penis of the

male, is pierced for the passage of urine, and thus constitutes a

true urinary penis, Beyond this is a transverse slit of a slightly

erescentic form which constitutes the opening of the vagina.

There are none of those duplicatures of the integument which in

other mammalia constitute the labia and nymphe, but the skin

is smooth. But one of the most curious points in the structure

of these parts is that in the virgin state this vaginal aperture does

not exist, the skin being perfectly and tightly drawn over the

entrance ; so that there are in this state but two openings, the

urethral and the intestinal. So perfectly is this the case that it

is very difficult to know a virgin female mole from the male by

mere external examination, As this covering is so tense, the

utility of the little bone at the extremity of the penis in the male

is very obvious, and its pointed and tapering form is at once

accounted for; for it is clearly intended to perforate this tense

covering to the vagina.”

After another thirty years the subject is again brought forward,

in the form of references to Bell’s article, by Owen in his ‘ Com-

parative Anatomy of the Vertebrates’ (3). But Owen’s plain

statement that “‘ the mole shows a complete closure of the vaginal

orifice in the virgin state” tended rather to suggest that nothing

more strange than an unusual development of the hymen ac-

counted for the condition. The fact that no vaginal orifice existed

was rather lost sight of in the suggestion that the outlet was

merely closed in the virgin state. By 1868, therefore, this extra-

ordinary discovery had found its place, but only as a brief

allusion, and in a rather modified form, in the standard work on

comparative anatomy ; and it still rested apparently entirely upon

the isolated investigations of Geoffroy carried out forty years

before.

So far as I can ascertain the whole question was then practically

lost sight of, or forgotten, until 1902, when Lionel E. Adams,

after a wide experience of work asa field-naturalist, read a paper,

entitled ‘‘ A Contribution to our Knowledge of the Mole,” before

the Manchester Literary and Philosophical Society. To the older

work of Geoffroy, Adams added original observations based on a

very wide first-hand knowledge of the mole; and in addition

to describing in detail the naked-eye external changes which

accompany the formation of the vagina, it was made clear that

this vaginal development was in a way spontaneous and not

due to any action of the male.

The work of Adams was published in 1903 (4); it remains the

standard account of the progress of this wonderful change, and

as such is referred to in most recent works on zoology. Quoting

from his original paper, the results are summarised as follows :—

‘“‘ My observations show that about March Ist a wrinkle appears

at the base of the clitoris which in a few days assumes a purple

hue, and by the middle of March a perforation appears in this

livid wrinkle on each side of the middle line. Towards the end

Proc. Zoo. Soc.—1914, No, XIII, 13

194 DR. F. WOOD-JONES ON THE

of March these two perforations coalesce and the vagina is then

open for penetration in the usual way. I have not found any

internal hymen whatever.”

Such a statement may be considered as a summing up of our

knowledge of a process which one would suppose would arrest the

attention of any anatomist, a process by which an animal evolves

in its post-natal stages an entirely new orifice for the genital

system, and changes an apparently male arrangement of external

genitalia into one that is obviously female.

This brief history of the subject, incomplete in its details as it

very possibly is, seems to me to be highly instructive, if only

for the light it throws upon the relation of what we are apt to

despise as mere nature lore to the more rigid demands of accurate

anatomical knowledge.

It is now more than twenty years ago that a mole-catcher told

me that all first-season moles were alike as to their external

genital organs, and that only in their breeding year could they be

distinguished as males and females. I do not imagine that the

knowledge originated with this man, but would be inclined to

regard him as merely a link in a very long chain of folk whose

knowledge was certainly not gained from books, but was culled

from real observation and backed by the weight of tradition. It

was easy to regard a mole-catcher’s statements as mere super-

stitions until the work of Professor J. P. Hill (5) on Perameles,

and some study of the developing vagina in the human embryo,

compelled a belief in the enormous plasticity and adaptability of

the female genital tract, and lent colour to the suggestion

that a vagina could be produced de novo in post-natal life. It

was only then that preliminary investigations showed that the

superstitions regarding this transformation of the mole were well

founded; and a search of the literature revealed the work of

Geoffroy and Adams.

B. Embryonic Stages of Hxternal Genitalia.

I shall begin my account of the development and transforma-

tions of the genitalia of the Mole by a brief description of the

formation of the external genitalia in the embryo. I have been

fortunate in the examination of a very large series of embryos of

all ages, and for the bulk of this material I am indebted to

Mr. R. H. Burne, of the Royal College of Surgeons, who has

placed at my disposal a large number of fcetal families collected

with the greatest care, and in a perfect state of preservation.

Other embryos I have collected for myself, or have received

from time to time from friends, and the entire series has per-

mitted the examination of the external genitalia in all phases of

embryonic development.

I have been very careful in following, and attempting to

interpret, these stages correctly, and the reason for this especial

caution demands explanation. Hitherto my actual first-hand

REPRODUCTIVE HISTORY OF THE MOLE, 195

knowledge of the process of development of the external genitalia

had been limited to the detailed study of human embryos. I had

long since satisfied myself as to the method of development of the

human external genitalia, and I had become convinced that some

of the processes which are usually described did not in fact exist.

The most cursory examination of my series of mole embryos,

however, seemed to show that what I had regarded as, and stated

to be (6), inaccuracies in the account of human development, were

actual and obvious phases in the mole. Such a result necessi-

tated a most careful review of my material, for it seemed that

what was so obvious in the mole embryo was sufficient to stultify

my assertions regarding human development. ‘To assure myself

of the correctness of my interpretation of the phases in the mole

was therefore essential, and I have accordingly submitted all my

material to a very critical examination. The results given here

may therefore be taken as those which are so evidently true

that, after repeated examination, they had to be accepted—though

unwillingly—as correct.

The reinvestigation of the phases in human development was

the next step, and the results of this part of the research I reserve

for a further paper.

In the mole embryo measuring 9 mm. along the dorsal surface

from the vertex to the caudal bend, the rudiments of all the

essential elements of the external genitalia are readily recognisable.

Such a stage is shown in text-fig. 1(A & B). The cloaca is present

as a wide orifice, from the ventral margin of which the genital

tubercle projects. The urethral groove is present upon the cloacal

aspect of the genital tubercle, and the,margins of the groove (inner

genital folds) are widely separated at the base of the tubercle and

fade away posteriorly within the cloacal margins. The separate

visceral orifices opening into the cloaca are not recognisable by

external examination at this stage. Upon the lateral aspects of

the base of the genital tubercle, continued in a caudal direction

as the lips of the cloacal opening, are the labia majora (labio-

scrotal folds). The labio-scrotal folds are lost in a cephalic

direction upon the skin of the abdomen in front of the genital

tubercle ; they swell out as two ovoid prominences at its base,

and diminish again towards the tail by skirting, as slightly

elevated margins, around the cloacal orifice.

In the next stage, an embryo of 18 mm. shows that the genital

tubercle has elongated, and the urethral groove become closed by

the meeting of the labia minora along its cloacal aspect; but the

outstanding feature of the changes consists of an extensive growth

of the labio-scrotal folds (text-fig. 1,C & D). So extensive is this

growth that the base of the genital tubercle has been buried

beneath the ingrowing mesial margins of the two ovoid promi-

nences present in this position in the 9 mm. stage. The urethral

channel has been completed by closure of the inner genital folds,

and now, over this closed urethra, the outer genital folds have

met in the mid line. The dorsal portion of the cloaca is now

13*

196 __<DR._F. WOOD-JONES ON THE

Text-figure 1.

A, Embryo of 9 mm.

B. External genitalia; 9 mm. stage.

C. Embryo of 18 mm.

External genitalia: 18 msn. stage.

REPRODUCTIVE HISTORY OF THE MOLE.

A. Embryo of 23 mm.

B. Embryo of 27 mm.

Text-figure 2,

C. External genitalia: 27 mm. stage.

198 DR. F. WOOD-JONES ON THE

cut off from the base of the genital tubercle and the urethra

by the meeting of the outer genital folds: and in this way the

anus becomes a separate orifice.

The ingrowth and meeting of the outer genital folds take place

first at the base of the genital tubercle at the site of their first

prominence; and a constriction in their breadth in front of the

anus marks this first union. At this stage the anus is triangular

in outline, the apex of the triangle being directed towards the

genital tubercle and corresponding with the primary ingrowth of

the outer genital folds. In addition to this ingrowth, the outer

genital folds have also become elongated, so that although the

genital tubercle has increased greatly in length, it has become

buried to a great extent beneath the overgrowing outer genital

folds. Upon the portion of the genital tubercle which is still

Text-figure 3,

A. Hairless nestling: 9,59 mm.

B. External genitalia at this stage.

exposed beyond the fused outer genital folds, the median raphé,

formed by the union of the inner genital folds, is plainly visible.

This raphé does not extend to the free extremity of the exposed

genital tubercle.

By the time the embryo is approaching full term and is 27 mm.

long, the growth of the outer genital folds has become so pre-

ponderant that the genital tubercle is practically hidden beneath

REPRODUCTIVE HISTORY OF THE MOLE. 199

their invading margin (text-fig. 2, B & C). The anus has become

circular in outline and its margins have become more distinctly

separated from the skin which forms the perineum, for although

by no means so protuberant as in the adult, the anus of the more

mature embryo is situated upon the apex of a conical tumid

mound which rises from the general surface of the perineum.

The genital tubercle ensheathed in the overgrown outer genital

folds has become recognisable as the adult penis, and between

this penis and the anus the prominence of the outer genital folds

has diminished to make a more or less flattened area—the adult

perineum. By full term (30-40 mm.) the genital tubercle is

completely ensheathed, and hidden from view, within a prepuce

derived from the outer genital folds; the anus is situated at the

summit of a conical elevation of skin, and the perineum shows

neither a median raphé nor an elevation due to the presence of the

outer genital folds. The naked nestling shows the same condition

with all the parts considerably enlarged (text-fig. 3).

The stages that have been described take place in exactly the

same manner in both sexes; so it seems obvious, from external

examination, that in the female, as well as in the male, the

urogenital sinus must be carried forwards, by closure of the inner

genital folds, to the tip of the genital tubercle, which then

becomes secondarily ensheathed within the’ overgrowing outer

genital folds. In other words, it appears as though the outlet for

the genital products in both sexes must be at the extremity of

the penis or clitoris.

C. Post-natal Stages of External Genitalia.

As Thomas Bell truly said, “it is very difficult to know a

virgin female mole from the male.”

Moles breed but once a year, and an extended series of obser-

vations by Adams and others proves that in the great majority

of cases the three or four young are born during the month of

May.

These young grow very rapidly. According to Adams (7) they

double their length within the first ten days of their life; three

weeks after birth they are three-quarters grown (H. E. Forrest, 8),

and by the later months of the year they resemble the previous

year’s adults in all obvious characters.

I started collecting moles for examination in the autumn

months; and from among the many individuals secured it was

not difficult, after some experience, to pick out the females of the

year.

Text-fig. 4 shows the condition of the external genitalia present

in these young females. The particular specimen figured was

caught in November, and it shows but little modification of the

late foetal and early nestling conditions already described. Text-

fig. 5 shows the condition of a male taken at the same time, and

900 DR. F. WOOD-JONES ON THE :

in a general way its external genitalia may be said to be similar

to those of the female. Still, though it is admittedly a matter of

difficulty to be absolutely certain of the sex of an individual,

there are minor characters, which in typical cases furnish some

indications upon which to make a correct diagnosis fairly certain.

Text-figure 4.

External genitalia of a young female in the autumn of the first year.

In the first place the clitoris of the female, though remarkably

like the penis of the male, is, as a rule, slightly smaller, and in

most cases, its root slopes more gently to the perineum, so that

its base merges with the perineal skin with less line of demarca-

tion than is present in the male. Another fairly obvious dis-

tinction is the shorter distance between the root of the clitoris

and the anus in the female. In some cases, but I think not in

all, this sexual difference is noticeable; it was remarked on by

Geoffroy, and, taken with the condition of the genital organ,

it gives the best guide for determining the sex of young

individuals. RE

It will be noticed that in the virgin female in the late autumn

months, just as in the mature embryo, the perineum presents a

smooth unbroken surface. There is no wrinkle or pucker; no

depression or blind pit, to mark a closed female genital orifice.

It is not the case, as the description of Owen might lead one to

REPRODUCTIVE HISTORY OF THE MOLE, 201

suppose, that a vagina closed by a thickened hymen is present ;

for in the female as in the male, the perineal skin passes without

interruption from the base of the genital tubercle to the margin

of the anus. Such a description holds good, I think, for all first

year females during the autumn and winter months; but in the

early spring external changes are apparent. In the second week

-Text-figure 5.

External genitalia of a normal male.

of March in moles caught in Epsom, and during the third week

in moles caught in the neighbourhood of East Grinstead, the

perineum had become pigmented. This pigmentation, so far as

my observations go, occurs at first in the absence of any anatomical

change in the perineum.

202 DR. F. WOOD-JONES ON THE

The genital tubercle becomes more vascular, and, in the recently

dead specimen, is intensely injected and vivid red; the anus

shares to a lesser degree in this change, and although the vivid

redness may be in part a mere post-mortem phenomenon there can

be no doubt as to the increased vascularity of the parts. This

phase of pigmentation is shown in text-fig. 6. In well-marked

Text-figure 6.

External genitalia of a young female taken in March.

Stage at which pigmentation is present.

contrast to this redness of the genital tubercle, the perineum

shows a bluish coloration which is reminiscent of the sexual

perineal pigmentation displayed in some Primates. This blue

pigmentation is certainly not a post-mortem change, and indeed

there are some indications that it tends to fade after death.

Later in March this pigmentation becomes more vivid; the

whole perineal structures are enlarged and highly vascular, and

REPRODUCTIVE HISTORY OF THE MOLE, 203

the blueness of the base of the genital tubercle is increasingly

conspicuous. :

The next stage that I have had the opportunity of examining

in a perfectly fresh condition is that seen in specimens taken in

the first week of April, and in these a vaginal orifice is evidently

recently established in the midst of the blue area. This orifice

is placed with its long axis transversely in the perineum, and it

Text-figure 7.

External genitalia of a young female taken in April.

Stage at which perforation is present.

is a mere break in the continuity of the perineal tissues; its

margins show no signs of any genital folds such as are

commonly present in some form or other, nor has it even the

regular outline of the ordinary non-genital visceral orifices. This

opening is situated near to the base of the genital tubercle, but

I fancy that even its exact site is subject to some slight varia-

tions within the narrow limits of the perineum, and its distance

204 E “DR. F. WOOD-JONES ON THE-

from the genital tubercle -does not seem to be absolutely

constant (text-fig. 7). °

Towards the end of April the colour fades from the perineum,

but the female genital orifice remains widely patent, and during

the month of May parturition usually takes place. I have not

actually seen the bilateral perforations described by Adams, for

all the specimens I was able to procure at about the critical

period had either an entirely imperforate blue area, or else had a

vaginal cleft established. It has seemed to me in several

instances that the purple pigmentation was most marked im-

mediately upon either side of the middle line, and though I have

not myself yet come upon a stage in which the perforation was

bilateral, I have every reason to suppose such a condition to be

extremely probable.

Text-figure 8.

Winter female with pigmented perineal scar (Noy. 20th). Second year.

After parturition is effected retrogressive changes evidently

take place, and in the later months of the year some most

interesting phases are met with. This vaginal orifice, which is

formed in inch a curious way, t tends again to become occluded’ in

the autumn months, and in by far the greater number of

specimens that I have examined, the outlet for the female

REPRODUCTIVE HISTORY OF THE MOLE, 205:

genital ducts becomes entirely closed again within a very short

space of time.

Some specimens examined in the autumn months show nothing

more than a wrinkled surface at the base of the genital tubercle,

and somewhere in this wrinkled area is commonly a patch, or a

series of minute patches of dark pigment (text-fig. 8). Others

show a very definite transverse scar, more or less puckered,

and generally pigmented in some portion of its length (text-

fig. 9). At times this scar formation is irregular, and pockets

will penetrate some distance into the perineal tissues. In one

instance, a probe could be passed far into the vaginal mouth of

a November mole, although this vagina did not form an open

channel communicating with the uterus (text-fig. 10).

Text-figure 9,

Winter female with perineal scar (Dec. 5th). Second year.

It would therefore seem that in virgin moles a vagina is

formed in the early months of the spring; that after parturition

has taken place it closes again in whole, or in part, and that by

the autumn its orifice is represented by a mere perineal scar.

I do not know how long moles live, nor how many times they

breed, but it would seem that at every recurrent breeding season

the process was repeated, and the vaginal orifice re- -established.

So far as I have been able to determine, the vagina as a patent

206 DR. F. WOOD-JONES ON THE

canal communicating with the uterus does not usually become

established as a permanency in the mole, but the experience of

meeting with one winter female with a patent vagina would

make me prepared to believe that at times the communication

may be permanently established. We have in this process a

most remarkable parallel to the evolution of the medina vagina

of the Marsupials, to which reference will be made later.

Text-figure 10,

Winter female with a perineal orifice (Dec. 5th). Second year.

D. Embryonic Stages of Internal Genitalia.

In order to follow the embryonic development of the female

genital ducts and orifices, I have been compelled to prepare a

very large series of sections, for the difficulty of distinguishing

the sexes during embryonic life renders the selection of material

exceedingly difficult. The female ducts have been followed

in their entire length in embryos of 27 mm, and 33 mm.

respectively.

In the earlier of these stages the arrangement of the ducts is

comparatively simple, for in the female, just as in the male, the

genital ducts end separately in the floor of the closed uro-genital

REPRODUCTIVE HISTORY OF THE MOLE. 207

sinus. As the urinary passage passes from the bladder towards

the urethral outlet at the tip of the genital tubercle it describes

a curve of which the convexity is directed caudad: it is at the

lowest part of this bend that the female genital ducts terminate

in the urethra. The Mullerian ducts pass down dorsal to the

bladder and urethra, and then running ventrally with a curve

similar to that of the urethra, they terminate at the caudal floor

of the urethral bend.

This termination is made in the immediate neighbourhood of

the orifice of the Wolffian ducts, and a considerable proliferation

of epithelium marks the actual site of the ends of the ducts of

Muller (PI. I. figs. 1 & 2).

In the embryo of 27 mm. the continuity of the Mullerian

ducts and the urethral passage is easy to trace, although the

proliferation of the Mullerian epithelium makes the continuity of

the lumina difficult to establish with certainty.

So far this must be considered merely as a primitive arrange-

ment in which separate Mullerian ducts terminate in a uro-

genital sinus in the neighbourhood of the Wolttian ducts ; and

the peculiarity which exists in Zalpa at this stage is that the

uro-genital sinus is closed in, and carried forwards to the tip of

the genital tubercle, in the female as it is in the male. Were it

not for this penile prolongation of the female uro-genital sinus,

there would be nothing very remarkable in the primitive

condition of Zalpa at this stage.

Even in this peculiarity Zalpa is not unique; for very similar

conditions are known to be present in some other animals.

Among Insectivores several species have been described as

having the genital tubercle tunnelled by the urethra in the

female; this condition occurs again among the Lemurs, and,

according to Chapman, Capromys pilorides among the Rodents

shows a like conformation of the clitoris in the female. Tn all

these types, however, the Mullerian ducts open in the normal

way by a separate vagina, and no great physiological questions

are raised by this anatomical arrangement. But the condition

described by Morrison Watson (9) and by Chapman (10) in

Hyena crocuta appears to be an exact parallel to the virgin

condition of Zalpa. Assuming that it is not by the mere

coincidence that only virgin examples of Hycna crocuta have

ever been examined by anatomists, we must suppose that the

condition in this case is permanent; that no new vagina is ever

formed; that copulation takes place via the urethra, and that the

young are born by the same route—through the tip of the

clitoris.

In Talpa it is evident that this primitive condition does not

last long, for already in the embryo of 33 mm. the Mullerian

epithelial proliferation has proceeded a stage further and the

lower ends of the Mullerian ducts are somewhat more separated

from the wall of the urinary passage. The condition seen at this

stage is as follows. The Mullerian ducts which above are united

208 : -DR. F. WOOD-JONES ON THE

into a common chamber having a wide lumen, separate again as

they approach the pelvis, and the lumen of each individual duct

becomes occluded by the proliferating epithelium of its walls

(Pl. II. fig. 2). In this occluded state they pass as epithelial

cords dorsad to the urinary canal and, sweeping into the pelvis

caudad to it, may be traced past the openings of the minute

Wolffian ducts.

From this point onward each Mullerian cord may be traced as

a delicate epithelial strand caudad and ventrad of the urethra,

still in intimate relation with the wall of the urethra, but

apparently altogether separated trom its lumen.

This is the state of affairs at the caudad bend of the urethra,

From here the urethra and the crura clitoridis turn cephalad

again to the genital tubercle, the tissues of which they penetrate ;

but the Mullerian strands do not take part in this cephalad bend,

for they continue their course towards the base of the genital

tubercle rather than towards the central mass of its tissues. It

is here that the most curious feature in the development of the

female genital system of Talpais seen. The genital tubercle itself

is, of course, covered by the general many-layered cutaneous epl-

thelium which here shows but few hair rudiments. At the lateral

and inferior margins of the genital tubercle just where it joins

the general skin of the perineal region, the deeper layer of the

perineal skin dips inwards into the perineal tissues as two lateral

cords which become continuous below the sides of the urethra

with the similarly solid Mullerian cord.

I have spoken of these solid cords as being bilateral, but in the

embryo of 33 mm., of which I have serial pelvic sections, I have

been able to trace the continuity of the surface epithelium with

the Mullerian cord only upon the left side, although in the same

situation upon the right side is a well-marked stream-line of

the cells of the perineal tissues (Pl. III. fig. 1, the figure is

reversed), This unilateral condition of the epithelial ingrowth

is worthy of notice, for again at a later stage I have observed the

same thing : and it becomes significant when taken in conjunction

with the observation of Adams that the adult vagina is perforated

by bilateral breaking down of the perineal tissues. The changes

that have taken place in the interval between the 27 mm. stage

and the 33 mm. stage are not really very great, for the rudiments

of all the conditions described in the 33 mm. embryo are already

present in that of the 27 mm. embryo, Although in the younger

example the Mullerian ducts definitely join the urethral wall at

their lower ends, still, even at this stage, an epithelial prolongation

from them may be followed around the caudal bend of the urethra

towards the base of the genital tubercle, and, moreover, an epi-

thelial ingrowth from the skin of the base of the genital tubercle

is already in process of development. In this younger embryo

the ingrowth is again unilateral, being found in my series only

upon the right side (Pl. IT. fig. 1: the figure is reversed).

The embryonic condition of the female passages is therefore a

REPRODUCTIVE HISTORY OF THE MOLE. 209

curious one—though it must be remembered that it has its

parallel in general mammalian, and even in human development.

The Mullerian ducts open first into a cloacal uro-genital sinus ;

at a later stage this opening is lost and the lumen of the ducts

becomes obliterated. The Mullerian ducts become mere Mul-

lerian cell-strands, and these cell-strands pass forwards below the

urethral floor, and mesial to the crura clitoridis, towards the base

of the genital tubercle. It is at the base of the genital tubercle

that the uro-genital sinus approaches nearest to the surface of

the body, and here a proliferation of surface epithelium dips in

towards it.

This epithelial invasion is crescent-shaped in outline (correspond-

ing with the base of the genital tubercle), and the extremities

of the crescent dip into the perineal tissues and meet the

Mullerian epithelial strand. In the embryos examined the right

horn of the crescent in one case, and the left in the other, alone

extends inwards to the termination of the Mullerian strand.

E. Postnatal Stages of Internal Genitalia.

(i.) The Nestling Mole.

The young mole is born in a naked and rather immature

condition, and although the growth within the first few days of

life is extraordinarily rapid, it is not until about the 9th day

that the colour changes from the primitive pink to the dull lead-

colour that marks the advent of hair. At about the end of the

first fortnight the actual hair appears.

This period of life has been fully observed from the field-

naturalist’s point of view by Adams, Cocks, and Evans (all quoted

by Barrett-Hamilton, 7).

It was obvious from the outset of my investigation that the

nestling period was one of the utmost importance, for it was

evident that it was not only a time of extremely active body

growth, but that remarkably rapid internal changes must be

taking place as well. Indeed there may be said to be two

well-marked phases of wonderfully rapid evolution in the repro-

ductive system of the mole, the one being present during this

nestling period, and the other being marked by those almost

violent changes which occur during the short but intense period

of sexual activity.

I have found specimens of nestling moles particularly difficult

to obtain, and during two breeding seasons I have failed to find

nestlings, although I have had adults in all phases of sexual

activity. For the only material which I have been enabled to

examine microscopically in serial sections, I am indebted to

Professor L. Doncaster. This material, which consisted of two

specimens, is of interest, for the nestlings were presented to the

Cambridge University Museum of Zoology by the Rev. Leonard

Jenyns, and had been in their bottle of spirit for at least some

Proc. Zoo. Soc.—i914, No. XIV. 14

210 DR. F. WOOD-JONES ON THE

seventy or eighty years when the sections were cut, and yet the

histological condition of the tissues remains extremely good.

The nestlings are immature and pink, and are devoid of obvious

hair except along the margins of the tail. The eye has become

further reduced from the late feetal condition; the ear appears

in extraordinary proximity to the shoulder, and the genitalia are

in practically their adult condition. The vertex-rump measure-

ment of.one (2) is 59 mm. (text-fig. 3, A) and of the other (¢)

62 mm.

The serial sections of the female specimen show that a great

adyance has been made on the condition seen in the full-term

embryo, the development of the genital system taking place

between the 33 mm. and 59 mm. stages being remarkable.

In the first place the utero-vaginal canal has lost all trace of

its bilateral origin, for the Mullerian tract is now a wide canal

with well differentiated walls. The whole picture of the pelvic

relations has altered by this rapid growth of the female genital

passages, for whereas in the embryonic stages the urinary tract

was far wider than the genital passage, in the nestling the utero-

vaginal canal is the most conspicuous viscus in this region

(BIR esos. 2):

Another well-marked change is the complete separation of

the cavities and the almost complete separation of the walls of

these two passages, for it is only over a very small area that the

musculature of the genital and urinary tracts becomes united.

But although the separation of the two channels has thus become

more marked, there is as yet but little advance on the embryonic

condition towards the establishment of a new orifice for the

female ducts. The epithelial ingrowth at the base of the genital

tubercle has become more conspicuous, and with the increased

growth of the parts it is still further separated from the

centrally-lying urethra.

Advanced as is the development of the female passages at this

stage, there is, however, still a very imperfect union of the

epithelial ingrowth and the utero-vaginal canal.

In the specimen examined in serial sections it is only upon

the right side that the ingrowth meets the patent utero-vaginal

canal; so that it is only on the right of the urinary passage that

there is even an epithelial continuity between the female genital

system and the surface of the body. The left side of the epithelial

crescent is well developed, but it remains widely separated from

the utero-vaginal lumen (PI. ITI. fig. 2: the figure is reversed).

(ii.) The Adult.

There is no need to describe the uterus and its adnexa in the

adult, for this description has been furnished many times by

comparative anatomists who have examined second year females.

It was evidently only the female of the second year that was

examined by John Hunter, for he says (11):—‘‘ There is no

common vagina; the vagina is very long and runs in a serpentine

REPRODUCTIVE HISTORY OF THE MOLE, 211

eourse forwards and backwards.” By “‘common vagina” Hunter

doubtless means urogenital sinus, and the serpentine condition of

the vagina may be taken as an indication that the mole dissected

was not a virgin female in its first year. It is remarkable that

no chance ever brought to John Hunter an example of a female

mole in its virgin condition. Our knowledge is the poorer as a

consequence; for the reflections of the man who observed that

‘there is very little fat on a mole” upon discovering the absence

of an outlet for the female genital tract can well be imagined.

Text-figure 11.

Z oe

= aoa E

. arg sp, Z

. INTE ELL

(head

\ e Z by \ 4.

€ 4d, 3\ ES iY }

ee Wet

yA SNS

Section through the pelvis of a female of the first year (November).

The female genital passages are represented by the solid epithelial strand (ep.sti.).

r., rectum ; gl., gland; w., urethra; gen. tub., genital tubercle.

The descriptions of Owen and others are obviously founded

upon the examination of similar material.

In describing the genitalia of any mammal—and this applies

more especially to the Insectivora—it is almost essential to record

the condition of sexual activity in which the organs were at the

14*

212 DR. F. WOOD-JONES ON THE

time of examination. In most of the Insectivora there is an

almost complete anatomical transformation during the brief

flare up of sexual activity at the pairing season.

During a very few weeks, parts which were altogether insig-

nificant have become increased ten times, or more, in size; and

this is even more obvious in the male than in the female.

Text-figure 12.

SOS

YS o

ul SF

Yaxs

Ay,

f CSRS

og: rs n

ieee

Section through the pelvis of a female of the second year (April).

The epithelial strand has now broken down into the wide open vaginal canal (0.2.c.).

r., rectum ; gl., land; wr., urethra; gen. tub., genital tubercle.

The wonderful transformations which take place in the male

genital glands have been described in detail by histologists. The

whole of the female genital tract also undergoes these changes

during the months of March and April, and an enormously

increased blood supply is furnished to the genitalia during these

months.

Examined during the autumn months of its first year, the

REPRODUCTIVE HISTORY OF THE MOLE. 213

female shows but little advance upon the condition described as

existing in the nestling of 59 mm, ‘The utero-vaginal canal is a

wide cavity which narrows below the neck of the bladder and

passes, still as an open tube, towards the perineum at the base

of the genital tubercle. But at a point some distance from the

perineal surface it becomes closed by the approximation of its

anterior and posterior walls, and is continued as a wide area of

epithelial cells which becomes continuous with the skin at the

base of the genital tubercle (text-fig. 11, p. 211).

There is, therefore, no outlet for the female genital passages,

for the whole of the lower part of the tract is a solid epithelial

mass, just as it is in the 59 mm. nestling.

The greatest advance to be noted in the adult examples of

which I have prepared serial sections, is that the solid epithelial

area is of greater lateral extent and becomes continuous with the

whole of the lower, patent, extremity of the utero-vaginal canal.

In the pigmented young females taken in April a very great

change is seen.

All parts of the genital tract are much enlarged, their tissues

are all more vascular, and the large pre-anal gland mass has

increased greatly in size. But the most important change

consists in the complete opening of the lower part of the utero-

vaginal canal by the desquamation of the central cells of the

previously solid epithelial mass (text-figs. 12, 18, A). So far as

can be determined from the material at my disposal, it would

appear that with the increased vascularity of the parts in the

months of March and April, a proliferation of the epithelial mass

is initiated, that the central cells of the mass degenerate and are

shed, and the peripheral cells become the lining epithelium of

the cavity thus opened up (text-fig. 13, B).

In sections of the genital tract of non-virgin females caught

in the autumn months the retrogressive changes are seen. The

lower portion of the utero-vaginal canal is again occluded, but

the occlusion is not uniform throughout the epithelial portion.

Pockets run in from the surface, and in some cases these are deep

enough to admit the tip of a probe, but beyond these pockets the

occlusion is again complete in whole or in part.

F. Summary and Conclusions.

(a) Although the peculiarities of the female reproductive

system of Zalpa ewropea appear at first sight to be extremely

anomalous, nevertheless they have, for the most part, some

foreshadowings in the normal processes of development of other

mammalian types.

(0) Everything that is unusual concerning the female repro-

ductive system is initiated in the early stages of embryonic

development when, between the 9 mm. and 18 mm. stages, the

labio-scrotal folds begin to grow towards the middle line over

the closed urethra in the base of the genital tubercle.

214 DR. F. WOOD-JONES ON THE

may be said that, in the method of development of the external

genitalia, the male falls into line with some other members of

the Insectivora, the Rodentia, Ungulata, and some other Orders.

(d) It is the female alone that is anomalous in following from

the first a male type of formation of the external genitalia.

(e) Even this state of affairs would seem to be found again in

the female of Hycwna crocuta.

Text-figure 13.

A. Median section of pelvic viscera of a female of the first year (November).

The utero-vaginal canal ceases to be an open passage some distance from the surface,

and is continued to the surface only by an epithelial strand.

B. Similar section of a female of the second year (April).

The utero-vaginal canal is open to the exterior at the base of the genital tubercle.

(/) There is nothing really remarkable in the shifting of the

site of opening of the Mullerian ducts, nor is it unparalleled for

them to end in solid epithelial prolongations.

(g) Both these phases are found in the normal development of

REPRODUCTIVE HISTORY OF THE MOLE, 215

other animals, and both are stages in the normal formation of

the female genital system of Homo (12). The imperforate vagina

is a normal stage in the female human embryo, and this solid

vagina becomes patent in very much the same way as does that.

of the mole—by desquamation of its central cells.

{h) The moje is exceptional in that the opening up of the

solid vagina is so long delayed.

(2) Again, it is not without parallel that a vagina once formed

should become occluded again, only to be reopened when next.

functional activity demands a passage; these phases have been

established by Hill (5) with regard to the median vagina of

Perameles.

Bibliography.

(1) Errenne Grorrroy (St. Hilaire).—Cours de histoire

naturelle des Mammifeéres. Paris, 1829.

(2) Tuomas Betu.—Article “ Insectivora” in Todd’s Cyclopedia

of Anatomy and Physiology, vol. ii. p. 1005. 1839.

(3) RicHarD OwEn.—Comparative Anatomy and Physiology of

the Vertebrates, vol. iii. p. 609. 1868.

(4) Lionet E. Apams.—Manchester Memoirs, vol. xlvii. No. 4.

1903.

(5) James P. Hini.—Proc. Linn. Soc. New South Wales, 1889,

pt. 1., March 29th, p. 42.

(6) F. Woop-Jones.—The Development and Malformations of

the Glans and Prepuce. Brit. Med. Journ., Jan. 15th,

ILO.

(7) Various authors quoted by G. EK. H. Barrert-HAminron.—A

History of British Mammals, pt. vii. p. 37.

(8) H. E. Forrest.—The Fauna of Shropshire, 1899, p. 45.

(9) Morrison Watson.—On the Female Generative Organs of

Hyena crocuta. Proc. Zool. Soc. Lond., May 1877,

p- 369.

(10) Henry C. Cuapman.—Observations on the Female Gene-

rative Apparatus of Hyena crocuta. Proc. Acad. Nat. Sei.

Philadelphia, 1888, p. 189.

(11) Joun Hunrer.—Essays and Observations. Edited by Owen,

1861, vol. i. p. 188.

(12) F. Woop-Jonzs.—The Development and Malformations of

the Urogenital Passages, etc. Brit. Med. Journ.,

Dec. 17th, 1904.

EXPLANATION OF THE PLATES.

Puate I.

Fig. 1. Section of the pelvis of the 27 mm. embryo showing the lower ends of the

Mullerian ducts.

2. Section of pelvis of 27 mm. embryo, slightly lower than fig. 1, showing the

extreme limit of the caudal bend of the urethra, and the tissue in which

the Mullerian ducts end.

216 ON THE REPRODUCTIVE HISTORY OF THE MOLE.

Puate II.

Fig.3. Section of pelvis of 27 mm. embryo at a higher level than figs. 1 & 2 (Pl. I.).

Section passes through the base of the genital tubercle and cuts through

the urethra in three places.

3a. Key to show the planes of sections illustrated in figs. 1-3.

4, Section of pelvis of 33 mm. embryo showing the double Mullerian ducts.

Puate III.

Fig.5. Section of pelvis of 33 mm. embryo at a slightly lower level than fig. 4

(Pl. II.). The section passes below the caudal bend of the urethra and

shows the lower ends of the Mullerian ducts, the strand of tissue in which

they terminate, and the epithelial ingrowths from the base of the genital

tubercle.

6. Section of pelvis of 59 mm. nestling, just shaving the base of the genital

tubercle. In more caudal sections the lumen A becomes continuous with

the lumen B.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 217

EXHIBITIONS AND NOTICES.

February 3, 1914.

Sir JoHn Rose Braprorp, K.C.M.G., D.Sc., F.R.S.,

Vice-President, in the Chair.

The Srcrerary read the following report on the additions to

the Society’s Menagerie during the months of November and

December, 1913 :—

NoveEMBER.

The number of registered additions to the Society’s Menagerie

during the month of November was 350. Of these 251 were

acquired by presentation, 51 by purchase, 19 were received on

deposit, 21 in exchange, and 8 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 237.

Amongst the additions special attention may be directed to :—

1 White-tailed Guereza (Colobus caudatus), from East Africa,

new to the Collection, presented by G. St. J. Orde Browne, Esq.,

on November 3rd.

1 Cheetah (Cynelurus jubatus), from East Africa, presented

by H.G. The Duke of Sutherland, F.Z.S., on November 7th.

1 Argentine Blackbird (Jerula fuscatra), new to the Collection,

presented by R. Suggitt, Esq., on November 12th.

10 White-headed Starlings (Poliopsar lewcocephalus), from

Cochin China, new to the Collection, presented by W. O.

Danckwerts, Hsq., K.C., F.Z.S., on November 26th.

2 Red-billed Hornbills (Lophoceros erythrorhynchus), from

Africa, received in exchange on November 11th.

1 Crowned Pigeon (Goura coronata), bred in the Menagerie

on November 29th.

1 Hybrid between a Black-winged Peacock (Pavo nigripennis)

and a Domestic Hen (Gallus domesticus), presented by R. P.

Wheadon, Esq., on November 7th.

2 Kagus (Rhinochetus jubatus), from New Caledonia, presented

by the Marquess of Tavistock, F.Z.S., on November 1st.

2 Masai Ostriches (Struthio massaicus), from Hast Africa,

purchased on November 3rd. .

A large Collection of Small Birds and Waterfowl, presented by

Heatley Noble, Esq., F.Z.S., on November 24th.

DECEMBER.

The number of registered additions to the Society's Menagerie

during the month of December was 166. Of these 89 were

acquired by presentation, 31 by purchase, 19 were received on

deposit, 26 in exchange, and 1 was born in the Gardens.

The number of departures during the same period, by death

and removals, was 277.

218 MR. D. SETH-SMITH ON HYBRID BIRDS.

Among the additions special attention may be directed to :—

1 Pigmy Hippopotamus (Cheropsis liberiensis) 3 , from Liberia,

presented by H.G. The Duke of Bedford, K.G., Pres.Z.S., on

December 16th.

2 Vicunas (Lama vicugna) $ 2, from the Andes, presented

by F. Lesser, Esq., on December 20th.

1 Cheetah (Cynelurus jubatus) 2, from British East Africa,

presented by Mrs. McMillan, F.Z.S., on December 20th.

1 Pine-Marten (Mustela martes), from Kerry, presented by

H. G. Constable, Esq., F.Z.S., on December 11th.

3 Four-banded Chipping-Squirrels (Hutamias quadrivittatus),

new to the Collection, deposited on December Ist.

1 Olive-backed Thrush (Hylocichla ustulata swainsoni), from

North America, new to the Collection, received in exchange on

December 23rd.

A large Collection of Birds, including a pair of Red-collared

Lorikeets (Lrichoglossus rubritorques), a number of Waterfowl

and Small Birds, presented by Heatley Noble, Esq., F.Z.8., on

December Ist.

4 South-African Ostriches (Struthio australis), bred in South

Australia, deposited on December 12th.

29 Ornamented Ceratophrys (Ceratophrys ornata), from the

Argentine, presented by E. G. Robinson, Hsq., on December

23rd.

Mr. D. Seru-Smiru, F.Z.8., Curator of Birds, exhibited a

photograph of two hybrids between a Peacock and a hen Guinea-

fowl which were bred in Germany and were now in the Berlin

Zoological Gardens, and remarked that there were three or four

cases on record of such hybrids being produced, whereas there

were only two instances known of the Peafowl crossing with

the domestic fowl, and none, so far as he knew, with birds of

any other genus.

Mr. Seth-Smith also showed the skin of a hybrid Pheasant hen,

which had been sent to him by Mr. H. J. Elwes, F.R.S. This

bird was one of a number bred in Sussex by Mrs. Johnstone,

between a cock Calophasis mikado and hen (’, elliott. It bore a

very strong resemblance to the female C’. mikado, but showed

a trace of the,black throat and rufous barring to the tail of

C. ellioti.

Mr. G. A. Boutencer, F.R.S., F.Z.S., gave an account of the

collections of Batrachians and Reptiles made by the British

Ornithologists’ Union and the Wollaston Expeditions in Dutch

New Guinea. Four species of Batrachians and eight species of

Reptiles were described as new.

This paper will be published in the ‘ Transactions.’ .

MR. G. JENNISON ON A HYBRID SEA-LION, 219

February 17, 1914.

Prof. E. A. Mincutn, M.A., F.R.S., F.Z.S., Vice-President,

in the Chair.

The Srecrerary read the following report on the additions

to the Society's Menagerie during the month of January

1914 :—

The number of registered additions to the Society’s Menagerie

during the month of January was 151. Of these 73 were

acquired by presentation, 23 by purchase, 23 were received on

deposit, and 32 in exchange.

The number of departures during the same period, by death

and removals, was 209.

Amongst the additions special attention may be directed to :—

A pair of Park Cattle (Bos tawrws), from Chillingham, presented

by the Karl of Tankerville, F.Z.8., on January 28th.

A pair of Indian Wild Dogs (Cyon dukhunensis), from Nagpur,

presented by Mrs. H. A. Crump on January 23rd.

1 Brown-shouldered Hangnest (/eterus pyrrhopterus), from

Brazil, new to the Collection, presented by T. W. Smith, Esq.,

on January 20th.

2 Cuban Conures (Conurus ewops), new to the Collection,

presented by the Marquess of Tavistock, F.Z.S.,on January 23rd.

1 Abingdon-Island Tortoise (Zestudo abingdoni), from the

Galapagos Islands, new to the Collection, purchased on January

12th. j

1 Queensland Long-necked Terrapin (Chelodina expansa), from

Queensland, new to the Collection, purchased on January 19th.

A Hybrid Sea-Lion.

Mr. Grorce JENNISON sent for exhibition a mounted specimen

of a hybrid Sea-Lion (Ofaria pusilla § x O. californiana 9),

which had been born in the Belle Vue Gardens, Manchester,

together with photographs of another specimen now living in the

same Gardens, and the following notes :—

“Both the parents, O. pusilla g and O. californiana 9°,

arrived here as immature specimens. The male had been over

four years, the female six years in the Collection when the

first pup was born in June, 1911. It was found dead. The

next one, in 1912, lived only about a day. It was taken

from the parents who were tossing it about. The keeper

(J. Craythorne) came to the conclusion that the ill-treatment

was nervousness and excessive solicitude, and accordingly

separated the female a few days before the next birth was

expected, and prepared a corner fenced from the water into

which he took the pup—they are always born on land—when

it was born on June 15th, 1913. The mother followed the little

220 MR. E. G. BOULENGER ON THE

one, pushed the pup with her head until it found the teats, and

the rearing has presented no difficulty. A few interesting features

presented themselves.

“On June 21st I had the partition removed, and the pup

immediately went into the water and continued to return there,

although the mother threw it out by the neck repeatedly, finally

in so rough a manner as toendanger its life. June 28th, the pup

climbed over the two-foot fence. This was then raised to three

feet, which was beyond its powers. ‘The keeper tells me that on

July 2nd the mother carried the pup by the neck to the water,

kept it in the tank and on the side for half an hour, and then

carried it back to the safe retreat. Neither then nor subsequently

did I see this interesting sight. And later the mother seemed

rather careless of the young one when in the water. An occasional

bath seemed necessary, and as the little one had no notion of sur-

mounting the three inches from the water to the side of the bath,

the keepers had to watch his swim—he is a male—and then lift

him out, a duty that increased in difficulty as he grew stronger

and, strangely enough, more shy.

“‘ At five months old it learned to leap out. At six months it

began to play with fish given to the mother, and after a fortnight

it ate a little. It still sucks. It is very lively in the water,

fond of swimming on its back and stroking its nose with the

fore flippers, a trait common to Cape Sea-Lions which I have

never noticed among the Californians. The pup has a pearly-

grey pelage; the head is round and well bombed, and the whole

animal bears a close resemblance to the male parent.

“The adult animals are kept in the open with free access to a

shelter which they never use.

“The male is usually in the water and generally drives the

female out very quickly. This hostility is suspended in severe

weather, when both animals swim continually to keep the ice

from forming.”

Giant Saddle-backed Tortoise.

Mr. E. G. BouLencer, F.Z.S., Curator of Reptiles, exhibited a

photograph (text-fig. 1) of a female example* of the Giant Saddle-

backed Tortoise (Testudo abingdonii), recently purchased by the

Society. On the arrival of the tortoise, the origin of which was

unknown, Mr. Boulenger was somewhat puzzled as to the species

to which it should be referred, but, on carefully comparing it with

the Saddle-backed Tortoises in the British Museum, came to the

conclusion that the specimen was none other than the hitherto

unknown female of ZY. abingdonii, a species which had never

previously been brought to Europe alive, and which was thought

to be extinct. An inspection of the very representative collec-

tion of Giant Tortoises in the Tring Museum strengthened the

conclusion arrived at.

* [The tortoise has since died, and turned out to be a male-——EprTor, P. Z. 8.] ,

GIANT SADDLE-BACKED TORTOISE. 22%

From the few male specimens in museums this tortoise differs

by the fore part of the shell being less strongly compressed and

not reaching the same height, more as in 7. ephippium, by the

“comparatively greater breadth of the hinder part of the carapace,

by the broader bridge, and by the smaller size of the marginals

bordering the bridge.

Text-figure 1.

Giant Saddle-backed Tortoise (Lestudo abingdonii).

Measurements. ties

Length of carapace in straight line .........-....+--. 25

Length of carapace OVer CULVE ...-..-.e ese eeeeee sree 27

Width of carapace in straight line.................-+. 17

Width of carapace over curve.......... Ras as 5 AA

Width of hinder part of carapace in straight line } 17

(from centre of 9th marginal)................++++++

Width of hinder part of carapace over curve .... 2 4

Vertical height in front ........... MEN A PENA ET eee 92

Length of plastrom ......-.....-.:seeeeeeeeee terns tee eee 19

Width of plastron ..........seseseee sense nets eee ne eens 163

Width of bridge ..,.........c:ceeceeeee eens ee enenenenenes 10

Depth of marginals, bordering WSIS.’ Seb ecesontecncn 4

222 DR. R. T. LEIPER AND SURGEON E. L. ATKINSON

Mr. Boulenger pointed out that the neck, which exactly

equalled the shell in length, was not as represented in the single

stuffed specimen in the British Museum, without any folds, but

that the skin was loose, with numerous folds, a pair of very

strong lateral ones extending from the sides of the head down

almost the entire length of the neck.

The tortoise, although very lively, is extremely shy, and on

one’s approach has the peculiar habit of withdrawing its head,

standing as high up as possible on its hind limbs, and bending

over until the fore part of the carapace almost touches the

ground. The purpose of this perfgrmance seems to be in order

to protect the head and neck which, owing to the peculiar shape

of the front part of the shell, remain completely exposed on

withdrawal.

The shell in this species being of extraordinary thinness and

lightness, the creature is able to progress at a pace which,

compared with that of the other tortoise in the collection, is

remarkable for its rapidity.

Helminthes of the British Antarctic Expedition, 1910-1913 *.

Dr. R. T. Lerper, M.B., F.Z.8., and Surgeon E. L. Arkinson,

R.N., gave a lantern demonstration of the Helminthes collected

by the British Antarctic (‘Terra~-Nova’) Expedition, 1910-1913.

In all, thirty-seven species of Helminthes were collected by

Surgeon Atkinson on the voyage of the ‘ Terra-Nova,’ and whilst

a member of the shore party. One species was a free-living

Nematode, Leptosomatum setoswm, the remainder were parasitic.

Of these latter,

8 species had been recorded from the Antarctic zone, viz. :—

In Leptonychotes weddelli:

Ascaris osculata Rud.

Ascaris radiata v. Linstow.

Ascaris rectangula v. Linstow.

Corynosoma antarcticum Rennie (=C. hamanni v. L.).

Dibothriocephalus mobilis Rennie & Reid.

Dibothriocephalus coatsi Rennie & Reid.

Diphyllobothrium perfoliatum Railliet & Henry.

In Aptenodytes forstert :

Anomotenia zedert (Baird),

Free-living Nematode:

Leptosomatum setosum v. Linstow.

* From the Helminthological Department of the London School of Tropical

Medicine.

ON HELMINTHES. Se 223

3 species previously found in the Arctic zone are now recorded

from the Antarctic for the first time, viz. :—

In Megaptera longimana ;

Filaria crassicauda Creplin.

Echinorhynchus turbinella Dies.

In Leptonychotes weddelli :

- Ogmogaster plicatus (Creplin). This species isa parasite

of whales in northern seas.

1 species recorded outside the Antarctic zone has now been

found within the Circle, viz. :—

In Megalestris maccormicki :

Tetrabothrius cylindraceus Dies.

15 species found within the Antarctic circle are new, viz. :—

NEMATODA.

KATHLEENA SCOTTI, sp. n., from Diomedea melanophrys.

An Ascarid nearly related to Ascaris osculata which is

designated type, infra, of a new genus Kathleena.

Whitish, firm, round-worms. Male 15x0-9 mm. Female the

same or slightly larger. Interlabia very large, pentagonal.

Short-curved cesophageal appendage 0°2 mm. Intestinal cecum

18 mm. (ésophagus 2°53x0-4 mm. Spicules 3x2°7 mm.

Tail of male terminates in blunt digitate process.

ACANTHOCEPHALA.

ECHINORHYNCHUS CAMPBELLI, sp. n., from Zrematomus ber-

nacchit.

Male 9mm. Female 10 mm. Thin-walled. 2:5 mm. broad.

Proboscis 2 mm. Hook-bearing rostellum 0°5 mm. Hooks 14

linear series of 8 hooks each. ‘Testes oval, occupy the third

fourth of the body.

EcCHINORHYNCHUS RENNICKI, sp. n., from Zrematomus ber-

nacchit.

Male3°7mm. Female4mm. Proboscis] mm. Hook-bearin

rostellum 0°73 mm. Hooks in 12 linear series of 6 each. Those

of alternate rows are in line transversely. Each hook protrudes

from a transparent cuticular lapel. Lemnisci are long and

slender.

ECHINORHYNCHUS DEBENHAMI, sp. n., from Zrematomus ber-

nacchi.

Male 2°2 mm. Female 2:2 mm. Sickle-shaped. Stout,

cylindrical rostellum with hooks in 12 linear series of 6 each.

Lemnisci bag-like, extending but little behind proboscis. Testes

large, occupying anterior half of body-cavity, deeply lobed.

Female crowded with eggs.

224 DR. R. T. LEIPER AND SURGEON E. L. ATKINSON

TREMATODA.

HEMIURUS OATESI, Sp. n., from 7rematomus bernacchii.

Length 2mm. Abdomen present but retracted wholly. Skin

sharply striated. Ventral sucker 0°34 mm. diam., twice that of

oral sucker. Enormous muscular seminal vesicle. Yolk-masses

compact, lobulated. Eggs exceedingly numerous and small.

APONURUS BOWERSI, sp. n., from Zrematonmus bernacchit.

Length 1 mm. Oral sucker has characteristic fleshy lip along

dorsal rim only. Gut-branches greatly dilated extend to posterior.

end of body. The yolk-glands are peculiar: two halfmoon-shaped

solid masses lying in apposition immediately in front of the

ovary.

LEPODORA GARRARDI, sp. n., from Zrematomus bernacchii.

Flat fleshy forms 3x0°9 mm. Brownish colour due _ to

numerous yolk-glands. Skin covered with delicate spines.

Ventral sucker 0°27 mm., oral sucker 0°37 mm. Stout pyriform

pharynx 0°2 mm. Eggs few but large. Testes tandem. Gut-

branches wide and extending to posterior end of body.

PoDOCOTYLE PENNELLI, sp. n., from 7'rematomus bernacchii.

Small forms tapering from large pouting ventral sucker.

Armed cirrus extends to posterior level of the ventral sucker.

Yolk-glands large and discrete. Testes smooth, tandem. Eggs

large, with flat knob-like protrusion at one pole.

ALLOCREADIUM FOWLERI, sp. n., from Zrematomus bernacchit.

Immature forms 0°74 mm. in length, 0°4 mm. broad. Skin

smooth. Cylindrical excretory vesicle with fine black pigment

granules. Large ventral sucker 0°36 mm. Three small round

bodies 0°1 mm. in diam. represent the genital glands.

CESTODA.

DIBOTHRIOCEPHALUS LASHLEYI, sp. n., from Leptonychotes

weddelli.

3 to 4 cms. Young segments quadrate. Mature segments

3 to 4 times longer than broad. Head 1:2 mm. long x 0°77 mm.

broad. Suckers, situated laterally, are almost circular and do not

extend much down the head. Eggs commence at 14th segment,

0-06 mm. The testes extend inward in each segment in single

series of three.

DIBOTHRIOCEPHALUS ARCHERI, sp. n., from Leptonychotes weddelli.

6 to 12 cms. Large square head 2:04 mm. broad. Lips of the

suckers folded inwards. Eggs start at 57th segment and measure

0:07 mm. ‘Testes scattered diffusely.

DIPHYLLOBOTHRIUM RUFUM, sp. n., from Leptonychotes weddelli.

3 to 6 cms. The head is characteristically pigmented brick-red

around the base of the suckers. Head. measures 1°64 x 1:44 mm.

ON HELMINTHES. - 225

Suckers are dorsal and ventral, The segments overlap markedly

as in D, perfoliatum. Hggs measure 0°025 mm.

ORIANA WILSONI, gen. et sp. n., from Lalenoptera borealis.

Segments all immature. Strobila 13 ems. Head discoidal,

3 mm. in diam., quadrate in outline, 4 round suckers present

terminally. Neck very slender. Testes arranged in two definite

groups of 7-8 and 17-18. Near to Diplobothrium.

TETRABOTHRIUS WRIGHTI, sp. n., from Pygoscelis adelie.

Strobila 2°2 mm., but none contains eggs. Head 0:4 mm. in

length. Testes constantly twelve, auricular appendages of suckers

well developed.

ANTHOBOTHRIUM WYATTI, sp. n., from Trematomus bernacchii.

Small scolices, unsegmented. Four large auricular appendages

each occupied by two tandem suckers. A bright pigmented band

crosses the neck in the living state.

9 species were collected in Tropical and Temperate Zones during

the voyage of the ‘Terra-Nova.’ Of these three have been

recorded previously, viz. :—

ABOTHROS CARCHARIAS and a larval TrerrarHyncHus from

Carcharias sp., and TETRABOTHRIUS HETEROCLITUS Dies. from

Puffinus cinereus.

Five Cestodes, all of the genus TETRABOTHRIUS, are new, viz. :—

TErRABOTHRIUS CREANT, sp. n., from Wstrelata trinitatis and

i. arminjoniana.

Strobila 4°5 ems. Head 0°84 mm. broad, carries four suckers

but no rostellum. Testes are numerous. Yolk-gland large.

Cirrus 0:06 mm.

TETRABOTHRIUS CATHERINA, sp. n., from Wstrelata trinitatis.

Stouter than preceding. Head comparatively small. Suckers

most on top of head. Segments overlap succeeding segments

by one-third. Testes 30 to 45, bunched in middle of segment.

Genital organs very characteristic. Cloaca divided into outer

and inner portions. ‘There is a large pyriform seminal vesicle

internal to the cirrus.

TETRABOTHRIUS AICHESONI, sp. n., from Wstrelata trinitatis.

Strobila 3 cms. Moreslender than preceding. Segments more

uniform, only overlap slightly. Testes arranged in three distinct

sets, are very numerous, far in excess of those of the previous

forms.

TETRABOTHRIUS PRIFSTLEYI, sp. n., from a Frigate-bird (Fregata

aquila or F’. ariel).

Strobila 10 ems., excessively slender with large tulip-like heads.

Testes 17-20. Near to 7’. pelecanit Fuhrmann.

Proc. Zoou. Soc.—1914, No. XV. 15

226 ON HELMINTHES.

TETRABOTHRIUS NELSONI, sp. n., from Phebetria palpebrata.

Fragments only. Head absent. Testes 6 to 8, clumped at

opposite side of segment from the cirrus.

One Nematode is new and the type of a new genus :—

TERRANOVA ANTARCTICA, from Mustelus antarcticus.

A single Ascarid female 32 mm. long. Three squat fleshy lips

with paired anterior lobes. No labia intermedia. Cisophagus

without appendage. The intestine has a long cecum. The anus

lies at the base of a deep sulcus.

The following new genera are proposed :—

CrassrcaupA, gen. n., for Filaria crassicauda (Creplin), t. sp.

TmRRANOVA, gen. n., for Terranova antarctica, sp. n., t. sp.

An Ascarid with three large simple lips. No interlabia

(Esophagus simple. Gut with anterior cecal prolongation. No

cesophageal appendage.

KaTHLEENA, gen. n., for Ascaris osculata Rud., t. sp.

An Ascarid with three large fleshy lips and three interlabia.

(Esophagus has a solid appendage and the intestine has an anterior

cecal prolongation. In this genus may also be placed Ascaris

radiata, A. rectangula, and K. scotti.

OrIANA, gen. n., for Oriana wilsoni, sp. n., t. sp. (vide supra).

Cyclophyllid with large quadrate discoidal head carrying four

round suckers. Rostellum absent.

The types of the above forms are in the British Museum

(Natural History), London. An illustrated account of the col-

lection will appear in one of the zoological volumes of the results

of the Expedition, to be published by the Trustees of the

Museum.

March 3, 1914.

Prof. E. W. MacBripe, M.A., D.Sc., F.B.8., Vice-President,

in the Chair.

Mr. C. Tare Reean, M.A., F.Z.5., reported on the fresh-

water fishes from Dutch New Guinea collected by the British

Ornithologists’ Union and the Wollaston Expeditions. Symbran-

chus bengalensis was obtained for the first time in New Guinea.

The collections included examples of two species of Melanotzniine

Atherinids, and Mr. Regan had revised this group of fishes.

This paper will be published in the ‘ Transactions.’

No. 127.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON.*

February 3rd, 1914.

Sir Joun Rose BraprorD, K.C.M.G., D.Sc., F.R.S.,

Vice-President, in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Sucrerary read a Report on the Additions made to the

Society’s Menagerie during the months of N ovember and Decem-

ber, 1913.

Mr. D. Sera-Surrn, F.Z.8., Curator of Birds, exhibited a

photograph of two hybrids between a Peacock and a hen Guinea-

fowl which were bred in Germany and were now in the Berlin Zoo-

logical Gardens, and remarked that there were three or four cases

on record of such hybrids being produced, whereas there were

only two instances known of the Peafowl crossing with the

domestic fowl, and none, so far as he knew, with birds of any

other genus.

Mr. Seth-Smith also showed the skin of a hybrid Pheasant

hen, which had been sent him by Mr. H. J. Elwes, F.R.S. This

bird was one of a number bred in Sussex by Mrs. Johnstone,

between a cock Calophasis mikado and hen C. ellioti. It bore a

very strong resemblance to the female QO. mikado, but showed

a trace of the black throat and rufous barring to the tail of

C. elliott.

Mr. G. A. Boutencer, F.R.S., F.Z.S., gave an account of the

collections of Batrachians and Reptiles made by the British

Ornithologists’ Union and the Wollaston Expeditions in Dutch

New Guinea. Four species of Batrachians and eight species of

Reptiles were described as new.

This paper will be published in the ‘ Transactions’ in due

course.

* 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

‘+ 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.

Dr. F. E. Bepparp, M.A., F.R.S., F.Z.8., Prosector to the

Society, read a paper containing further observations upon the

Cestode genus Urocystidiwm Beddard, in which he described

more fully, from fresh material, the structure and life-history of

this genus of Tapeworms.

Mr. H. G. Purm™er, F.R.S., F.Z.8., Pathologist to the Society,

reported on the Deaths which had occurred in the Society’s

Gardens during 1913, and on tke Blood-parasites found during

the same period, and illustrated his communication with an in-

teresting series of lantern- slides. .

The next Meeting of the Society for Scientific Business will

be held on Tuesday, February 17th, 1914, at half-past Hight

o’clock #.m., when the following communications will be made :—

EXHIBITIONS AND NOTICES.

Dr ny AREY ADE LEIPER, M.B., Ch.B., E.Z. S., onl Siemezet ATKINSON,

Lantern demonstration of the Helminthes collected by

Scott’s Antarctic Expedition.

. SELIGMANN, M.B., M.R.C.P., E.Z.S8. ., and 8. _ SHATTOCK.

Observations made o ascertain iin any relation subsists

between the Seasonal assumption of the ‘‘ Eclipse” plumage in

the Mallard (Anas boscas) and the condition of the Testicle.

F. Woop Jonss, M.B., B.Sc., F.Z.8.

Some phases in the female Reproductive System of the Mole

(Talpa europea).

F. F. Larpnaw, M.A., F.Z.8.

Contributions to a Study of the Dragon-fly Fauna of Borneo.

—Part II. The Gomphine and Chlorogomphine.

H. C. Cuapwick, A.L.S.

Note on an imperfectly-developed Specimen of Hehinus

esculentus.

C. F. U. Mzex, M.8c., F.L.S., F.Z.8.

The Possible Connection between Spindle- ‘length | and Cell-

volume.

The following papers have been received :—

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

Further Contributions to the Anatomy of the Ophidia.

Witu1am Nicouu, M.A., D.Se., M.D., F.Z:S.

Trematode Pao from anor dying in the Zoological

Society’ s Gardens during 1911-1912. ;

F. J. Mraerrr, M.Sc.

The Structure and Life- History of a Tapeworm (Jchthyotenia

filicollus Rud.) parasitic in the Stickleback.

L. N. G. Ramsay, M.A., B.Sc.

1. On the Annelids of the Family Nereide collected by

Mr. F. A. Potts at Puget Sound, British Columbia, in 1911,

» with a Note on Micronereis as a Representative of the Ancestral

Type of the Nereide.

2. On the Genera Ceratocephale Malmgren and Tylor-

rhynchus Grube. 2

A. Kyyverr Torron.

The Structure and Development of the Caudal Skeleton of

‘the Teleostean Fish, Plewragramma antarcticum. ~

D. M.S. Watson, M.Sc., F.Z.8.

_ On the Skull of a Pariasaurian Reptile, and on the Relation-

ships of that Type.

H. R. Hoae, M.A., F.Z.8.

Spiders from the Montebello Islands.

Miss ALBERTINA CARLSSON. -

_ On Cynodictis CUES and Cynodon gr acilis from the.

‘Phosphorites of Quercy.

G. C. Rosson.

Report on the Mollusca collected by the British Ornitho-

logists’ Union Expedition and the Wollaston Expedition in

Dutch New Guinea.

HH. "WRewaine Kew, F.Z.S8.

On the Nests of Pecailcecenicnee with Historical Notes

on the Spinning-Organs and Observations on the Building and

Spinning of the Nests.

C. Tate Rzaean, M.A., F.Z.S.

Report on the Freshwater Fishes collected by the British

Ornithologists’ Union Expedition and the Wollaston Expedition

in Dutch New Guinea.

Prof. W. J. Dakin, D.Sc., F.L.S., F.Z.8.

Fauna of Western Australia.—l. The Onychophora of W.

Australia. II. The Phyllopoda of W. Australia.

Lt.-Col. J. M. Fawcert.

Description of Two new Species of African Sphingide.

GEORGE JENNISON.

Note on a hybrid Sea-Lion.

P. R. Awatt, B.A., C.I.C.

The Mechanism of Suction in the Potato Capsid Bug (Lygus

pabulinus Linn.).

EK. W. SHann, B.Sc.

On the Nature of the Lateral Muscle in Teleostei.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Socrery oF Lonpon,

ReGENtT’s Park, Lonpon, N.W

February 10th, 1914.

No. 128.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON."

February 17th, 1914.

Prof, EK. A. Mincuin, 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 made to the

Society’s Menagerie during the month of January, 1914.

Mr. E. G. Boutencer, F.Z.8., Curator of Reptiles, exhibited a

photograph of a female example of the Giant Saddle-backed

Tortoise (Zestudo abingdonit) recently purchased by the Society.

On the arrival of the tortoise, the origin of which was unknown,

Mr. Boulenger was somewhat puzzled as to the species to which

it should be referred, but, on carefully comparing it with the

Saddle-backed Tortoise in the British Museum, came to the

conclusion that the specimen was none other than the hitherto-

unknown female of 7. abingdonti, a species which had never

previously been brought to Europe alive, and which was thought

to be extinct. An inspection of the very representative collec-

tion of Giant Tortoises in the Tring Museum strengthened the

conclusion arrived at.

From the few male specimens in museums this Tortoise differs

by the fore-part of the shell being less strongly compressed and

not reaching the same height, more as in 7’. ephipiwm, by the

comparatively greater breadth of the hinder part of the carapace,

by the broader bridge, and by the smaller size of the marginals

bordering the bridge.

Mr. Boulenger also remarked upon some of the habits of this

species.

* 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 Stxpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance.

Mv, Grorcr JENNISON sent for exhibition a mounted specimen

of a hybrid Sea-Lion (Otaria pusilla g x O. californiana @),

which had been born in the Belle Vue Gardens, Manchester,

together with photographs of another specimen now living in the

same Gardens, and notes upon its habits.

Dr. R. T. Lerreer, F.Z.S8., and Surgeon HE. L. Atkinson, R.N.,

gave a lantern demonstration of the Helminthes collected by the

British Antarctic Expedition (Terra-Nova), 1910-1913. The

collection contained nine forms previously recorded from the

Antarctic Zone, three previously recorded only from the Arctic

Regions and one other previously recorded elsewhere and now

found in the Antarctic Zone, and fifteen new species and four

new genera. Of the forms obtained in Tropical and Temperate

Zones during the voyage, three had been recorded previously and

five were new species.

Messrs. CO. G. SreLiamMann, F.Z.S., and S. G. SHarrock com-

municated the results of some observations made to ascertain

whether any relation subsists between the seasonal assumption of

the “eclipse” plumage in the Mallard (Anas boscas) and the func-

tion of the testicle. The observations consisted in the micro-

scopic examination of the testicle each month of the year, and in

a study of the results of castration. The first observation showed

that the passage from the eclipse to the full winter plumage was

not associated with the advent of spermatogenesis. The Mallard

had passed out of eclipse by September, but at this date and till

early spring the spermatogenic function was in abeyance. Castra-

tion carried out during the eclipse (July and August) did not

delay the assumption of the winter plumage. If carried out in

December, however, the eclipse was delayed, though only for a

few weeks. This delay had its parallel in the well-established

fact that, if a colt is castrated when shedding its winter coat, the

shedding was for a time arrested, and then proceeded only slowly.

The complete castration of birds was a difficult procedure, even

under anesthetics and after free incision and inspection of the

abdominal organs. And in no instance in the case of the Mallard

was no testicular tissue reproduced, though the amount was quite

small.

Whilst it would appear that the seasonal change of plumage did

not correspond with the spermatogenic function of the testicle,

its connection with the production of an internal secretion could

only be settled by castration followed absolutely without regen-

eration; this could be ensured only by re-opening the abdomen

under an anesthetic and removing any reproduced tissue found.

Dr. F. Woop-Jones, F.Z.8., read a paper on some phases in

the reproductive history of the female Mole (Talpa europea). He

dealt with the nature-lore connected with the sexual evolution of

the Mole, the embryological development of the reproductive

system, seasonal changes in the external sexual characters of the

adult female, and the character of these changes as seen in

microscopical sections of embryos, nestlings, and adults.

Mr. H. C. Caapwick contributed some notes on an imperfectly

developed specimen of the Sea-Urchin (Hchinus esculentus). Two

well-marked depressions in the test attracted attention to the

living animal. Denuded of the spines the test presents the

appearance of a lump of plastic material which has been pinched

with thumb and forefinger. Two abnormalities appear in the

apical system, and abnormal plates occur in ambulacra II, III,

and IV. Inambulacrum III two plates of normal composition

are exceptional, in that the pore-pairs of their respective demi-

plates are almost completely outside the limits of the plate, and

perforate the adjoining interambulacral plate. Similar irregu-

larities, though not so well marked, occur in ambulacrum IV.

Mr. C. F. U. Meex, M.Sc., F.Z.S., communicated a paper

entitled “The Possible Connection between Spindle-length and

Cell-volume.” In Sorficula auricularia, Helix pomatia, and Man

the ratio between the lengths of the mitotic spindle in the two

spermatocyte metaphases seemed to be identical or almost identical

with the ratio beween the radii of two spheres, of which the

volume of one is equal to twice that of the other ; and, since the

volume of the primary spermatocyte cell in the metaphase is

presumably equal to twice that of the secondary spermatocyte,

connection was suggested between the spindle-length and cell-

volume at this stage. Photo-micrographs are now given, showing

this ratio in the two first-named organisms; but, in the case of

Forficula, new preparations have been observed to contain primary

spermatocyte spindles of excessive lengths, and unless a satisfactory

explanation can be found for these, the original suggestion must be

either modified or discarded.

The study of spermatocyte cells in Man, Triton cristatus,

Stenobothrus viridulus, S. curtipennis, Forficula awricularia, and

Helix pomatia proves that the volume may be similar in widely-

separated organisms, and very different in organisms that are

closely allied. In the circumstances, increasing somatic com-

plexity of the organism is not necessarily accompanied by increase

of the volumes of these cells.

A paper received from Mr. F. F. Larptaw, M.A., F.Z.S., con-

tained a further contribution to the study of the Dragon-fly

fauna of Borneo, and dealt with the Gomphine and Chloro-

gomphinz, of which a number of new species and subspecies was

described.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, March 3rd, 1914, at half-past Hight

o'clock P.m., when the following communications will be made :—

EXHIBITIONS AND NoTIcEs.

————

C. Tare Recan, M.A., F.Z.S.

= ee

Report on the Freshwater Fishes collected by the British

Ornithologists’ Union Expedition and the Wollaston Expedition

in Dutch New Guinea.

H. Wauus Kew, F.Z.S.

On the Nests of Pseudoscorpiones; with Historical Notes

on the Spinning-Organs and Observations on the Building and

Spinning of the Nests.

H. R. Hoae, M.A., F.Z.8.

Spiders from the Montebello Islands.

D. M. 8. Watson, M.Sc., F.Z.8.

On the Skull of a Pariasaurian Reptile, and on the Relation-

ships of that Type.

F. J. Mecerrr, M.Sc.

The Structure and Life-History of a Tapeworm (Lchthyotenia

jilicollis Rud.) parasitic in the Stickleback.

Wituram Nicott, M.A., D.Se., M.D., F.Z.S.

Trematode Parasites from Animals dying in the Zoological

Society’s Gardens during 1911-1912.

The following papers have been received :—

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

Are pate eee No lanes SD

Further Contributions to the Anatomy of the Ophidia.

L. N. G. Ramsay, M.A., B.Sc.

1. On the Annelids of the Family Nereide collected by

Mr. F. A. Potts at Puget Sound, British Columbia, in 1911,

witha Note on Micronereis asa Representative of the Ancestral

Type of the Nereide.

2. On the Genera Ceratocephale Malmgren and Tylor-

rhynchus Grube.

A. Knyvett Totton.

Soc ee

The Structure and Development of the Caudal Skeleton of

the Teleostean Fish, Plewragramma antarcticum.

Miss ALBERTINA CARLSSON.

ieee eee

On the Fossil Carnivores Oynodictis intermedius and Cynodon

gracilis from the Phosphorites of Quercy.

G. C. Rosson.

Report on the Mollusca collected by the British Ornitho-

logists’ Union Expedition and the Wollaston Expedition in

Dutch New Guinea.

Prof. W. J. Daxty, D.Sc., F.L.S., F.Z.8.

Fauna of Western Australia.—tI. The Onychophora of W.

Australia. II. The Phyllopoda of W. Australia.

P. R. Awatt, B.A., C.LC.

The Mechanism of Suction in the Potato Capsid Bug (Lygus

pabulinus Linn.).

KE. W. Susann, B.Sc.

On the Nature of the Lateral Muscle in Teleostei.

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

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.— XIII. On Two new Species belonging to the

Genera Oochoristica and Linstowia, with Remarks upon those

Genera.

Guy DouLMAN.

SANs ei pep Sisto eee

Notes on a Collection of East African Mammals presented

to the British Museum by Mr. G. P. Cosens.

W. T. Catan, D.Sc., F.Z.8.

Report on the River-Crabs (Potamonide) collected by the

British Ornithologists’ Union Expedition and the Wollaston

Expedition in Dutch New Guinea.

OnpFIELD Tuomas, F.R.S., F.Z.S.

Report on the Mammals collected by the British Orni-

thologists’ Union Expedition and the Wollaston Expedition in

Dutch New Guinea.

Communications intended for the Scientific Meetings should

be addressed to

P, CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL SOCIETY OF LONDON,

Recent’s Park, Lonpon, N.W.

February 24th, 1914.

: bape.

ae:

aarp aay

ss sili moyen

No. 129.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

March 3rd, 1914.

Prof. E. W. MacBripz, M.A., D.Sc., F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. C. Tare Recay, M.A., F.Z.S., reported on the fresh-

water fishes from Dutch New Guinea collected by the British

Ornithologists’ Union and Wollaston Expeditions. Symbranchus

bengalensis was obtained for the first time in New Guinea.

The collections included examples of two species of Melanoteeniine

Atherinids, and Mr. Regan had revised this group of fishes.

This paper will be published in the ‘Transactions.’

Mr. H. Watuis Kew, F.Z.S., contributed a paper ‘On the

Nests of Pseudoscorpiones: with Historical Notes on the Spinning-

Organs and Observations on the Building and Spinning of the

Nests.” The paper described the nests in which these animals

enclose themselves for moulting, for brood-purposes, and in some

cases for hibernation. ‘They are closed cells of spun tissue, with

or without a covering of earthy or vegetable matters. ‘The tissue

is of innumerable threads crossed and coalesced irregularly,

without interspaces, and almost like silk-paper. With regard to

the spinning-apparatus, confusion has existed ; but the Author’s

observations on living animals place it beyond doubt that the

cephalothoracic glands, whose ducts traverse the chelicerz to near

the apex of the movable finger and open in the galea, or in 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 Siapence, or, if desired, sent post-free for

the sum of Sia Shillings per annum, payable in advance.

tubercle which replaces it in some groups, are the organs con-

cerned. Contrary to previous statements, the “combs” of the

chelicerze have nothing to do with the silk. The manner in

which the nests are built and spun was described in detail. The

spinning was associated with forward and backward movements

of the body and with lateral movements of the chelicere ; and

the silk issued from the galea or tubercle in several viscid, very

fine threads. The methods of three species, representing both

main divisions of Pseudoscorpiones, were observed; and, being

essentially identical, were believed to be characteristic of the

order.

Mr. H. R. Hoae, M.A., F.Z.8., read a paper on a collection of

Spiders made by Mr. P. D. Montague, of Caius College, Cambridge,

supplemented by a few specimens sent by Mr. T. H. Haynes

from the Montebello Islands off Onslow, on the N.W. Coast of

Australia.

These islands, from geological evidence, were part of the old

coast-line, though now about 90 miles away. Although the

larger specimens are mostly widely spread and possibly more or

less recent importations, the smaller are nearly all new species,

showing evidence of a much longer separation from their con-

generic relations on the mainland.

Out of seventeen species ten are new, as well as a new genus

and two new varieties.

Mr. D. M.S. Watson, M.Se., F.Z.S., gave an account of his

paper on “ The Skull of a Pariasaurian Reptile and the Relation-

ships of that Type.” The skull of Pariasaurus is completely

described therein, with the exception of the bony labyrinth of

the ear. It is compared with all the members of the Order

Cotylosauria which are well enough known to make a comparison

of any value, and shown to differ in the very important characters

of the brain-case from all of them, representing an entirely

distinct branch.

The characters of the various orders of the superorder Thera-

psida are then detailed and the characteristics of the superorder

deduced from their comparison ; Pariasaurus is shown to possess

none of them, and hence to be quite unconnected with their

ancestry.

A sphenethmoid is recognised in Pariasaurus for the first time

in any reptile, and the homologies and distinguishing features of

the single temporal bone, the squamosal, are discussed.

Mr. F. J. Mecairr, M.Sc., contributed a paper, communicated

by Prof. F. W. Gamste, F.R.8., F.Z.8., containing a description of

a Tapeworm parasitic in the Stickleback (Gasterosteus aculeatus),

with an account of its anatomy and histology, and its life-

history.

Dr. Witti1am Nicouu, M.A., D.Sc., F.Z.S., sent a paper dealing

with the Trematode Parasites obtained from animals that had died

in the Society’s Gardens during 1911-912, in which he described

four new genera and eleven new species.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, March 17th, 1914, at half-past Hight

o'clock P.m., when the following communications will be made :—

EXHIBITIONS AND NoTIcEs.

L. N. G. Ramsay, M.A., B.Sc.

1. On the Annelids of the Family Nereide collected by

Mr. F. A. Potts in the N.E. Pacific in 1911, with a Note on

Micronereis a8 a Representative of the Ancestral Type of the

Nereide.

2. On the Genera Ceratocephale Malmgren and Tylor-

rhynchus Grube,

A. Kwyyvert Torron.

The Structure and Development of the Caudal Skeleton of

the Teleostean Fish, Plewragramma antarcticus.

G. C. Rosson.

Report on the Mollusca collected by the British Ornitho-

logists’ Union Expedition and the Wollaston Expedition in

Dutch New Guinea.

P. R. Awart, B.A., D.I.C.

The Mechanism of Suction in the Potato Capsid Bug (Lygus

pabulinus Linn.),

K. G. Buatr, B.Sc., F.E.S.

Coleoptera Heteromera collected by the British Ornithologists’

Union Expedition and the Wollaston Expedition in Dutch. New

Guinea,

R. Lypexx«eEr, F.R.S., F.Z.S.

The Malay Race of the Indian Elephant.

Prof. W. J. Dakin, D.Sc., F.L.S., F.Z.8.

Fauna of Western Australia.—I. The Onychophora of W.

Australia. II, The Phyllopoda of W. Australia.

The following papers have been received :—

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

a a ad

Further Contributions to the Anatomy of the Ophidia.

Miss ALBERTINA CARLSSON.

at Ps

On the Fossil Carnivores Cynodietis intermedius and Cynodou

gracilis from the Phosphorites of Quercy.

KE. W. Suany, B.Sc.

On the Nature of the Lateral Muscle in Teleostei.

F. E. Bepparp, M.A., D.Sc., F.R.S., F.Z.S.

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.— XIII. On Two new Species belonging to the

Genera Oochoristica and Linstowia, with Remarks upon those

Genera.

Guy DoLiMAN.

Gon

Notes on a Collection of East African Mammals presented

to the British Museum by Mr. G. P. Cosens.

W. T. Catan, D.Sc., F.Z.S.

Report on the River-Crabs (Potamonidz) collected by the

British Ornithologists’ Union Expedition and the Wollaston

Expedition in Dutch New Guinea.

OLDFIELD THomas, F.R.S., F.Z.S.

Report on the Mammals collected by the British Orni-

thologists’ Union Expedition and the Wollaston Expedition in

Dutch New Guinea.

H. A. BAYtis.

Report on the Nematodes and Gordiacea collected by the

British Ornithologists’ Union Expedition and the Wollaston

Expedition in Dutch New Guinea.

The Rev. T. R. R. Stespine, M.A., F.R.S., F.L.S., F.Z.S.

Crustacea from the Falkland Islands collected by Mr. Rupert

Vallentin, F.L.S.—Part 2.

Communications intended for the Scientific Meetings should

be addressed to

P, CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpon,

Recent’s Park, Lonpon, N.W.

March 10th, 1914.

Pi As AL awk a

iG OE aot

Papers (continued).

Page

. Observations made to ascertain whetber any Relation subsists between the Seasonal

Assumption of the “Eclipse” Plumage in the Mallard (Anas boscas) and the

Functions of the Testicle. C. G. Suriemany, F.Z.8., and 8. G. Suarrock. (Text-

Frepuric Woop-Jonus, D.Sec., F.Z.8. (Plates I,-I1I., and Vext-figures 1-15.) ......

figures 1-6.) ° O20 2020000 Pee ee se se ses ere soseses0e eso 2520298 29 8 23

3. The Possible Connection between Spindle-Length and Oell-Volume. By C.F. U. Muzx,

Mises -E.:8:, 1.2.8. ~ (Plates TP. 6 VE) wees vs lors on vice Foddne CE cohOko pA amo oO GOeS 45

4, Contributions to a Study of the Dragonfly Fauna of Borneo.—Part IJ. The Gomphinz

and. Chlorogommphine. By F. F. Laipuaw, M.A., F.Z.8. (Plate I.) ............ ol

5. Note on an imperfectly developed specimen of the Sea-Urchin (Hehinus esculentus). By

H. C. Cuapwicr, A.L.S. (Text-figures 1-4.)...... Se ae SARA MRI Sires NT or a 65

6. Spiders from the Montebello Islands, ‘ By H. R. Hoaa, M.A.,F.Z.8. (Plates I-III). 69 :

7. On the Nests of Pseudoscorpiones: with Historical Notes on the Spinning-Organs and

Observations on the Building and Spinning of the Nests. By H. Wauuis Knew, F.Z.8.. 93

8, The Structure and Life-History of a Tapeworm (Ichthyotenia filicollis Rud.) Parasitic

in the Stickleback. By F. J. Muaeitr, M.Sc. (Birm.), Board of Agriculture and

Fisheries Research Scholar, Eleni of Birmingham. (Plates I.-IV., and Text-

figures 1-5.).......... RGca ROO OUOS HOO COD GOO TU cU AMOO SCOR Co Kt Ob GOe rein co 113

9. Trematode Parasites from Animals dying in the Zoological Society’s Gardens during

1911-1912, By Wiux1am Nicoxt, M.A., D.Sc., M.D., F.Z.8. (Plates I-IV.) noe. JB!)

10. On the Skull of a Pariasaurian Reptile, and on the Relationship of that Type. By

D. M. S. Watson, M.Sc., F.Z.8., Lecturer on Vertebrate a esa gy) in University

College, London. @aceeae Tf) tion a eaoc AG SIS eS eM Rei cer aASa ras Amare ns 155

11. Report on the Deaths which occurred in the Zoological Gardens during 1918, together

f with a List of the Blood-Parasites found during the Year. By H.G, Privmar,l.R.S., .

Mie pk Tut OLOIBt ee O) PME SOCIOL) vier a) a/ctelin'elm =) sia Sic. ¢atclevanicleis cs s'las stelmjetsla = ojnc se eels 181

12. Some Phases in the Reproductive History of the Female Mole (Zalpa europea). By

191

ist

Menk:

DLarpraw: $

Hosa: re

Mecert:

‘Nico:

JONES:

The ‘ Been dines for the year are issued in four

80 that the complete reference i is now P. Z. 8. ‘1914, 2 ieee

is as follows :—

Pl Ly

LIST OF PLATES,

1914, Parr I. (pp. 1-226).

Pl. I.

II.

Pl. I. Bornean Dragonflies de eae DE CE ge

\ eee ise Cells in various Organisms ee

ED.

tv.)

Pe i ae

ay Spiders from the Montebello Islands oer esenrecs

USteucnne of Ichthyotenia filicollis....-..

| : SE »

a ronal Parasites .. “t setter esters cease ae ws

Ty.) se Pees iss

Pl) Biers Gesu

iil WN cikeenal Genitalia of the Bees Re See arate eign

mL)

NOTICE.

‘ Part LL. issued in Maret, ai a = d

ads June.

32 ”

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCLETY

OF LONDON.

1914.

PART II. 7.

CONTAINING PacEs 227 to 490, wiTH 26 PLATES

‘AnD 47 TEXT-FIGURES.

JUNE 1914.

- PRINTED FOR THE SOCIETY,

_ SOLD AT ITS HOUSE IN REGENT’S PARK,

LONDON :

MESSRS. LONGMANS, GREEN, AND CO.,

PATERNOSTER ROW.

ie :

ig Mele

tf J <d if t

‘aay | Se ee ee

[Price Twelve Shiliangs.|

LIST OF CONTENTS.

1914, Parr IL. (pp. 227-490).

EXHIBITIONS AND NOTICES.

The Sucrzrary, Report on the Additions to the Society’s Menagerie during the month

Ot Melina WOM (aa ea oo oa auadsgbU use GoU0ONcocOUdcdob odo bE Udo SSO eeu seas

Mr. G. C. Rogson, B. A. Notice of Report on Mollusca from Dutch New Guinea......

Mr. K. G. Buair, B. Pe. Notice of Report on Heteromerous Coleoptera from Dutch New

(Glia enies ae NOE SAO on oO OUD COME oar tad 2 COMO COUBEOOAGU DUOC GOs onc a

Mr, R. H. Burns, M. A., F.Z.S. Palatal growth in mouth of Camel. (Text-fig. 1.) ....

- Mr. R. I. Pocooxs, F.R.S., F.Z.S8. On the Feet of Domestic Dogs. (Text-figs. 1-3.) ....

The Sucrutary. Exhibition of the photograph of a female Orang-utan (Simia satyrus) -.

The Sucrutary. Exhibition of photographs of large-tailed Punjab Domestic Sheep ....

Dr. W. T. Cauman, F.Z.8S. Notice of Report on River-Crabs (Potamonidz) from Dutch

UNG ws Giimear cb sect stant cerca a enero guise eee tere we ole taiallehanaie alan eta eteievaite Sie( sds Se ie Een ote ie eaRD

Mr, Oxpriexp ‘nomas, F.R.S., F.Z.S. Notice of Report on Mammals from Dutch New

(EN ihaC rine aren aera Arie SS Cert ee entrar A i erm ISS QC OOS jc G2 6

The Srcrerary. Report on the Additions to the Society’s Menagerie during the month

Ot Wire NM Soaks nausea dopo noaanmbo os a0 Co oo so oKnd sah S00GDOCoD ODO NOSSO:

The Srcretary. Notice of change in time of Scientific Meetings ....cessceseueseceee

Mr, D. Szra-Surra, F.ZS. Hxhibition of egg of Mantell’s Kiwi (Apteryx mantelle) i

in the Society’s Gardens ...- 62.020. e ce cc ee ce sew e et eee ences crac cscteneneacs .

Mr, Srantuy Hirst, F.Z.S. Notice of Report on Arachnida (other than eee) aH

Myriopoda from Dutch New Guinea... .<.scsers icet ots ots ee

Surgeon G. Murray Levicn, R.N. Notice of Lecture on the Manners and Customs of

Ndéhekbenguins| CReygoscelis Gdelve)\ aera tale ele teleleate w\<) she tale ale <eieisicle)aleisioperersi ait teenetetets

Sir Epmunp G. Lopmr, Bt., F.Z.S. Exhibition of Antlers of Red Deer (Gorvns elaphus) ..

The Secretary. Report on the Additions to the Society’s Menagerie during the month

of April 1914......... EeMere internals Soalstane enueae's Gane vinta te ala oP cnpera ite weenie

Mrs. R. Hate THOMAS, F.Z.S. Exhibition of Skulls of Hornless Antelopes ...

Mr. D. M. 8. Watson, M.Sc., F.Z.S. Exhibition of Procolophon eee a Cotyla:

saurian Reptile ...... a usletteieielevectcaniereieate rataeneiee- pe ieietelec ras ejellesofeiel netateiers tenets UAGOS

PAPERS.

13. On the Fossil Oarnivores Cynodictis intermedius and Cynodon gracilis from the Phospho-

rites of Quercy. By AuBertIna Carusson, Zootomical Institute, University of

Stockholm. (Plate 1.)...........eeee: NE GURGTCO On odo NOU nDOaEOG og cO ou oo Cod-

14. On the Genera Ceratocephale Malmgren and Tylorhynchus ay By L. N. a.

- Ramsay, M.A., B.Sc, Carnegie Research Scholar, Christ’s College, Cambridge ......

15. On the ee of the Family Nereidx collected by Mr. F. A. Potts in the N.E.

Pacific, in 1911. With a Note on the Morphology of Micronerets as a Representative

of the Ancestral Type of the Nereida. By L. N. G. Ramsay, M.A., B.Se., Carnegie

Research Scholar, Christ’s College, Cambr wae) (Text-figures 1-7.) ..... aku

Contents continued on page 3 of Wrapper. :

Page

{

‘ 4

Se ee td can ee eR ee tet

227 ‘ “4

231

ZOOLOGICAL SOCIETY OF LONDON.

Tarts Society was founded in 1826 by Sir Sramrorp Rarruzs,

Mr. J. Sastne, Mr. N. A. Vigors, and other eminent Naturalists,

for the advancement of Zoology and Animal Physiology, and for the

introduction of new and curious subjects of the Animal Kingdom,

and was incorporated by Royal Charter in 1829.

Patron.

HIS MAJESTY THE KING.

COUNCIL.

HIS GRACE THE DUKE OF BEDFORD, K.G., F.RS, President.

Ricnarp H. Burne, Esa., M.A., || E. G. B. Mravr-Watpo, Esa.

Vice-President. | Prorrsson Epwarp A. Mixcutn,

Aurrep H. Cocks, Ese., M.A. | M.A., F.R.S., Vice-President.

Tar Rr. Hoy. Tar Earn or P. CHatmers Mircnetr, Ese.,

Cromer, P.C., G.C.B., Nee) IDASeh IbIbID.. IIR,

GeMG.; K-CSI, F.RS.,

Vice-President.

)

Secretary.

W. R. Ocitvre-Grant, Ese.

F. G. Dawrrey Drewirr, Ese.,

M.A., M.D. | ALBerT Pam, Ese.

Cuartes Druamsonp, Ese., Aprran D. W. Pottock, Esa.

ee: | Sre Ronarp Ross, K.C.B., F.RS.

F. Dy Canz Gopman, Ese.,D.C.L., |

ERS. | Tue Marevgss or Srico, F.S.A.

d 1

Str Watrer Rover Lawrence, | OLprietp Tuomas, Ese., F.R.S.

Br., Gere Ene Vice-President. ANTHONY H. WINGFIELD, Ksa.

_ Ervest W. MacBripz, Ese., Heyry Woopwarp, Ese., LL.D.

a eI) isu iaeePiesident.

President. |

The Society consists of Fellows, and Honorary, Foreign, and

Corresponding Members, elected according to the By-Laws. It

carries out the objects of its foundation by means of the collection

of living animals, by its Library, and by its Scientific Publications.

The Office of the Society, Regent’s Park, N.W., where all com-

munications should be sent, addressed to ‘The Secretary,” is open

from Ten till Five, except on Saturdays, when it closes at One P.M.

The Library, under the superintendence of Mr. Henry G. J. Peavot,

is open daily (except Sunday) from Ten a.m. till Five p.m.; on

Saturdays, Ten a.m. till Two p.m.

The Library is closed from Good Friday to Easter Monday, and

upon all other Bank Holidays. It is also closed annually for

cleaning purposes during the whole month of September.

The Meetings of the Society for General Business are held in the

Meeting Room at the Society’s Office on the third Wednesday in

every month of the year, except in September and October, at half-

past Four o’clock p.m.

The Meetings for Scientific Business are held in the Meeting

Room at the Society’s Office fortnightly on Tuesdays, except in

July, August, September, and December and January, at half-past

Five o'clock p.m.

The Anniversary Meeting is held on the 29th. of April, or the

nearest convenient day, at Four p.m.

The Society's Gardens are open daily from Nine o’clock until

Sunset. Mr. R. I. Pocock, F.R.S., F.LS., is the resident Super-

intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator

of Birds and Inspector of Works, Mr. E. G. Boulenger is Curator

of Reptiles, and Prof. H. M. Lefroy is Curator of Insects.

The Prosectorium for Anatomical and Pathological work is under

the charge of Mr. Frank E. Beddard, M.A., D.Sc., F.R.S., Prosector,

assisted by Mr. H. G. Phimmer, F.R.S., M-R.C.S., Pathologist to

the Society.

TERMS FOR THE ADMISSION OF FELLOWS.

Fetrows pay an Admission Fee of £5, and an Annual Contri-

bution of £3, due on the Ist. of January, and payable in advance,

or a Composition of £45 in lieu thereof; the whole payment,

including the Admission Fee, being £50.

No person can become a Frrtow until the Admission Fee and

first Annual Subscription have been paid, or the annual payments

have been compounded for.

Fetztows elected in November and December are not liable for

the Subscription for the vear in which they are elected.

[S)

PRIVILEGES OF FELLOWS.

Frttows have Personal Admission to the Gardens upon signing

their names in the book at the entrance gate, and may introduce

Two Companions daily.

The Wire or Huspanp of a Fettow can exercise these privileges

in the absence of the Fellow.

Until further notice, Frttows will receive 40 undated Green

Cards, available on any Sunday or week-day up to the end of

February of the year following the year of issue, and 20 White

Cards available on any week-day up to the same date. Twenty

of the Green Cards may be exchanged for a book containing two

Orders for each Sunday in the year. Twenty White Cards may

be exchanged for a book of dated Saturday Orders, each Order

available for any day except Sunday in the week ending with the

Saturday for which it is dated. Special children’s tickets will uo

longer be issued, but the Green and White Cards will be perforated,

and each half will be valid for a Child under twelve years of age.

It is particularly requested that Fellows will sign every ticket before

it goes out of their possession. Unsigned tickets are not available.

Frttows are not allowed to pass in friends on their written

order or on presentation of their visiting cards.

Frtiows have the privilege of receiving the Society’s ordinary

Publications issued during the year upon payment of an additional

Subseription of One Guinea. This Subscription is due upon the

1st. of January, and must be paid before the day of the Anniversary

Meeting, after which the privilege lapses. Fxrttows are likewise

entitled to purchase these Publications at 25 per cent. less than

the price charged to the public. A further reduction of 25 per

cent. is also made upon all purchases of Publications issued prior

to 1881, if above the value of Five Pounds.

Frttows also have the privilege of subscribing to the Annual

Volume of ‘ The Zoological Record,’ which gives a list of the Works

and Publications relating to Zoology in each year, for the sum of

One Pound Ten Shillings. Separate divisions of volumes 39 to

42 can also be supplied. Full particulars of these publications can

be had on application to the Secretary.

Frettows may obtain a TRANSFERABLE Ivory Ticker admitting

two persons, available throughout the whole period of Fellowship,

on payment of Ten Pounds in one sum. A second similar ticket

may be obtained on payment of a further sum of Twenty Pounds.

Any Fetiow who intends to be absent from the United Kingdom

during the space of at least one year, may, upon giving to the

Secretary notice in writing, have his or her name placed upon the

“dormant list,” and will then be called upon to pay an annual

subscription of £1 only during such absence, but after three years:

must make a further application to be retained on that list.

Any Fextow, having paid all fees due to the Society, is at liberty

to withdraw his or her name upon giving notice in writing to the

Secretary.

Ladies or Gentlemen wishing to become Fellows of the Society

are requested to communicate with “The Secretary.”

P, CHALMERS MITCHELL,

Secretary.

Regent’s Park, London, N.W.

June, 1914.

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

FOR

SCIENTIFIC BUSINESS.

TupspaY, OcTOBER ...... 27th.

NovempBer .... 10th.

bh)

” o)e)

The Chair will be taken at half-past Five o clock precisely.

LOOESGVWW SE SOCIETY -OF LONDON:

LIST OF PUBLICATIONS.

Tue scientific publications of the Zoological Society of London

are of two kinds—“ Proceedings,” published in an octavo

form, and “ Transactions,” in quarto.

According to the present arrangements, the “ Proceedings”’

contain not only notices of all business transacted at the scien-

tific meetings, but also all the papers read at such meetings

and recommended to be published in the “‘ Proceedings ”’ by

the Committee of Publication. A large number of coloured

plates and engravings are issued in the volumes of the

“ Proceedings,” to illustrate the new or otherwise remark-

able species of animals described therein. Amongst such

illustrations, figures of the new or rare species acquired in a

living state for the Society’s Gardens are often given.

The “ Proceedings” for each year are issued in four parts,

paged consecutively, during the months of March, June,

September, and December. From January 1901 they have

been issued as two half-yearly volumes, indexed separately.

An “ Abstract of the Proceedings” is published by the

Society on the Tuesday followimg the date of the Scientific

Meeting to which it refers. It is issued along with the “ Pro-

ceedings,” free of extra charge, to all Fellows who subscribe to

the Publications, but it may be cbtaimed on the day of publi-

cation at the price of Sixpence, or, if desired, sent post free

for the sum of Six Shillings per annum, payable in advance.

The ‘‘ Transactions” contain such of the communications

made to the Scientific Meetings of the Society as, on account of

the nature of the plates required toillustrate them, are better

adapted for publication in the quarto form. They are issued

at irregular intervals.

Fellows and Corresponding Members, upon payment of

a Subscription of One Guinea before the day of the Anni-

versary Meeting, are entitled to receive the Society’s

Publications for the year. They are likewise entitled to

purchase the Publications of the Society at 25 per cent. less

than the price charged to the Public. A further reduction

of 25 per cent. is made upon purchases of Publications

issued prior to 1881, if they exceed the value of Five

Pounds.

Fellows also have the privilege of subscribing to the

Zoological Record for a sum of One Pound Ten Shillings

(which includes cost of delivery), payable on the Ist. of July

in each year; but this privilege is forfeited unless the

subscription be paid before the 1st. of December following.

The following is a complete list of the publications of the

Society already issued.

TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 19 vols. and Index. Price to Price to the

Fellows. Public.

Vols. L.—-LYV. (out of print).

Wolke We a CO oon CIGRHGS) geoe on ZS obo. G1 ©

rane ibs. i" O27 ya) ee ChS66269), 0... UL 25. 0) yo Ome

pee Wil Li . 13 9 nat LeOo=g) .... 10, 4:0 eae

Pee walliles 5 82) eect Creel Sioa) . 2.5 "9 $8 (Sek Sewn UG

cope Oe < QO OORT). 3... 12 TONG aa Geo mme

Xe s BD oy p500 Cheat) hoe MOOS oo44 18 7 ©

Index, Wols. Xe arrrck pero eine @s33—79) i250) 7 6) aa OO RG

Vol, Xi containme 9 lates ya (lss0-85))_. 5. 9:12 0 Pes I2aIGeG

¥ DCG eno amen (SSO—OON une. O ) eo mau, i & ©

se Alle Oo Mn mee ML OOU-OD)) wsnee- Os GOO. 811 0

AO GIN eo pee eee SOO—IS) heres ko) sole “ © ©

i OWS x OE ese as MORES SIO ig 5 a ey Gl 714 0

Bi NAVI: 8S) ong oo ICOISIOOS) oa Sy SO", “ & @

SOL, Py A ne oe, (CIOS AI0G) eae lS) (Ol mean. aégelommey

oy oN WAU fy AS Rae oes (OUR eI OMI pee OSG) ako 1) mmm)

PaO Xe. ee ao (LOD SIND) 2, IO Ze Octso, IB 12 ©

" NO ieart i, (els a. (heb: 192) Owls Ocoee Loa ©

se NOKXe == 1/5) 25 1(Pisy Vi XOV a) g(Avpril O22 Oe an OME)

se SOX == oy Be CARDCVIEOONITE UMein: GIS). I BE. 110 O

a SOC ae GES CONN ESOL) (tae 9 0. 12 ©

a XOX, 5. (Rls) XONV 1 Ox) (March 1914) 90): 12 ©

se OCS) , (Rb SOONIE) (Gilera UGAa) 3 © . en)

*, XOX See i | (p ESeXUXOXGIATI SXOXGXCTITS) (Miers). Gi 10 O

ie DROXG =e Bau (Manvel OA) Ga rset. soctonspone woolen Oe aie 3. 0

a DOKG ag Ore (Maye ONA) i wine ke ag Mahe, 3 0

ss PONG er Opa NOMA i tare e ia scaslarseh ame Oat 3 0

now able to offer for sale, at the reduced price of £30, sets of Vols. V.—XVI. inclusive, and

In consequence of a re-arrangement of the stock of the ‘Transactions,’ the Society is

separate papers, of which a list can be supplied, at about one-fourth their published price.

PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND

CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF

LONDON. 8vo. 2 vols. (Letterpress only). Erice te Price cote

Fellows. Publie.

Part I. 1830-31. 1 vol. 8vo., out of print.

4 iil, EB ite ae nee AsvGd; tien

PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.

First SERIEs.

Parts 1—XV. (1833-1847). 8vo. 15 vols. (Letterpress only.) Price to

Fellows : 4s. 6d. each part ; to the Public, 6s.

Index 1830-1847. Price to Fellows: 4s. 6d.: to the Public, 6s.

Parts I., VII.-IX., XI, XIV., XV., out of print.

SECOND SERIES.

Parts XVI.-XXVIII. (1848-1860). 8vo. 13 vols. (Letterpress only.)

Price to Fellows: 4s. 6d. each part ; to the Public, 6s.

Index 1848-1860. Price to Feliows: 4s. 6d.; to the Public, 6s.

The Parts of this series containing Coloured Plates are out of print.

PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE

ZOOLOGICAL SOCIETY OF LONDON. 8vo. 40 vols. and 4 Indices.

Letterpress only. With Plates uncoloured. With Plates coloured.

Price to Price to the Price to Price to the Price to Price to the

Fellows. Public. Fellows. Publie. Fellows. Publie.

MS GilGe ASE Gia OSsa ees os 9s. Ee ane) DK o = Bea oss. 9d: .... 40s

NOY 56 4 GE bio no (8 “Sinema een Ae AO SRE ree Boe, Behe sooc 45s.

ISGB 45 4 Oth) ob06 OS clo oene 9s, Hos ai O Sub Tan ae Bae, See soon FRE

SCARS Ash Gd) ao OSehae oss 9s. sparen BOSAL ae ocr aos, 94. .... 45s.*

1365 5, 8a Gi, boon CSilooosoc 9s. LOSE Se SSS) Ean a ineeemaose

USGS oo 485 Ch ohbe0 Cio oosne Qs. ooo NESS eos oe om Gh shag) Zine

USC cit een ces ea eee 9s. S60 ees oso Soke OU eo ao 2am

GAS. 5 6 Sic Re ee ee 9s. ee SH ee OOS HOt Re AOS:

GD G56 oO REE De EO oa 9s RD sree eee Ba Wh ooco 45S

SS OMe eis Sec ees ake tSR ea iene 9s gD Es ae are te dos. 9d. .... 45s

indexes SGleNS7Or er 155 ake. S. ASiGdameos 6s.

TAMA Matric Cae hae rae SNC eee 9s TO SENS wee SOS) Ids see LOS

GD SS eit Saar teen eae Net Ys, NOG PS 3's 6 bos. Od. .... 45s8.F

SDs 35 3 ps com GeO RR ee eran eee 9s Ne, 26 5 ee Bh soon ais

WG AL 5. 25 eat See ce nee lea tae 9s Ness a 36s 48s.

ISDS hese tie Gea ee Ree ae rer eee 9s a HDS ae 3 36s 48s

LST. 5-8 da cea Rea ee ee ee ee 9s 5 Ds ree aera 36s 485.7

STM Tatler es cicres data eens 9s tri 36s 48s

JUSS; GaGa ROE OR ee Ear ame 9s AU GH Beale 36s 48s

WSTO): 3 SS ae HR er eae area seein ae 9s. Wt EDS eye OOS Aunts:

SS Opes sec cence ce es ase 9s. Bea eile) San eee 36s, Bey eaet koky,

linden: SIS MEV SSOR Ne sts! AS2 Odo an yn OS:

HSS I Sareea rem yeh ra ere Sie Rien 9s. seis LDS see ae AA OOS! SASS:

SSD Meme yee egies ey ler nee ie 9s NAS eesin s (Blo sooo. aes

NSB S55 SaSate Bet Ree eee ee Ores 9s Sliss 36s 48s

TSCXSML, ~ 6-3. Sasa ine an Rena NO Re 9s , 2s. 36s 48s

TICKS): 5 Gcaseen Hae eae eae ne 9s ls! 36s 48s

TRAGER Ne ta RRA ee 9s 5 19s. 36s 48s

RSS MAR sen Petes sstsce sala iea eaves 9s 5 25. 36s 48s

ISS Sm Ate set hs on cane t am tases 9s » 2s: 36s 48s

TICKERS) 5 vchaes Oe SiGe te Se eae ie eae 9s she tesla 36s 48s

LSD. 5 SYS Ge ee 9s OSU ate 36s 48s

Inidex1S81=1890 tee ee ASSOC eee 6s.

BITS Ope RE oe fe) Rec AE ot aren Sad he Ad ES tee fe 36s, s005 Ck

RS OP mnie pre aay Rte Woe heen ae (ee cial Hk ME be 36s sous 4kS5

SO SMM Lei art ent ck Shy MM tar ane ACU Some cach a ai 36s. 48s

Tee ope cate OUI ARS GE, nL TG Na tna cIRSHRAO nalRte . 36s 485

TIGOS 5 \bser a Sahel RSI aE RT er ne CAE gah Lakes Sn iON MIR 36s. 48s

SOC MPP enter aeye cro hcen an eee has cna ne CAL 36s . 48s

WS dicta SiS OORT ERE Ce REE EE nr ar eee 36s 48s

WEES. 5.6.5 8 6.0:00.0'5 6 COCR ee ot Re Ren on ee eee 36s 48s

He £4 SEs ocd oc RO RR eae ARUN a es Sg 36s. soo Zee

LS OO Pee eee ey rer gt Ut ly ie ns SOME gli 36s. ere LOS:

InidiexeplSO=1 900 paneer es AS, OG sono. (8

* No perfect copies in stock. Tt Out of print.

PROCEEDINGS or tot GENERAL MEETINGS ror SCIENTIFIC

‘BUSINESS or true ZOOLOGICAL SOCIETY OF LONDON.

8vo. 27 vols. and Index.

Price to Price to the

Fellows, Public.

OOD. Wolk hse ae eee aie en eS 6. 8). ot a aa eae Deetpanah arsiatae 1S.

st Bote Rar o DS ickd Dias ce SRA MEE ey 00 A 8 18s) eens 24s.

TSO 3 see SCENE eae Ron PSAP ERUEO Ui. Sa ca is co even MSH aS ad 5 EES

ee i LAD SUES au ceca aa Ee See GS ang arth «Se ep eee Mee a 3 4 eG

TEST OS sae cei Gs fe ait estan ial ea i aE eae RPT ra HATO Wek Pal Ala 24s.

s 531 RAIN Ne Pe REAR YEN © Se. Soe'gata ton) vel oe! a ca ee heres sag OMS.

MOA Ng 52s SNe eG tcea (os Ray thes cae ea ea ace ain Bhd iG, RENT eh save 1s: eae DAs.

‘i Na VCTIN Tein emera inn ed INEST EN MEAN ice ie ku ae SOE aR 18s\aaee DAs.

EGOS es eae UMN cyt vata teas, Gee's oudskicig cat dy ede ae eee 183: 2 cane PAs.

_ Zak bs 15th a aM IE Rae MR ET al 18s, 5 Gap aaelss

JUSTO SS SSO CESS eee Sr eRe ot 8S RS ee ee eae a 18s., [ease

Oba ie hd AS CO GMB VR ReZ ERA ets of oe nner Acta ERD 18s;. ieee ase

SURG TO eave atige Eats atte ae aia MEE LEST BE Atte ps one eae Reenter 188.525 aes

a A Uae Cee RRR Te, Nhe oR Ce RT Rr 1 S's: Gener 24s,

TAS) OST ig act apa ee eca aa PAR eS co oUt ere Rn ESM NERD aan ey 18s. (2. ae 2a

Ae wg SLUGS) Acsiioe Bes ballot 4a Ege ee IRON RRR cP am Cereal 188) AS eerie

TIGIONS RAO eee) Uncen Mera L, os cry MEO Fees aunts Aya ment oC ANS les. 2 sees

Pa ALTA SARS Ey «aN SRS oe eet IE gE rc PARC we csc ge TSoe: ce 24s,

MONO sek Dea Ws oie ek, cla Beane, uh Gera ake Wet sue hede baa a LBS. 055 ee 24s,

eer par Ov Leu yeier en eects MEET NE APG Hee Tite nicia cA See os 18s." oes

Hnclesxe OO WT OM ON fe 20): AV al tl caee eeaned scree uk see ee as eee ine EPG, eee) OR.

TOWANDA aap sshake 2, feet of co mae Recess gs Ro aie Sean aR a TS sc eres DAs.

‘ Art Aedes rn eR, ot ee anions mica dls Gia 18s. venQas:

BPO Onl ikS AMR srs ge, Sines co R cy ue cold ole at aNEDS eal Re cere eR gaa 18s) eae 21s.

¥ rl ON Na AA eer nee Peers OA aA Re TOTO ST NTER GIG Gis eo alerG [Sse See ees QAs

NUS ities SRO aL ca Bae dsr, eoceice hie (aiiee sl cetesehe a at ycatte dace nt oe NG ISS 24s.

‘i, VME Feat i psc: Ni oe ee ae ER gee Cee ec dee Oe 18s. 24s

ay er MeL eas ml shar ok erste cele: < eae Coal eur Wend acts ane Sener Ree 18s. ska eeas

LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS.

List of the Vertebrated Animals now or lately Living in the Gardens

of the Zoological Society of London. (Kighth Edition.) 8vo.

1883. Cloth, 4s. 6d.

List of the Vertebrated Animals now or lately Living in the Gardens

of the Zoological Society of London. (Ninth Edition.) 8vo.

1896. Cloth, 6s.

Catalogue of the Library of the Zoological Society of London

(Fifth Edition.) 8vo. 1902. Cloth, 6s.

THE OFFICIAL ILLUSTRATED GARDEN GUIDE— 12th Kdition

—with (1) a Railway and Street Map, showing a direct

route to the ‘ Zoo” from all parts of London and the Suburbs ;

(2) a Plan of the Grounds, showing at a glance the location of

the animals; (3) a short description of some of the principal

animals in the Collection (now containing over 3000 spe-

cimens), together with 50 Photographic Illustrations and

Index. Price 6d. in Stiff Paper Cover, postage l3d., or in

Green Cloth Cover price 1s, 2d. post free.

P. CHALMERS MITCHELL,

Secretary.

Regent’s Park, London, N.W.

Jnue, 1914.

ZOCROGICNE SOCIETY OF LONDON.

THE ZOOLOGICAL RECORD.

ies object of the ZootoaicaL Record is to give, by means of an

annual Volume, complete lists of the Works and Publications

relating to Zoology in all its branches that have appeared during

the year preceding the issue of the Volume, together with full

information as to the points they deal with, arranged in such a

manner as to serve as an Index to the literature of Zoology in all

parts of the globe, and thus to form a repertory that will retain its

value for the Student in future years.

The ‘ Zoological Record’ having been amalgamated with the

International Catalogue of Scientific Literature, Zoology, volumes

from 43 onwards can now be obtained only from Messrs. Harrison

& Sons, except when purchasing complete sets from the Zoological

Society.

Under the scheme of amalgamation, Fellows of the Society, and

Institutions already on the subscription-list, have the privilege of

subscribing at the old rate of 30s. per annum, which covers the

cost of carriage of the volume. The subscription becomes due on

July lst. ineach year, and lapsesif not paid by the 1st. of December

following.

The Society is able to supply complete sets of the Record on the

following terms :—

Vols. 1 to 42, price £16 10s. net. Vol. 43 and onwards at 40s. each.

The prices for separate volumes are as follows :—_

Vols. 1 to 42 (except Vols. 4 and 6 which are sold with sets only) 10s.

The price of the ‘Zoological Record,’ Vol. 43 and subsequent volumes,

obtainable separately only from Messrs. Harrison and Co., is 40s.

each. .

Index Zoologicus. An alphabetical list of names of genera

and subgenera proposed for use in Zoology, as recorded in the

‘Zoological Record,’ 1880-1900; . together with other names not

included in the ‘ Nomenclator Zoologicus’ of S. H. Scudder. Com-

piled (for the Zoological Society of London) by CHarnzs Owen

WarERHOUSE and edited by Davin Suarp, Editor of the ‘ Zoological

Record.’ London, 1902. Price to Fellows, 18s.; price to the

public, 20s., or if sold with a set of the ‘ Zoological Record,’ 10s.

Index Zoologicus, No. II. An alphabetical list of names of

genera and subgenera proposed for use in Zoology, as recorded in

the ‘ Zoological Record,’ Vols. 38-47 inclusive (1901-1910), and

the Zoology volumes of the ‘ International Catalogue of Scientific

Literature, Annual Issues 1-10.. Compiled (for the Zoological

Society of London) by Cuartes Owen WareErnovuse, I.8.0., and

edited by Davrp Suarp, M.A., F.R.S., Editor of the ‘ Zoological

Record. London, 1912. Price to Fellows,-12s. 6d. net; price

to the public, 15s. net., or if sold with a set of the ‘ Zoological

Record, 10s.

Divisions of Vols. 39 to 42 of the ‘ Zoological Record’ can be

supplied by the Society, but those of Vol. 43 onwards can be had only

of Messrs. Harrison & Sons, 46 St. Martin’s Lane, W.C.

[pp «©.

SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.

Divisions of the ‘ Zoological Record,’ Vols. 39-42, containing

the literature of the years 1902-1905, may be obtained separately

as follows :—

cae

List of abbreviations of journals, etc.

Special Records, viz. :—

cal

are

XIII.

XIV.

KW.

XVI.

XVIL.

XVIII.

Index of new names of genera and subgenera.

. General Subjects ..

. Mammalia

. Aves ls Ale So pte:

. Reptilia and Batrachia. .

. Pisces

. Tunicata

. Mollusca

. Brachiopoda ..

. Bryozoa

Crustacea

Arachnida

Myriopoda

Insecta

Echinoderma

Vermes ..

Coelenterata ..

Spongize

Protozoa

an A

WNworPwwntsrPwoNnwre £Fe te Ww Sb bw

SOS C28 OG SO AQAeeana GO Dpnoogeo ag &@ OA Ee

Divisions from Vol. 43 onwards are now supplied by

Harrison & Sons, 46 St. Martin’s Lane, London, W.C.

P. CHALMERS MITCHELL,

Secretary.

Regent's Park, Lonpon, N.W.

June, 1914.

P.Z.S. 1914. Carlsson, Pl. I.

West,Newman Iith.

1-3. CYNODICTIS INTERMEDIUS. ‘

Bu 13), (CE NANIOIDOIN, [C1 RUA(C ULL S.

ON FOSSIL CARNIVORA. Dah

PAPERS.

13. On the Fossil Carnivores Cynodictis intermedius and

“Cynodon gracilis from the Phosphorites of Querey. By

ALBERTINA CARLSSON *, Zootomical Institute, University

of Stockholm.

[Received December 23, 1913: Read Aprul 21, 1914. ]

(Plate I.+)

INDEX.

Morphology : Page

(OOTOOMMOLIS CILTAAIERIONS. 555 0909000 soe 22 Sebqncoasee waoueees PAT

(COON. GRECOUOS cos nas ond oaahon ssa0ce opasva dracon babeceBaacconae [eAls)

Among the fragmentary fossils of Carnivora from the Phos-

phorites of Quercy which belong to the collections of the

Zootomical Institute in Stockholm, and which Professor Leche

has requested me to examine, there were two which excited my

special interest.

CYNODICTIS INTERMEDIUS. (PI. I. figs. 1-3.)

One was a part of a skull, consisting of the well-preserved

brain-case and a fragment of the face. In the Catalogue it is

designated No. 2216, Cynodictis sp. Comparing it with the

descriptions of Filhol (1, p. 116) and Schlosser (4, pp. 40 & 47),

I have identified it as Cynodictis intermedius.

As we know very little about the skull of this species, or

about that of most species of the same genus—Filhol knows

only the foremost part of the face,—I will here give its characters.

The skull in question belongs to an adult animal, for only the

sutures between the squamosals and the frontals, as well as those

between the basioccipital and the basisphenoid, can be seen.

The head, which, as in other forms of Cynodictis, is long and

not very broad, has a brain-case which, in its narrow and elon-

gated form, vegeralolles more that of the recent Wirauetks than -

that of the now living Canide (fig. 1).

Filbol (1, p. 72) says of Cynodictis boried :—‘ La masse ae yell

devait tre proportionnellement petite par rapport a la taille de

Yanimal.” As in Cynodictis gryei (1, pl. 19), the face seems to.

be comparatively more developed than the brain-case, the frontals

—if it is allowed to judge from the distance between the orbits—

being comparatively broader than in the Viverride. The crista

sagittalis has attained a much higher degree of development than

in "Viverrides and Canidez of the same size. As in the Viverride,

the crista occipitalis is high, and consequently the height of the

supraoccipital is considerable (fig. 2).

* Communicated by OLpriELD THomas, F.R.S., F.Z.S.

+ For explanation of the Plate see p. 230.

Proc, Zoou. Soc.—1914, No. XVI. 16

228 MISS A. CARLSSON ON

As in the Canide, the foramen condyloideum (fig. 3, cond.) is

situated far behind the foramen lacerum posterius (lac.); in the

Viverridee, as is well known, these foramina open into a common

fossa, separated from each other only by a bony bridge. Also,

with’ regard to the foramen glenoideum, the skull in question

-ayrees with that of the Canidie, this foramen being here rather,

distinct; in the Viverride it is represented by an extremely

small opening.

The canalis alisphenoideus (alisph.) is longer in Cynodictis

intermedius than in the two above-mentioned families, and con-

sequently the alisphenoid is of greater length in the former than

in the latter. Below the posterior nares the border of the

palatines terminates in a ridge; such a ridge is not to be seen in

the Canide or in the Viverride, but is found, more or less

developed, in Propithecus, Hrinaceus, Perameles, Myrmecobius,

and Sinopa (3). The bullae ossee are not preserved, probably on

account of having been loosely attached to the skull, as they are

still found in young bears and in Paradoxurus. No doubt they

were ossified, as is the case in C. boriei and C. gryei (1, pp. 73

& 79). From the form and the direction of the processus par-

occipitales (pr.), it appears that they have not, as in the Viverridee,

been spread over the hinder surface of the bulle, but have been

separated from them as in the Canide.

The petrous bone is pear-shaped; on its lateral side there is a

deep fossa. As the foramen stylomastoideum (séyl.) opens into

it, it must be formed by a portion of the mastoid.

Thus the skull of Cynodictis agrees partly with the Canidz and

partly with the Viverride. Though in most respects it agrees

with the former, especially with regard to the basis cranu, the

form of the brain-case indicates kinship with the latter. Such

a kinship has been stated before by Filhol on account of the

result of his examination of the teeth and the tooth-formula of

the lower jaw of several species of Cynodictis.

Cynopon @raciuis. (PI. I. figs. 4, 5.)

The other fragment was part of a skull of Cynodon gracilis

designated No. 1621. Of this species Filhol has described only

the lower jaw (1, p. 120); the skull was as yet unknown.

A short description of the fragment in question may therefore be

of interest. It consists of a tolerably well-preserved part of the

face; on the right side the anterior part of the zygomatic arch

remains, on the left it is totally absent. The brain-case is not

entire, but its length and outlines can be seen. Of the basis

eranii, the portion of the basisphenoid which borders on the

pterygoids was preserved.

Of the lower jaw there are four fragments of different size ;

in two of them the three posterior premolars and the two

anterior molars are found; a left half of the lower jaw shows

FOSSIL CARNIVORA. 229

the alveoli of C,, P,, P,, and fragments of P,, P,, and M,; a

right half contains the canine, the premolars, and the molars.

The small size of the examined object is very remarkable.

According to Filhel (4, p. 12), ue, measures 5°5 and M, 8 mm.

in length; however, I found that the corresponding teeth

measure 4 and 5°5 mm. respectively. But it must be borne in

mind that, according to Schlosser (4, p. 54), in the specimen of

Cynodon gracilis in the Museum of Munich, M, is not quite

as long as that of the specimen examined by Filhol, and that

there is a possibility that this skull belonged to a female or

to a small specimen. How widely the size of the skull of Car-

nivora varies has been shown by Hensel (2).

That we have here to do with Cynodon gracilis, and not with

O. velaunus, C. aimardu, C. speciosus, or C. leptorhynchus, 1s

evident, as all these, which also occur in the Phosphorites of

Quercy, are larger than C. gracilis.

The sutures between the maxillaries and the frontals, those

between the latter and the nasals, those between the jugal and

the maxillary, as well as the outlines of the lachrymal, and those

of the orbitosphenoid are very distinct—showing that the

specimen is young. ‘There is, however, a crista sagittalis (fig. 5).

In Cynodon gracilis the canalis infraorbitalis opens above P*, in

Cynodon leptorhynchus, according to Schlosser (5, p. 115), above the

interspace between P* and P*, which probably depends on the

greater length of the jaws of the latter. The nasals are very

short, and consequently the anterior nares have an oblique

direction as in Cynodictis gryei, and as in the last-mentioned

animal the nasals stretch between the frontals above the

maxillaries (1, pl. 19. fig. 58). Judging from the strength and

direction of the preserved part of the jugal, the zygomatic arch

seems to have been strong and wide, thus differing from that of

Cynodon leptorhynchus, of which Schlosser states :—* Jochbogen

schlank, wenig gekrummt” (5, p. 115).

The upper incisors are separated by small gaps. The upper C

measures 7-5 mm. in height; P* has one root, and stands at some

distance from the canine. P? and P* are of the same form as

P, and P,, but of smaller size. As to P*, it has a strong cingulum,

as have the anterior premolars ; its anterior cusp is higher than

the posterior. Only its labial side was to be seen, which was

also the case with M' and M*. The height of the two latter teeth

is 5 mm. (fig. 4). ving

The lower jaw and its teeth have been most satisfactorily

deseribed by Filhol (1, p. 120).

List of References.

1. Finnor, H.—“ Rech. s. 1]. Phosphorites du Quercy,” in Ann.

d. Sci. Géol. t. 7. Paris, 1876.

16*

230 ON FOSSIL CARNIVORA.

2. Henset, T. R.—‘‘Craniologische Studien,” in Nova Acta der

Ksl. Leop.-Carol. Deutschen Akademie der Naturforscher,

Bd. xlii. Nr. 4. Halle, 1881.

3. Marraew, W. D.—‘‘The Osteology of Sinopa, a Creodont

Mammal of the Middle Eocene,’ in the Proc. of the

United States National Museum, vol. xxx.

4. Scutosser, M.—‘“‘ Die Affen, Lemuren etc. des europiiischen

Tertiars,” in Beitr. z. Paliontologie Cisterreich-Ungarns,

Balt vane yale airs

5. Scutosser, M.—‘ Ueber den Baren und den Birenahnlichen

Formen des europiiischen Tertiirs,” in Paleontographica,

Bd. xlvi. Stuttgart, 1899.

IXPLANATION OF PLATE IT.

Cynodictis intermedius.

Fig. 1. Skull, upper view. 1/1 nat. size.

2. Skull, seen from behind. 1/1 nat. size.

8. Basis cranii. 1/1 nat. size.

alisph. Canalis alisphenoideus ; cond. Foramen condyloideum ;

gl. Foramen glenoideum; fac. Foramen lacerum _posterius ;

ov. Foramen ovale; pr. Processus paroccipitalis ; sty/. Foramen

stylomastoideum.

The hatched parts of the figures represent lost portions

replaced by cement.

Cynodon gracilis.

Fig. 4. Part of the skull, right view. 1/1 nat. size.

Only the roots of M! and M? are left. The top of P? is lost.

5. Part of the skull, upper view. 1/1 nat. size.

ON TWO ANNELID GENERA. 931

14. On the Genera Ceratocephale Malmgren and Tylo-

rhynchus Grube. By L. N. G. Ramsay, MeAwy Basc:,

Carnegie Research Scholar, Christ’s Coliege, Cam-

bridge *.

[Received December 12, 1913: Read March 17, 1914]

InDEX.

Geographical distribution of Tylor hychus .......cccecereee serene 233

Systematic :

Ceratocephale osawai Izuka= Tylorhynchus chinensis Grube ... 231

Genera Ceratocephale and Tylorhynchus, characters and

relations 233

Along with the Nereide which he collected at Puget Sound in

1911, Mr. F. A. Potts recently handed over to me some specimens

of Ceratocephale osawai from the Sumida River, Tokyo, Japan.

The general facies of these worms recalled irresistibly to my

mind Grube’s figures of Tylorhynchus chinensis, n his paper on

the Annelids of the ‘ Novara’ Expedition (1867). Reference

to the literature shows conclusively that the two species are

identical.

The object of this note, then, is to draw attention to this fact,

and to indicate the characters and relationship of the two genera

more clearly than these have hitherto been recognized.

The identity of Tylorhynchus chinensis with Ceratocephaie

osawai.—Izuka, discussing the systematic position of C. osawai in

his original description of the species, remarks that he refers it

to the genus Ceratocephale tentatively, in preference to creating

» new genus for it (1903, p. 3). If we are to regard Tylo-

rhynchus as a distinct genus (and there appears to be sufficient

ground for this) Tzuka’s hesitation was fully justified. It is

probable that only lack of access to Grube’s paper prevented him

from recognizing the true position of the species.

The facts are as follows :—The Japanese specimens (Izuka’s

full description 1s illustrated by numerous excellent figures)

agree in every way with those described by Grube 7 from

Shanghai—as to parapodia, sete, general form, prostomium and

its appendages; but they differ in the fact that the basal ring

of the proboscis is devoid of papille, while those on the dorsal

surface of the maxillary ring are less numerous.

Among the specimens at present before me, however, three

which have the proboscis everted exhibit a number of large

swollen papille or protuberances on the basal ring, corresponding

approximately to those figured by Grube, while the condition of

* Communicated by Dr. A. E. SHIPLEY, E.RS., F.Z.S. ;

+ Grube’s description is fully detailed, and is accompanied by figures by

A. Afsmann, which are far above the usual standard. of the figures accompanying

this authority's papers.

232, MR. L. N. G. RAMSAY ON

the maxillary ring is intermediate between those described by

Grube and Izuka. Moreover, the smallest of these three speci-

mens (which are all in the heteronereid state) has these papille

distinctly less swollen and conspicuous than the two larger ones.

It is to be noted that Izuka’s specimens were from the same

locality as these (the Sumida River).

Further, Izuka himseif (1903, p. 9) states that the papille on

the neural surface of the maxillary ring seem to increase in

number with the age of the individual, and that in larger

specimens their number varies from 17 to 27.

Further still, Izuka describes the proboscis of the worm in the

nereid-form only, while Grube’s specimens as they are figured

were in the heteronereid condition. It is conceivable that as the

worms assume the heteronereid form, the changes in the posterior

region of the body may be accompanied by some swelling of the

areas between the deep furrows which exist in the basal ring

in the immature form. Unfortunately, none of the immature

specimens at present before me has the proboscis everted, and

the method of examination by slitting open this organ does not

give very satisfactory results, owing to ‘the difficulty of recognizing

the soft papille in the compr essed condition resulting from the

inverting of the proboscis.

It appears, therefore, that there is no ground for even the

specific separation of chinensis and osawai, and the latter name

must therefore stand as a synonym of Tylorhynchus chinensis

Grube.

The relationship of the Genera Ceratocephale and Tylo-

rhynchus.—These genera were described independently by

Malmgren and Grube in the year 1867. Since that year,

various authorities have mentioned them in their schemes of

classification of the Nereide, but their characters, especially those

ot Tylorhynchus, do not appear to have been properly com-

prehended. Matters were further confused when a species of

the latter was referred to the former genus by Izuka in 1903,

as explained above.

We must first consider whether the two genera should be

regarded as distinct. On this head I think it will be agreed

that the divergences are sufficient to warrant generie separation,

although the two, one of which is, so far as known, confined to

the Atlantic, the other to the Pacific Ocean, are certainly much

more closely allied to one another than to any other genus.

he chief points of difference appear to be as follows * :—

In Tylorhynchus,

(a) The parapodium lacks the neuro-ligule

(6) The neuro-cirrus is normal.

* T have not had an opportunity of examining specimens of Ceratocephale,

which has, however, been figured and described by Malmgren (1867), McIntosh

(1902), and Heinen (1911).

TWO ANNELID GENERA. UBS

In Ceratocephale, these characters ave negatived, the neuro-

ligule being well-developed, the neuro-cirrus double, and eyes

completely absent.

In other respects, the two show marked agreement.

‘Lhe two genera appear to form a small group, diverging from

the remainder of the family in the form of the parapodia, while

showing some resemblance to the genera Leonnates and Leptonereis

in the armature of the proboscis.

Hitherto most authors have stated that TZylorhynchus is

characterized by the presence of hard callosities on the proboscis,

as opposed to the soft papillae of Ceratocephale. This is clearly

due to a misapprehension of Grube’s original description of

T. chinensis, where he describes the proboscis-armature as

consisting of “calli” (Latin) or “ platten schwielchen.” There

is, however, no reason to suppose that these differ in nature

from the uncornified papille of Ceratocephale, Leonnates, ete.

Survey of the two Genera as known up to date:—

Genus TyLorHyNncHus Grube, 1867.

Characters.—Prostomium bearing palps, tentacles, and two

pairs of eyes, as in Wereis L. Peristomium apodous, with four

pairs of tentacular cirri. Proboscis armed only with soft papille,

and with a pair of jaws of typical nereid form. Parapodia

biramous, the rami not deeply divided, the notopodium con-

sisting of the mferior ligule only, with setigerous lobe at its

base, the superior ligule being absent. In the first two pairs the

aciculum and setigerous lobe of the notopodium are absent.

The noto-cirrus in the anterior region borne upon an enlarged

rounded lobe; the neuropodium consisting of setigerous lobe

alone, without the ventral ligule, and bearing two “lips,” anterior

and posterior; ventral cirrus simple. In the epitokous form no

special foliaceous lobes are developed, but the original lobes are

enlarged and flattened somewhat, in the middle region of the

body. The sete are homogomph and heterogomph spinigers and

falcigers as in WVereis, but some are of slightly peculiar type. In

the heteronereid cultrate setze appear, as in Nereis,

Range.—Western coasts of Japan (Izuka), Shanghai (Grube),

Java (Quatrefages, see below),

Species :—

T. CHINENSIS Grube, 1867.

Syn, Ceratocephale osawai Izuka, 1903 and 1912.

Nereis heterocheta Quatrefages, 1865.

** Die Japanischen Palolo,” Osawa, 1901.

Range.—As for the genus.

Quatretages (1865, p. 552) described, in his ‘ Histoire des

234 MR. L. N. G. RAMSAY ON

Annelés,’ a nereid which he called Nereis heterocheta, from Java.

Grube (1870, p. 312) states that this description refers to Tylo-

rhynchus chinensis. Quatrefages’ description is not sufficient

for identification, but as Grube personally examined Quatrefages’

material in the Paris Museum, there is no reason for hesitation

in accepting this record as a good one. As Quatrefages’ de-

scription was very inadequate, and was accompanied by no figures,

I have not thought it necessary to give his name preference to

that of Grube.

Genus CeraTocEPHALE Malmgren, 1867.

Characters.—As for Tylorhynchus, except that :—

Hyes are absent.

The neuropodium bears a well-developed ventral ligule

as well as the setigerous lobe.

The neuro-cirri are double, the two cirri arising from a

common base.

Range.—The North Atlantic (Scandinavia: Malmgren, Heinen,

Levinsen, etc.); Eastern North America (Gulf of St. Lawrence :

McIntosh ; Massachusetts: Verrill).

Only one species has been satisfactorily described, viz., C. loveni

Malmgren, 1867, from the Scandinavian waters. Those from

the American side may prove to be identical with this, although

Verrill (1879) named his specimens C’. websteri, without, however,

giving a satisfactory diagnosis, and without figures.

Bibliography of Ceratocephale and Tylorhynchus.

1865. QuatTREeFAGEs.—Histoire Naturelle des Annelés. Paris,

1865.

1867. Matmeren, A. J.—‘“‘ Annulata polycheta Spetsbergie,

Gronlandize, Islandiz et Scandinavie hactenus cognita.”

Ofvers. af K. Vet.-Akad. Férh. 1867, p. 176, tab. vi.

fig. 33, etc. ;

1867. Gruss, E.—Reise der Osterreichischen Fregatte ‘ Novara.’

Anneliden. Wien, 1867, p. 22, Taf. i1. fig. 3, ete. .

1868. Euurrs, E.—Die Borstenwiirmer. Leipzig, 1868, p. 577.

1870. Gruss, E.—‘“ Bemerkungen iiber Anneliden der Pariser

Museums.” Arch. f. Naturges. 1870.

1879. Verriti, A. E.—“ Recent additions to Marine Investigation

of the N.E. Coast of America.” Proc. U.S. Nat. Mus.

Tile Js IG,

1879. Tauser, P.—* Annulata Danica, i.” Kjobenhavn, 1879.

1893. Lrvinsen, G. M. R.—Annulata, etc. (‘“Sertryk af det

Videnskabelige udbytte af Kanonbaaden Hauchs ”),

Togter 1., 1883-86. Kjobenhavn, 1893.

1894. Brpenxap, O.—‘‘ Norges Annulata Polycheta.” Vidensk.

Selsk. Forh. Christiania, 1894.

ISTO

O02

1903.

WSL

1912.

TWO ANNELID GENERA 935

Osawa, K.—‘ Uber die Japanischen Palolo.” Verh. Congr.

Zool. Berlin, 1901, pp. 751-755.

McInrosH, W. C.—‘ Notes from the Gatty Marine

Laboratory, xxiii. 6. ‘On Canadian Nereide from the

Gulf of St. Lawrence.’” Ann. Mag. Nat. Hist. (7) x.

p. 258, pl. vi.

Tzuxa, A.—< Observations on the Japanese Palolo ( Cerato-

cephale osawai).” Journ. Coll. Japan, xvii. (1i.), pls. i.

& il.

Hemen, A.—‘‘ Die Nephthydeen und Lycoriden der

Nord- und Ostsee.” Wiss. Meeresunters. Kiel, N. F.

xiii. 1911, p. 62, Taf. i. figs. 11-15.

Izuxa, A.—‘‘ Errantiate Polycheta of Japan.” J. Coll.

Japan, xxx. 1912, p. 179.

7 ae ie ; Anti epee Bisie Anthea wi

ey 2 Re suiiaiag ai) boy op

- 5 ~~ an nif

aes

ON ANNELIDS FROM THE N.E. PACIFIC. Werth

15. On the Annelids of the Family Nereidz collected by

Mr. F. A. Potts in the N.E. Peific, in 1911.—With

a Note on the Morphology of Micronereis as a Repre-

sentative of the Ancestral Type of the Nereide. By

L. N. G. Ramsay, M.A., B.Se., Carnegie Research

Scholar, Christ’s College, Cambridge *

[Received December 12, 1913; Read March 17, 1914. |

(Text-figures 1-7.)

INDEX. Page

Morphology: -

SERS Olt INGINGHS COTAILYPIIS VIVRE, coss.ocabodo occ cos coobaceeueae sbebonsenenavos | BIS)

Micronereis, external morphology ..............00055 ceceeeeeeeee eee. 245

TOWONOINOIN OF ING RGED CRBICUUIS. oscac000 000500000 soa doo bus ont bos enoatansouesces | CAO

Geographical :

Micronereis variegata Clap., from Pacific Ocean .................. 243

Systematic :

Nereis shishidot Iauka,=N. cyclurus Harr. .......0..00c00 0c cece 289

PV. ezoensis Iauka,= N. vewillosa Grube ...............0c0.00 esses. 240

IN, dyamusi Vauka= I. virens Sars\......-..cc0cc0 eer cee see sen een san se, 2AQ

N. dumerilii, kobiensis, agassizi discussed ............000.00.000..... 242

Micronereis, relationship to Nereida ................0..6.0c0ee0cc0e000 247

The Annelid fauna of the Pacific coast of North America is

already fairly well known, and in the present paper only one

addition is made to it. This, however, is of great interest, as it

is a representative of the little known genus MJicronereis, and is

apparently identical with J/. variegata of European waters, the

solitary species of the genus.

The remaining five species contained in the collection are

already well known in this region; several of these are repre-

sented on the opposite shores of the Pacific by very nearly allied

species, some criticism of which will be found under their various

heads.

The collection also includes two examples of the remarkable

and extremely interesting Vereis cyclurus Harr., from which some

new facts as to the setz are brought to light.

NEREIS cycLuRUS Harr.

Nereis cyclurus Harrington (4), p. 214, 1897, pls. xvi., xvii.,

XViil.

N. -shishidoi Iizuka (5), p. 177, pl. xix. figs. 10-18.

Two examples, Nanoose Bay, June 13th, 1911. Dredged in

6 fathoms, in soft mud. Commensal with hermit-crabs.

In Mr. Potts’ notes these are described as of a ‘pale fiesh-

* Communicated by Prof, J. SranLEY GARDINER, F.R.S., F.Z.S.,

238 MR. L. N. G. RAMSAY ON

colour, the palps and anterior part of prostomium white, as also a

prominent nuchal band. Tentacles and tentacular cirri brownish.

In every segment a narrow anterior white band. Posteriorly the

notopodia increase in size and become white in colour. Dorsal

cirri flesh-coloured.”

This remarkable species has been fully described and figured by

Harrington and Johnson (6, pls. iv., v.). I have here to add only a

few further notes on the sete and some remarks on its affinities.

As to the bristles, both specimens under consideration exhibit

an important feature, which bas either been overlooked by other

writers, or as is perhaps less probable, occurs only in some spe-

cimens. This is the occurrence of a curious homogomph faleate

type of seta in the notopodial bundle of the posterior parapodia.

This form of seta, which is not unlike that which occurs in the

notopodium of the posterior parapodia of WV. procera Ebl., is

figured (text-fig. 1). In the larger of the two, which measures

Text-figure 1.

Homogomph falcigerous seta from notopodial bundle of 50th parapodium

of Nereis cyclurus. X 200.

75 mm. in length, with 107 pairs of parapodia, and has the

characters of the peristome and parapods fully developed, these

falcate setze appear at the 32nd parapod. They are at first two

in number, increasing to three or four about the 50th to 70th,

and decreasing to one or two near the posterior end. The

spinigers of the dorsal bundle, numerous anteriorly, decrease

steadily in number, until by the 50th there are only two or

three and further on none at all.

Anteriorly the lower neuropodial bundle consists entirely of

falcigers, but about the same region as in the case of the noto-

podial falcigers, heterogomph spinigers appear in small numbers,

and persist to the posterior end.

In the smaller specimen, which is about 26 mm. long with 59

setigerous segments (it is incomplete, and lacks perhaps 20 segments

posteriorly), the peristome is not fully developed. The distri-

bution of the sete agrees exactly with that given above, the

notopodial falcigers commencing about the 29th pair.

ANNELIDS FROM THE N.E. PACIFIC.

239

The seta-distribution in these two specimens may therefore be

summarised thus :—

Region of body

Anterior.

Mid.

Posterior,

Notopodial bundle.

Hom. spins.

(numerous).

Hom. spins. (few).

Hom. fales. (few).

Hom. fales. (1-2).

Hom. spins. (rare).

Upper neuropodial | Hom. spins. |Hom. spins. (fewer). Hom. spins.

bundle. (numerous).

Het. falcs. Het. fales. Het. fales.

(few).

Lower neuropodial Het. spins. (few). Het. spins. (few).

bundle.

Het. fales. | Het. fales. (fewer). Het. fales.

(numerous). |

Nereis cyclurus appears to have a considerable range along the

western sea-board of North America, from Southern California

(Moore, 12) to Puget Sound.

Izuka has described WV. shishidoi, a closely allied species, from

the eastern side of Japan. The description is based on the

solitary specimen which has been obtained there.

It is difficult to imagine what reasons Izuka can have had for

separating the Japanese example as a distinct species. His

description and figures, which are excellent, agree exactly with

NV. cyclurus. His only comment on the relationship of the

Japanese specimen to the American ones is that WV. shishidot

‘comes nearest” to WV. cyclurus. Harrington’s description of the

paragnaths of the basal ring (V—VITJ) is a httle difficult to under-

stand, and this may have led Izuka to separate his specimen.

Comparison, however, of the present specimens shows that the

paragnaths agree exactly with those of the Japanese worm.

Harrington, after summarising the differences between JV. cy-

elurus and NV. fucata Sav., concluded that these two worms, both

normally commensal with hermit-crabs, were closely related.

Moore (11, p. 343) suggests the generic separation of cyclurus

from other nereids on account of the extraordinary peristomium.

To the present writer it seems that these authorities are both

right in their way. cyclurus is possibly more nearly related to

fucata than to any other species, as the form of the parapods,

sete, arrangement of paragnaths, etc., would seem to indicate.

The enlarged bilobed noto-ligule in eyclwrus, bearing the noto-

cirrus below the upper lobe, is little more than an exaggerated

form of the noto-ligule of fwcata. On the other hand, the

development of the peristomium in the full-grown cyclurus is so

extraordinary that it might be regarded as ample ground for the

establishment of a new genus, were it not for the close resemblances

in other respects to certain other members of the genus Verets.

240 MR. L. N. G. RAMSAY ON

Here we have an interesting problem. JV. cyclurus and NV. fucata

are admittedly closely allied in some respects. The former inhabits

the North Pacific, the latter the North Atlantic. It is probable

that at a former epoch the distribution of the two was continuous

(around the north coast either of Eurasia or of America, or vid

Panama). The existence of certain other closely allied and even

identical species in the two regions points to this. Further, it

seems probable that the hypothetical ancestor-species of the two

had already formed its commensalistic habits before the distri-

bution-area was severed (it would be a remarkable coincidence

for the two allied species to have taken up the same habit

independently—a habit not exhibited by any other nereid).

Then, after the separation, the Pacific division of the ancestral

Species, apparently under the same conditions (both inhabit

Gastropod shells tenanted by species of Hupagurus, Pagurus, etc.),

has made an enormous stride in evelution, the while that the

Atlantic division has remained more or less stationary !

NEREIS VEXILLOSA Grube.

Nereis vexillosa Grube, Ehlers (2), p. 573, Taf. xxiii. figs. 3-5.

An example of this fine species, labelled “St. Paul Island,

Alaska, 1897,” accompanies the collection.

This form, originally described by Grube from the Sea of

Okhotsk (3, p. 3, Taf. i. figs. 1, 5,6), is now known to inhabit also

the west coast of America from Alaska to California. Izuka, in

1912 (5, p. 173), described a very closely allied species from Japan,

IV. ezoensis, He givesno reason for separating this, except that

‘‘ the falcate end-piece of bristles [is] much longer in this species

than in the closely allied Wereis vewillosa.” He figures a faleate

bristle, but emits to state from what region of the body it is

taken. Now, in JW. vewillesa, as in certain other nereids, the

length of the end-piece of the falcate bristles decreases very

markedly, proceeding from head to tail. In the specimen before

the writer at present, the falcigers from the posterior portion of

the body have certainly appendages very much shorter than that

figured by Izuka, but in those from the anterior portion (e. g.,

10th parapod) they are nearly as elongate as in the figure. I can

discern no other difference between the two species from Izuka’s

description and his excellent figures, and would therefore suggest

a more detailed comparison, with a view to determining the

specific distinctions in a way more satisfactory to others. Apart

from the progressive variation in length of the falcigers through-

out the body of a single individual, that in different individuals

must be taken into account.

NEREIS PELAGICA L.

Nereis pelagica L., Izuka (5), p. 154, pl. xvii. figs. 1-6.

One specimen, dredged near Turn Island, Friday Harbour,

Puget Sound, 18th July, 1911.

ANNELIDS FROM THE N.E. PACIFIC, 241

This specimen is referred to in Mr. Potts’ notes as a ‘blue

nereid, among kelp-roots.”” It is a female heteronereid, of rather

small size, measuring about 35 mm, in length, with 54 pairs of

parapodia (the body is not quite complete posteriorly). The

transition in the form of the parapodia occurs at nos. 16-17.

There is, I think, no doubt that this nereid is of the same

species as that figured and described by Izuka (loc. cit.) under this

name. But while it agrees exactly in other respects with Izuka’s

description and figures, it exhibits a difference in the paragnaths

of the basal division of the proboscis (V-VIII groups). These

are so numerous as to form a complete band around the proboscis,

consisting chiefly of small, pale brown, rounded (not sharply

pointed) paragnaths, covering closely almost the whole surface.

Group VI is represented by a well-marked little bunch of large

pointed paragnaths, which, if alone, would be almost exactly in

accordance with Izuka’s figure, but these are surrounded by the

small paragnaths just described, which invade area V and also

link the dorsal with the ventral areas. VII-VIII are affected

in much the same way.

Izuka’s figures do indicate that the paragnaths of the basal ring

are more numerous than those of European JV. pelagica, but I

suspect that the present example must be a more extreme one, if

not even an abnormality.

The proboscis was only partially everted, so that it would be

difficult to figure it satisfactorily.

The modified parapodia agree in form exactly with those of the

male heteronereid figured by M‘Intosh (8, pl. lxxi. fig. 7) and

with those of other specimens from the British Isles.

NEREIS VIRENS Sars.

Nereis virens Sars, Grube (8), p. 6, Taf. 1. figs. 2, 4, 5, 6.

NV. brandti Malmgren (9), p. 183; Ehlers (2), p. 563.

NV. dyamusi Izuka (5), p. 160, pl. xviii. figs. 1-12.

This large species was found in numbers in muddy beaches at

Departure Bay, Pleasant Beach (Seattle), Barkley Sound, and

Friday. Harbour ( fide collector’s notes).

Two large examples are in the collection (locality @).

Nereis virens is well known on the west coast of North America,

from California to Alaska, and was recorded from the Sea of

Okhotsk by Grube in 1849. The Japanese representative has

been named WV. dyamusi by Izuka (see below).

Malmgren (9, p. 183) decided that the specimens from the Sea

of Okhotsk represented a distinct species, which he named

NV. brandti. Ehlers (loc. cit.) followed him, but noted that the

two species were so nearly allied “dass wir nur die Variation

einer Grundform haben, welche circumpolar im Nordmeere ver-

breitet ist.”

More recent workers, however, agree in considering the Pacific

representative as identical with that of the Atlantic coasts

[Johnson (6), p. 398; Moore (11), p. 344], and I follow these.

QA MR. L. N. G. RAMSAY ON

Moore, in 1909 (12, p. 244) “provisionally ” bestowed the stib-

specific name of plenidentata on specimens from California, which

he found to be characterized by a larger number of segments and

more numerous paragnaths. He also suggested that the form from

the more northerly parts of the Pacific coast should be separated

as JV. virens brandti, on account of similar differences, which he

found to be present in a less marked degree. His final opinion

on the matter has not yet appeared.

Izuka, (5, loc. cit.) founds his new species on slight differences in

the numbers and arrangement of paragnaths in certain groups,

and on the number of teeth in the jaws. I should like here to

draw attention to an exceedingly good paper on Wereis virens

from the Atlantic coast of America (17). In this the writer

gives particulars of the variation of the paragnaths, which he

found to be very wide even in a small number of specimens. [

can see no reason for regarding the Japanese species as distinct

from that of Western America.

Nereis (PLATYNEREIS) AGASSIZI Ehlers.

Nereis agassizi Khlers (2), p. 542, pl. xxiii. fig. 1.

Two small] specimens from Departure Bay, 8th May, 1911.

These are both male heteronereids which were found amongst

material dredged in the Bay.

The paragnaths have the arrangement typical of the species

of the Platynereis group, I, II, and V being absent, the other

groups represented by pectiniform rows, which are not so con-

tinuous as usual, but more broken up. (I have noted this feature

occasionally in Huropean specimens of V. dumerilii.)

The large homogomph falcigerous sete of the notopodial

bundle are represented by a single large bristle with fused

appendage in each parapod, commencing about the 14th pair.

These have been figured by various authors (2, 5, 10).

These two nereids, each of which has about 76 pairs of para-

podia, are about 25-27 mm. in length. The heteronereid form

is not completely assumed, many of the ordinary sets not yet

having been cast in the posterior region. In the more advanced

of the two, the transition in the form of the parapodia occurs at

the 19-20th pair. The noto-cirri of the first seven pairs have

the characteristic swollen outline.

A considerable cloud hangs over the species of Platynereis

which are found in the Northern Pacific (as in the case of those

in other parts of the world).

We are concerned with the three species (?) :-—

NV. dumerilit Aud. et Edw.

XN. kobiensis McIntosh.

NV. agassizi Ehlers.

These species in the Pacific Ocean (dwmerilit, of course,

occurring elsewhere) are all characterized by the presence of

ANNELIDS FROM THE N.E. PACIFIC. 243

the curious fused falcigers in the notopodial bundle, figured by

various authors and mentioned above. The presence of these

setee is the only valid point distinguishing any of these species

from WV. dumerilit of the Atlantic coasts which has hitherto been

mentioned by any author. Further, no author has yet given any

point of distinction between these three Pacific species which has

the least weight whatsoever. For example, Izuka describes and

figures them, one after the other, in his fine work on Japanese

Polycheta, but gives no means whatsoever of distinguishing the

three. They are apparently three absolutely indistinguishable

species, and they even inhabit the same localities. | Nor does he

compare them in any way; in not one of the three fully detailed

descriptions does he even mention the names of the other two

species. Systematic zoology becomes impossible under these

conditions.

It does appear that the Platynereis of the North Pacific in the

adult form * is characterized by these peculiar fused falcigers

(which there can scarcely be a doubt are fully homologous with

the corresponding unfused homogomph falcigers occurring in

Platynereis from all other parts of the Oceans).

If this isa valid specific distinction, then we have the following

synonymy of V. agassizi Ehlers :—

Nereis kobiensis : McIntosh (7), p. 210, pl. xxxiv. figs.

3-6; pl. xviA. figs. 2-4.

Treadwell (15), p. 226; (16), p. 1161.

Izuka (5), p. 162, pl. xvii. figs. 12-13.

Nereis dumerilii A. et E.: Izuka (5), p. 158, pl. xvii. figs. 7-8.

Marenzeller (10), p. 123, pl. ii. fig. 4.

Moore (11, p. 344) has already suggested ‘that the Japanese

specimens referred to WV. dumerilii by Marenzeller belong to this

closely related but perfectly distinct species” (i. e., V. agassizi).

There remains, however, considerable doubt as to whether

N. agassizi should not be regarded as a mere subspecies of

N. dumerilii, and whether the latter has not a world-wide range

in the Southern as well as the Northern Hemisphere. This

I propose to discuss at a future date, after a more complete

examination of the material from all parts of the world which is

at my command.

MICRONEREIS VARIEGATA Claparede.

Micronereis variegata Claparede (1), p. 57, Taf. xi. figs. 5-7,

1863.

Four specimens, Departure Bay, May 1911.

* Von Marenzeller (10) figures an unfused homogomph falciger from a young

Japanese example of N. dwmerilii, which differs in no way from the ordinary type,

and suggests that the unjointed type of bristle in old specimens is derived from this

by fusion of the appendage in its socket.

Proc. Zoou. Soc.— 1914, No, XVII. Lf

Q44 MR. L. N. G. RAMSAY ON

I have compared these with specimens from the South of

England, kindly lent by Major Elwes, and have been unable to

find any points of specific difference between them.

Clapareéde’s (loc. cit.) and McIntosh’s (8, p. 261) descriptions

and figures fit the British Columbia specimens as well as the

European ones.

These tiny nereids, which range’from about 4°5 to 7 mm. in

leneth, were found inhabiting globular masses of mucus on

brown Ulva dredged in Departure Bay. In one case the mucus

contained a mass of eggs as well as the worm. On removal the

mucus was again secreted. The general colour of the living

animals was green, with transverse brown markings on the

dorsum of each segment. The anal cirri were in one case dull

erimson (Collector’s notes).

One of the specimens exhibitsa remarkable peculiarity, namely,

the presence of a number of slender simple set interspersed

among the ordinary compound spinigers. Whether these occur

throughout the body I was unable to ascertain, but in the

mid-region they are certainly present in several pairs of

parapodia.

In the 10th, for example, we have the following :—

Notopodium: 15 spinigers, 17 simple sete.

Neuropodium: 17 3 A airar eae a

The setz are arranged in a single vertical series, the simple

sete appearing singly or two together between the compound.

The former are from one-fourth to one-third the diameter of the

latter, and from about one-third to nearly three-fourths the

length ; they are apparently perfectly smooth, and taper gradually

to a fine point.

I have not found any trace of these in the other three

specimens, so that it is not possible to regard their presence as

a specific distinction.

It should be noted that the four specimens of Micronereis in

this collection are of two different types—in this respect, that

in two of them the trunk and especially the parapodia are so

much swollen that adjacent parapodia are in contact with one

another; while the remaining two have a scraggy, thin appear-

ance, with wide intervals between adjacent parapodia. I am

unable to account for this difference. The “fat” type is

represented in text-fig. 2, while text-fig. 5 is of a parapodium

of the other.

Micronereis has hitherto been recorded only from the French

coasts, both Mediterranean and Atlantic (Claparéde, St. Joseph),

and from the South Coast of England (Zlwes).

ANNELIDS FROM THE N.E. PACIFIC, 24.5

Note on the Morphology of Micronereis, as @ supposed

Representative of the Ancestral Type of the Nereida.

In order to gain a satisfactory idea of the relationship of

Micronereis to the rest of the family Nereide, a proper under-

standing of the external structure of the head and its appen-

dages is, among other things, necessary. As these have not

been very clearly or fully described by previous workers, I have

taken some trouble to determine their relations in the Puget

Sound specimens. Owing to the small size of the animals and

the crowded condition of their appendages, this was a matter of

some difficulty. Specimens were examined microscopically, both

by reflected light and, after clearing, by transmitted light, and

serial transverse sections of the anterior region of one were also

eut with the microtome. The following description is based on

an examination by these methods.

Text-figure 2.

Micronereis variegata, Puget Sound.

Anterior extremity from below, showing peristomial cirri and first

five pairs of parapodia. X 45.

The first two setigerous segments bear parapodia consisting of

one lobe only (text-fig. 2) corresponding to the neuropodium

of the remaining feet. Anterior to these only one region 1s

distinguishable, viz., the cephalic or peristomial region, which

bears the two pairs of eyes on its dorsal surface. On each side

this region bears also two rounded knobs or humps, anterior and

posterior (the posterior being also situated rather higher on the

side of the head than the anterior). Each of these four knobs

bears two large subulate cirri, one dorsal, the other ventral.

The points of especial significance are :-—

(a) There is no trace of palps or tentacles.

(b) No separate prostomial lobe, as distinct from the peri-

stomium, is developed. The eyes are situated on the

dorsal surface of the head in the same transverse plane

as the peristomial cirri.

246 MR. L. N. G. RAMSAY ON

The above views differ from those of Claparede in several

particulars. This authority considered that the first pair of

parapodia was borne by the buecal segment, and that the first

two pairs of parapodia represented the notopodia of the rest.

As to the first point, I think it is plain, both in the case of

Major Elwes’ specimen and in the Puget Sound material, that

the first pair of feet arises from the segment next behind that

which bears the tentacular cirri. They are, however, directed

forward in such a way that it is not unnatural that even a

careful observer should conclude that they arose from the buccal

segment, if one takes into account the minute size of the worm.

The transverse sections make this point still clearer. As to the

second point, the sections again show clearly that the first two

pairs of feet bear cirri on the under surface—not on the dorsal, as

would be the case if they were notopodia. Further, these feet

plainly arise from the same level as the neuropodia of the

following ones, as can be seen both in the sections and in the

complete specimens.

Text-figure 3. 5 Text-figure 4.

Text-figure 3.—Micronereis variegata.

Transverse section (oblique) through anterior region, showing one of the posterior

pair of tentacular cirri (Z’.C.) and their common base, and the first parapodium

(Z., on the left); on the right, Z., IL., III., IV. indicate the first four para-

podia. Vasc., vascular tissue of same. X 56.

Text-figure 4.—WMicronereis variegata.

Oblique section through the 2nd parapodium, showing the ventral cirrus.

X about 137.

These points have an important bearing on the systematic

position of the genus.

Claparéde also suggested that the parapodial cirri were more

of the nature of ligules (‘Ziingelchen”) than true cirri, on

account of their position and form. This view, however, seems

to me to be quite unnecessary. The cirri have the position, size,

and shape of true cirri, and in their cytological structure they

also agree. (I have compared the sections with those of Lepto-

nereis from Plymouth, which possesses typical nereid parapodia ;

the only cytological difference between the ligules and cirri in

Leptonereis is in the presence of numerous blood-capillaries in

the former, and their absence in the latter; otherwise they

both consist of epithelial tissue and large glandular cells. The

ANNELIDS FROM THE N.E. PACIFIC. 247

parapodial cirri of Jicronereis exhibit the same structure so

far as can be seen from the material available, which was not

preserved for cytological examination.)

With regard to the four pairs of tentacular cirri, Claparéde

stated that he was unable to decide whether some of these

should not be regarded as tentacles (‘‘ Kopf-ftihler”). As has

been indicated by McIntosh, these should probably all be

regarded as cirri corresponding to the four pairs present in other

nereids.

We are now in a position to compare MMicronereis morpho-

logically with other nereids. According to the above revised

views as to the head-parts and parapodia, it would seem that

we have in Micronereis a surviving descendant of the primitive

ancestral type from which the successful nereid line of to-day has

arisen.

The most characteristic features of this well-marked family

are to be found in the peculiar, lobed, biramous parapodia, the

well-developed prostomium or cephalic lobe with its palps and

tentacles, the four pairs of peristomial cirri, and the armature

of the proboscis with conical papille. All these features can be

directly derived from the Micronereis-type. This I shall now

endeavour to explain :—

Micronereis as a@ True Nereid.—The right of this genus to

be included in the family Nereide is upheld by the following

characters: the body-region bears on each segment a pair of

deeply cleft biramous parapodia, each ramus supported by a

Text-figure 9.

—

= ca

é SS =

Micronereis variegata. 10th parapodium, X 45.

lance-like aciculum and giving rise to a series of compound

homogomph spinigerous sete of typical nereid form, and bearing

a subulate cirrus on its external* surface. The first two

pairs of parapodia are incomplete, the dorsal division (noto-

podium) with its spine and seta-bundle being entirely absent.

Thirdly, the head-region bears dorsally two pairs of well-

developed eyes situated at the angles of a trapezium, and

* T. e., respectively dorsal and ventral in the noto- and neuro-podium.

248 MR. L. N. G. RAMSAY ON

laterally on each side two pairs of large tentacular civ.

Fourthly, the eversible proboscis is provided with a pair of curved

toothed jaws.

Micronereis as a Primitive Nereid.—The primitive, undiffer-

entiated condition of the genus is indicated by the following

points: in the parapodia, the two rami consist of a simple

setigerous lobe and cirrus alone, without the development of

“lips” and ligules of the more advanced nereids. The occur-

rence in one of the Puget Sound specimens of simple setz in

addition to the compound spinigers may also be a primitive

feature, as may the absence of falcigers. The prostomium

and peristomium are fused*, or to put it in another way, 20

distinct prostomial lobe has been differentiated from the buccal

“ segment,” and, further, palps and tentacles have not begun to

be acquired. Thirdly (and this I mention with some doubt,

having had no opportunity of examining the everted proboscis),

the proboscis is unprovided with papille, whether soft or hard.

Advances made by other Nereide.—The main ways in which

other Nereidz have progressed are in the differentiation of the

parapodia, the arming of the proboscis with papille, usually

cornified and hard, and the differentiation of a distinct pro-

stomial or cephalic lobe, and the development of two pairs of

sensory appendages (palps and tentacles) borne by this lobe.

Text-figure 6.

Parapodium from mid-region of Nereis kerguelensis, for comparison.

X 20.

As to the parapodia, the more advanced type is easily to be

derived from that of Micronereis by the gradual development and

elaboration of subsidiary protuberances on the surface of the

original simple setigerous ramus. The setigerous lobes of the

typical parapodium will thus represent the apices of the original

rami of the ancestral type, while the elongate ligules and the

smaller “lips” are secondary developments from the sides of

* As Quatrefages has pointed out (14, p. 578).

ANNELIDS FROM THE N.E. PACIFIC. 249

the same. In Dendronereis, this lobation of the rami is carried

still further (apart from the dendritic development of the

cirtl), while in Ceratocephale and Tylorhynchus the process has

probably not gone so far.

Coming to the first two pairs of parapodia, we meet with a

ditficulty, In these, in the higher forms, the notopodium,

although without the setigerous lobe with its spine and bristles,

is nevertheless represented by a single ligule and a dorsal cirrus.

Well, this is a difficulty, and for the present I shall leave it

as such. I hope to elaborate this theory further in a larger

work.

Text-figure 7.

FC,

KO PERL +PRO.

Poly

ae Es,

Bre

A. Sagittal section of anterior region of Nereis pelagica, Plymouth.

X about 6.

B. Micronereis variegata. Sagittal section of anterior region.

(Diagrammatic.) XX about 50.

PRO., prostomium; PERL, peristominm; T., IT, first and second, seti-

gerous segments; Z., tentacle; Plp., palp; H., position of eyes; 7.C.,

position of tentacular cirri; Pp.Z., position of first parapodium.

The severed surfaces are indicated by shading,

And now we come to the prostomial lobe of the higher forms.

This would seem to have been derived from the condition in

Microneveis by the forward elongation and constricting off of the

upper median surface of the head-region, in such a way that

the eyes are carried forward so as to be included in the con-

stricted-off lobe. Then on the anterior margin of this lobe

palps and tentacles have been developed, in response to the need

for more efficient sense-organs consequent on the enlarged ac-

tivities and “ wider outlook on life” which the higher Nereidze

have probably assumed,

250 ON ANNELIDS FROM THE N.E. PACIFIC.

Text-fig. 7, A, B, will help to elucidate the foregoing explana-

tion of the homologies of the cephalic region in the two types.

Conclusion.—According to the foregoing reasoning, Micronereis

should be regarded as a more or less unaltered representative of

a primitive ancestral form of the Nereide. - It does not appear

to me that the features of Aficronereis can be regarded with as

much probability as degenerations from the more highly developed

type.

I have set forth these views of the facts with some difhdence,

owing to my studies having been mainly confined to the single

family, the Nereide. A wider knowledge of the nereidiform

group of the Polycheta might perhaps produce an alteration

of them.

REFERENCES.

- (1) Charareprs, E.—Beobachtungen itiber .Anat. u. Entwick-

lungsges. wirbelloser Thiere. Leipzig, 1863.

(2) Enters, E.—Die Borstenwiirmer. Leipzig; 1868.

(3) Gruss, E.—Middendorft’s Sibirischer Reise: Anneliden.

St. Petersburg, 1849.

(4) Harrrneton, N. R.—‘“‘ On Nereids commensal with Hermit-

Crabs.” Trans. N. York Acad. xvi. 1897.

(5) Izuxa, A.—Journ, Coll. Sci. Japan, xxx. 1912.

(6) Jounson, H. P.—“ Polycheta of the Puget Sound Region.”

Pr. Boston Soc. Nat. Hist. xxix. 1901.

(7) McINrosu, W. C.—Scientific Results of H.M.S. ‘ Challenger,’

Zoology, xu. London, 1885.

(8) Mcinrosu, W. C.—The British Annelids.—II. London,

1910.

(9) Maumeren, A. J.—“ Nordiska Hafs—Annulater.” Ofvers.

K. Vet.-Akad. Forh. 1865.

(10) Marexzeuuer, EH. v.—‘ Siidjapanische Anneliden (1.).”

Denks. K. Akad. Wiss. Wien (Math.-Naturw. IEG a) oxlite

1879.

(11) Moors, J. P.—‘‘ Polychzetous Annelids from the N. Pacific

Coast of N. America.” Proce. Ac. Sci. Philadelphia,

1908.

(12) Moors, J. P.—* Polycheetous Annelids from California.” Jd.

1909.

(13) Moors, J. P.— Polychetous Annelids dredged by U.S.S.

‘ Albatross’ in 1904.” Jd. 1911.

(14) Quarreraces.—Histoire des Annelés. Paris, 1865.

(15) TRrEapwELL, A. L.—‘‘ On the Heteronereis Stage of J. kobi-

ensis McIntosh.” Biol. Bulletin, Woods Hole, Mass.,

ix. 190d:

(16) TReapwet, A. L.—‘ Polychetous Annelids of the Hawaiian

Islands.” Bull. U.S. Fish Comm. xxii. 111. 1903.

(17) Turnsuit, F. M.—‘ On the Anatomy and Habits of Werezs

virens.” Trans. Connect. Acad. 11. 1875.

JRUOLO AG OREO All INO SAINI ele Neti GC Incl whOk INUO ME SMILES) ISIN)

TOpao T syatT Tor hor)

tli he

am \ =

A SS)

KN ANY SF aan SS SS

A \ = BARNS t

eNO Od WMS IES) Zl

ra!

WOOILLOYVLINVY VNWNVYEDVHNaId 10 INF OMREIEEDNS Is Ce 8

‘mOpuoTy sya y YyaMyY

ic NGO he wlet Sy cl

—

ON A TELEOSTEAN FISH. 2

16. The Structure and Development of the Caudal Skeleton

of the Teleostean Fish, Plewragramma antarcticum. By.

A. Kyyvert Torton *.

[Received December 17, 1913: Read March 17, 1914. |

(Plates I. & II.)

INDEX. Page

Mionpholo cyt istiakea: 2 aa: neuen ieee ae watneciaeat ate EOS!

evelopment. hese a Aare Lh etgae aeea espana ann OL

In this paper I have recorded several points of interest in

connection with the development of the vertebral column of a

Teleostean fish, Plewragramma antarcticwm. My material con-

sisted of a collection of the post-larval stages collected on the

Southern Cross Expedition, and handed over to the Imperial

College of Science for description on behalf of the Trustees of the

Natural History Museum by Professor Jeftrey Bell, to whom 1

here wish to express my thanks. ‘lo Professor EH. W. MacBride,

F.R.S., V.P.Z.S., at whose suggestion I undertook this piece of

work, and under whose supervision it has taken its present form,

and to Mr. H. G. Newth, for very much useful help, my thanks

are due and are very gratefully tendered.

Unfortunately the Plewragramma material does not contain

stages quite late enough to show the actual formation of centra,

though the largest specimens are about 4°5 cm. long,

I believe that no specimens of this fish have been taken inter-

mediate in size between those at my disposal and certain speci-

mens—adults 16-17 em. long, taken from the stomachs of Seals,—

one of which Mr. Tate Regan very kindly allowed me to examine,

together with a specimen of Zrematomus newnesii, a closely allied

species in which the notochord is relatively smaller and the centra

better ossified than in Plewragramma antarcticum.

The fixative used in most cases appears to have been formalin

or spirit, so that the material is not very good from a histological

point of view.

Pleuragramma antarcticum is a specialised Teleost of the family

Nototheniide, the genus differing from the related Antarctic

genus Zrematomus chiefly in the feeble ossification of the skeleton.

Boulenger included Pleuragramma in the family leptoscopide,

but Regan has pointed out that it does not resemble Lepto-

scopus, but, on the other band, is very near Zrematomus. Most

of the Nototheniiform fishes are littoral and feed on crus-

taceans, molluses, ete. Zrematomus newnesti and Pleuragramma

antarcticum are amongst the more southern types, and appear to

be circumpolar (cf. Regan, Trans. R. Soc. Edinburgh, xlix. 1913,

pp. 251, 257, 264, &e.).

* Communicated by Prof. E. W. MacBripg, D.Se., F.R.S., V.P.Z.S.

+ For explanation of the Plates see p. 260.

252 MR. A. K. TOTTON ON

The caudal skeleton of an adult specimen of Plewragramma

antarcticum consists of a number of relatively large cylindrical

-bony centra of a papery nature. To the terminal one are

attached two expanded hypural bones, whose development proves

that they are compound (i. e. that they represent more than two),

and a third ventral hypural which is not so expanded. Between

the latter and the ventralmost of the former two hypurals, the

terminal branches of the caudal artery and vein come forth on

each side ina notch, This terminal centrum bears no arches, but

lying dorsal to it and the nerve-cord (not represented in PI. IT.

fig. 12) are two bones, the epiural apophyses (cf. Huxley, Q. J.

M.S. vol. vu. p. 41). Development shows that there were

originally three, and there are even indications of a fourth. The

penultimate centrum at first sight appears to be carrying two

neural and two hemal arches, but in the case of the neural arches,

at any rate, development shows clearly that they have arisen

through the division of one cartilaginous “ Anlage.”

I should like to make a digression here to discuss at greater

length this phenomenon of a centrum bearing two neural or

hemal arches. I have collected a few cases from various authors,

and have observed the phenomenon also in a few specimens in

which it has not been described, so far as I am aware.

Mormyrus kannume has two hypurals on the penultimate

centrum (¢f. Whitehouse, “Caudal Fin of Teleostomi,” P. Z.8.

1910, p. 596). I am not aware that the development of this

fish is known. It would be interesting to see whether the adult

condition was brought about by a division of a single cartilaginous

‘“‘ Anlage” or by a crowding together and displacement of arches,

as is more probable.

Lotz (Zeitschr. f. wiss. Zool. Bd. xiv. 1864, p. 88) gives the

following description of Cotéus gobio :—* Der vorletzte Wirbel

hat nur das Eigenthumliche, dass sein oberer Dorn sehr breit

und oben gespalten ist, so dass er aus zwei verschmolzenen

Dornen zu bestehen scheint.” The figure shows a much expanded

neural and a hemal arch on the penultimate centrum, and the

neural arch shows a line of division into two. I think this is a

clear case of ‘ division.” Moreover, ‘ division”? seems to be

connected in some way with expansion, as comparison with other

cases will show. He says of Barbus fluviatilis :— Der drittletzte

Wirbel besitzt in der Regel zwei obere Bogen deren Dornen

bereits als Stiitzen der kurzen Flossenstrahlen dienen, diess ist

iibrigens nicht constant; es kann auch nur ein oberer Bogen

vorhanden sein oder es kann der vorletzte Wirbel deren zwei

besitzen, wihrend er in der Regel nur einen besitzt, dessen Dorn

mit einem knorpeligen Ende versehen ist” (p. 86). On page 95

he says of Salmo salar :—‘“‘ Am. 17-18. Tag. wird tiber dem

Neural-canal hinter dem letzten normalen oberen Bogen (g) der

erste falsche Dorn (4) gebildet.” The ‘‘falschen Dornen” are

the epiural apophyses. It is to be noted that the neural and

heemal arches have already been formed. Lotz continues: ‘“ Oft

A TELEOSTEAN FISH. 953

der vordeste der falschen Dornen mit dem Dorn des letzten

oberen Bogen verschmilzt oder es ist ein ganzer iiberziihliger

Bogen vorhanden, so dass dann der viertletzte Wirbelkorper

zwei obere Bogen erhiilt.” The “ Wirbelkérper” appear towards

the end of the fourth month, doing so first in front.

O. Hertwig (‘ Entwickelungslehre der Wirbeltiere, III. 1.

pp. 456-82), in the account of the development of the vertebral

column of Teleosts by Schauinsland, says :—‘ Finally, it must be

pointed out, too, that there are vertebre in the tails of many

Teleosts, with fully formed and equally developed double upper

(or also both upper and lower) arches, the cartilaginous rudiment

of which can sometimes even be shown.” It is not very clear

what he means by the last statement. Does he mean that there

isasingle “ Anlage” or that there are two? He continues:

“These vertebre can be regarded: either as corresponding to

only two sklerotome halves, in which case, however, the arches

belonging to each half have become exceptionally developed to

an equal extent, or (which is more probable) as having arisen by

the later union of the two complete vertebre ; that is to say,

at least three, perhaps even four, sklerotome halves participate,

In these cases, then, there would be real (secondary) ‘* Diplo-

spondyly.”” He does not bring forward any embryological evi-

dence to prove his case, except the above-mentioned statement

of doubtful meaning. He refers to a figure (270) of the

“Dorsch ” (Gedus morrhua) as showing double upper and lower

arches on the penultimate centrum, but I believe there is some-

thing peculiar about the caudal fin-skeleton of Gadus which has

not been described, with reference to these double-arch bearing

centra. I understand that the appearance is due to partial

fusion of arches and radials.

T myself have met with the phenomenon of a centrum bearing

double arches in the case of Trematomus newnesri, where there

are two very well-defined neural arches on the penultimate

centrum (see Pl. IT. fig. 13); also in the case of 7. borchgrevinki

(see fig. 14), where, curiously, it is the antepenultimate centrum

which bears double upper arches and double hemal arches as

well. The penultimate centrum has greatly expanded arches,

and, indeed, there is some slight indication of a division in the

neural one, but the specimen was a dried skeleton and one not

entirely free from flesh, so that I had some difficulty in making

an accurate observation. In Votothenia macrocephala the arches

of both antepenultimate and penultimate centra are somewhat

more expanded than the rest, but only the penultimate neural

arch shows any sign of division. J examined a number of allied

forms, but did not come across any more cases of centra

with double arches. I may mention that my drawings of the

skeletons of the adult Pleuragramma antarcticum and Trema-

tomus newnesit were made from specimens dissected very carefully

in spirit, and from which every particle of flesh had been removed,

and as much connective tissue as possible also.

254. MR. A, K. TOTTON ON

I will now return to my description of the caudal skeleton of

Pleuragramma untarcticum.

One of the chief characteristics of this skeleton is the large

size of the cartilages supporting the procurrent rays. No doubt,

this 1s connected with the weakness of general ossification. The

caudal skeleton of an adult specimen of Zrematomus newnesit,

though generally similar to that of Plewragramma, differs in

several points. As mentioned above, the general ossification is

much stronger. Correlated with this probably is the smaller

relative size of the cartilages bearing the procurrent rays. The

epiural apophyses are three in number. The two terminal

hypurals still retain something of their compound nature; finally,

the penultimate centrum bears a well-defined double neural arch,

but a single unexpanded hemal arch. The free end of the noto-

chord extends further, and that of the nerve-cord not so far as

in Plewragramma.

In specimens of Pleuragramma antarcticwm of about 8 mm.

(Pl. I. fig. 1), the tip of the tail and the notochord are bent

slightly ventralwards, the angle thus formed being filled up by

skleroblastic tissue (sk). - This latter consists of mesenchyme-

cells, in which the cytoplasm has become clearer and the nucleus

more distinct (7.e. takes up stain more readily) than in the cells

of the surrounding tissue. The continuous fin-fold round the

tip of the notochord is expanded and supported by actinotrichia.

Whether this bending down of the tip of the notochord is appa-

rent in the living animals or not I do not know. It certainly

appears in large numbers of fixed specimens. One is reminded,

in connection with this, of the prolongation of the vertebral

column into the lower lobe of the vertical caudal fin of the

Ichthyosauria. I think the condition I have described can

searcely be due to fixation, because, since the skleroblastic tissue

is denser than the tissue dorsal to it, the tendency of the fixative

would be to shrink the upper tissue more than the lower, and so

turn the tail upwards instead of downwards. I suggest that the

presence of the skleroblastic mass under the notochord gives rise

to some sort of stimulus causing this flexure. 1 should like to

call attention to the fact that Ryder (U.S. Comm. of Fish &

Fisheries, 1884, p. 1057), in describing the development of Alosa,

says :—‘ Here the development of the hypurals is accompanied

by a pressing inward of the ventral wall of the chorda.” He

gives a figure (fig. 2, pl. i1.) to illustrate his point. I think that

the illustration would be more correctly interpreted as showing

a ventral flexure of the chorda, since the dorsal wall of it is also

involved,

Jn the next stage (-85 em.) (fig. 2), the skleroblastic cells in the

concavity of the notochordal flexure have become divided into

an anterior and a posterior mass, and a hyaline matrix has been

secreted around and between certain of them, and in this way

three blocks of cartilage are formed, two in the anterior mass

of skleroblastic tissue and one in the posterior mass. Lepido-

A TELEOSTEAN FISH. 255

trichia (/p.) have also been formed at the margin of these blocks

of cartilage.

Stage itl (1:05 em.) (fig. 3).—Darkly staining skleroblastic

tissue “(sk' .) has now made its appearance above the notochord im-

mediately dorsal to what will later become the ‘ventral hypural”

(a.). It is significant to note that just at this time the notochord

is becoming straight again. This secondary dorsal flexure is

started, I suggest, by some stimulus due to the appearance of this

new mass of skleroblastic tissue. More cartilage has now been

formed ventrally, in front of the two pieces already described

(Stage IT), in the anterior skleroblastic mass, which two pieces

have probably fused to form a, fig. 3. This new cartilage consists

of a pair of proximal pieces and a median distal piece. The

former, 6, represent a hemal arch and the latter is a radial.

That the proximal pair of cartilages represent, or are serially

homologous with, the arches which appear later and more ante-

viorly seems probable, because they correspond well enough in

size and in the manner of their appearance. I must confess that

1 am not quite satisfied that the proximal and distal parts are

actually separated, as the appearance of the line of separation

may be due to optical section. In any case, the fusion between

arch and radial in the next anterior segment is quite clearly seen

(figs. 5 & 6), so that the evidence for the fusion of the above-

mentioned elements is presumptive.

One may regard these cartilages, perhaps, as being in a con-

dition intermediate between that of the hypural behind, which

apparently is formed from the skleroblastic tissue as fused arch

and radial, and the condition of the arch in front, where fusion

of the originally separate elements can be seen as development

goeson. I may say here that the term “hypural” is limited to

those pieces of cartilage (which may later become ossified) which

represent or are actually made up of fused arches and radials.

The hypural (a, fig. 3), behind the one whose development I have

just traced, is a good deal larger than this latter, and is somewhat

rounded when seen laterally. It is beginning to form an anterior

peg which later on passes through the arch formed by the

proximal end of the hypural next in front, and lies just dorsal

to the caudal artery and vein which run in the arch. The

caudal fin is now beginning to be constricted off ventrally from

the median fin-fold.

Stage IV (fig. 4).—Examination of specimens 1:3 cm. long

shows that the upturning of the notochord has gone on further,

and produced from the straight condition seen in the specimens

1-05 em. long one in which there is a distinct dorsal flexure. The

epaxial elements (ep.), formed by the skleroblastic tissue mentioned

in the last stage, lie in the concavity thus formed. The posterior

hypural cartilaginous block (phy.) can now be seen to be made

up of two partially-fused blocks, or, rather, the original mass of

skleroblasts has secreted two masses of matrix, which are not

completely divided the one from the other. Similarly, the hypural

256 MR. A. K. TOLTON ON

in front (a.) is incompletely divided into two on the proximal

side, showing that it represents more than one hypural. There

is, in the specimen figured, a small piece of cartilage (£.) between

the “posterior” and “anterior” hypurals. This would appear

to be exceptional, and may be the vestige of another hypural.

Hemal arches are now beginning to appear anterior to the

hypurals, developing from behind forwards. The matrix secreted

by the skleroblasts diminishes In amount as one goes forward.

The lepidotrichia are still better developed and extend to the

margin of the fin. There is figured a row of lateral-line sense-

organs on each side, which extends on to the future mid-line

of the caudal fin, dividing the lepidotrichia into a dorsal and

a ventral series.

Stage V (1°55 cm.) (Pl. I. fig. 5).—There are now three dis-

tinct (a rudimentary fourth) epaxial elements (whose appearance

seemed to give the signal for the secondary flexure of the noto-

chord). In transverse section they show no signs of forking at

their proximal ends, and they originate from single median

masses of cartilage, not from paired pieces as do the arches.

They are much eloser together than the neural arches, which are

now appearing anterior to them, much in the same way as the

hemal arches arose. There is a gap between these epaxial

elements and the neural arches. These elements, moreover, stand

quite clear of the nerve-cord, not arching it over as the neural

arches do. They probably correspond to Huxley’s epiural apo-

physes in G'asterosteus (Huxley, Q.J.M.S. vol. vii. p. 41). The

fusion between the neural arch and radial, described as possibly

existing in Stage III, is now complete: there is no line of division

between the matrix of one and that of the other. A slight notch

has appeared in the posterior border, and it is here that the

caudal artery and vein run out on either side. A radial (7.) at

the distal end of the hemal arch next anterior to the one just

described has been formed at this stage, but no fusion of the two

elements has yet taken place.

In Stage VI (1'8 cm.) (fig. 6) there is still no sign of the

cartilages to which the procurrent caudal fin-rays are attached

in the adult. A typical neural arch, taken further forward than

those shown in fig. 6, extends at this. stage through a length of

about 136. It consists of a pair of cartilages lateral to the

nerve-cord. Their bases do not reach the notochord, and their

thickness in transverse section 1s about 8p. The hemal arches

also consist of pairs of cartilages which do not reach up to the

notochord. They are each about 16 thick, and are separated

at their distal ends by about 16 » of closely packed skleroblastic

cells, which have not secreted any matrix as yet. Further

forward the cartilages are considerably smaller, restricted to the

sides of the caudal vein, and extend only through a length of

about 30. Posteriorly the skleroblasts between “the distal ends

of both dorsal and ventral pairs of cartilages have secreted a

matrix, and in this way the pairs of cartilages have become

A TELEOSTEAN FISH. ONS

fused. Their proximal ends have grown till they now stand on

the notochord.

The hypurals are now beginning to expand laterally at their

proximal ends, so as to form a broad seat for the convexity of the

upturned notochord, so that the thrust of the tail is more widely

distributed. Fusion has taken place between the hemal arch

and radial (7.) described in Stage V. he epaxial elements

behind the neural arches have undergone a certain amount of

fusion, so that there is now a smaller posterior and a larger

anterior cartilage. The caudal artery and vein are both forked

at their posterior ends to pass round the large ‘ventral hypural,”

which is not arched, to admit of their passage between it and

the notochord. Fig. 6 is a reconstruction of this stage from a

series of transverse sections.

Stage VII (2-2 em.) (fig. 7),— Long narrow cartilaginous pieces

(car.) have now made their appearance, one dorsally to the two

posterior neural arches and the three original (apparently now two)

epiural apophyses described above, and another ventral to the most

anterior hemal arch. They give articulation to a dorsal and a

ventral group of procurrent lepidotvichia. From the shape and

position of this ventral piece of cartilage, it appears possible that

it may be the homologue of the two radials or hypural apophyses

which appeared just behind it and underwent fusion with two

hemal arches. The anterior ends of both these new cartilages

develop into two or three smaller and more or less separate

cartilages. The anterior peg of the “ventral hypural” now

extends between the proximal ends of the next anterior hypural,

with which it eventually fuses, giving rise to the appearance in

transverse section of a fusion of the halves of the arch themselves,

and making in fact a small bridge, dorsal to the caudal artery

and vein, which has rather a puzzling appearance.

The division between the halves of the “dorsal hypural”

(d, Pl. II. fig. 8) has now disappeared, or, in other words, the two

uppermost hypurals have fused (Stage VIII), and a certain

amount of absorption of the two large hypurals seems to be taking

place, which, together with growths in other points, such as the

posterior margins and the proximal ends, causes a considerable

change in shape. Between Stages II and VII the rate of growth

of the hypurals is greater than the rate of increase in girth of

the notochord; but between Stages VII and X there is a reversal

of this disparity which culminates in the condition where the

notochord has the relatively enormous proportions so character-

istic of the latest available postlarval stages of this fish.

Stage VIII shows the first step in this inflation of the notochord,

but here, as in later stages, the upturned tip or Chordastab

is not affected. In this respect Teleosts show an important

difference from Elasmobranchs (see Schauinsland, p. 462). Figs.

9 and 10 ave drawn under the binoculars with camera lucida,

and show a dissection of the posterior end of the notochord,

nerve-cord, and cartilaginous elements of a fish 4:2 cm. long.

258 MR. A. K. TOTLON ON

Fig. 9 is practically a lateral view and fig. 10 a dorso-lateral

view. It will be seen that the posterior neural arches are much

further developed than the anterior ones.

Reference to the adult fish (fig. 12) shows that the neural arch

has, on each side, two points of attachment to the centrum,

between which points the segmental nerve emerges. Hxamination

of Stage IX proves that the primary point of attachment is the

anterior one; the process of formation of the posterior one—as a

backwardly directed spur (sp.)—can be seen in figs. 9 and 10.

A comparison of these backwardly directed spurs with some

cartilages described by Schauinsland in Hertwig’s book (p. 467)

shows that it is probable that they represent intercalaria.

Schauinsland says: “The bulk of the base of the arch (carti-

laginous stage) is at the cranial end of the vertebra, but this base

sometimes extends caudalwards (Pike). The caudal part may

even be cut off to form a separate piece of cartilage (e. g. in the

tail of the Trout). This,” he considers, “ should be looked upon,

probably in the case of the Pike and pretty certainly in that of

the Trout, as the remnant of the second arch and as homologous

to the cranial arch of Amia” (i. e. it is formed by the cranial half

of a sklerotome: for this reason it is on the caudal end of a

vertebra).

Stage X.—The posterior edges of the three large hypurals

(fig. 11, 4:5 em.) have grown considerably, owing to additions from

the large enveloping cap of skleroblasts (not shown in figure) on

which the lepidotrichia are seated. The proximal ends of these

hypurals have grown also with the notochord, so that a large gap

(g, figs. 9-11) has appeared—and continually increases in size—

between the dorsal and ventral hypurals. It will be noticed

that the posterior neural arch is dividing into two.

The adult specimen I dissected was 16°5 cm. long. The centra

are very ‘‘ papery,” and constrict the notochord only very slightly.

The condition of the posterior neural arch is interesting. At

first sight the penultimate centrum appears to be carrying two

arches. Comparison with previous stages, however, shows that

this is not so, but that the appearance is due to the fact that the

division, incipient in Stage X, has now been completed. As a

result of this, the segmental nerve, which in Stage 1X emerged

beneath the undivided arch, now lies in the cleft, and a secondary

ossification beneath the nerve has finally reunited the two half-

arches at their points of attachment to the centrum. Similarly,

the hypural carried by this centrum has divided, and the two

halves are covered by confusing secondary ossifications.

The double neural arch in Plearagramma antarcticum and the

arch immediately in front of it are considerably more expanded

and better ossified than those in front of them.

The posterior part of the hypural borne on the penultimate

centrum of Pleuwragramma antarcticwm has a thick, well-ossified,

posterior edge, especially thick at the tip, while the rest of it

is very thin and membranous. The hypural behind the one just

A TELEOSTEAN FISH. 259

mentioned has similarly a thick, well-ossified, posterior edge and

proximal portion, whilst the rest 1s very thin and membranous.

The dorsal and ventral hypurals, which give attachment to the

majority of lepidotrichia, are now widely divergent, thin, and

membranous. These two, together with the hypural immediately

in front, are ankylosed to the last centrum. Of the two epiurals

whose development I have traced above, the posterior one is well

ossified and thick, while the anterior one has a thick, well-ossified

posterior edge and a thin, membranous, expanded anterior edge.

The uptur ned tip of the notochord is quite free and naked,

reaching about halfway up to the anterior margin of the dorsal

hypural (d.). The upturned nerve-cord also runs up to about

this level, beyond which J have failed to trace it in the dissection

T have made. The cartilaginous elements to which the fin-rays

ave attachel have grown a good deal by this time, but have not

become ossified.

SUMMARY.

1. The development of the vertebral column begins at the

caudal end, the hypaxial elements being the on to appear.

This eoincides with a down-bending of the notochord (compare

with this the analogous condition of the vertebral column in the

Ichthyosauria). Epaxial elements do not appear until this con-

dition has given way to the straight condition again. Arches

appear as paired cartilages at the sides of the caudal arter y and

vein and of the nerve- weed They are separated from the

notochord by connective tissue.

2. The notochord is of a relatively enormous size, and persists

with only slight constriction throughout life. The centra consist

of thin papery lamelle of membrane-bone. Ossifieation is

generally weak.

3. The neural and hemal arches of the penultimate centrum

are double, owing to the splitting of single rudiments. Similar

phenomena are to be observed in other fishes. ‘They may be

produced by different causes :—

(1) Splitting of an originally single rudiment.

(2) Crowding of two arches on to one centrum.

(3) Fusion of the first epiural apophysis with the last neural

arch.

(4) Exceptional equal development of both arch and inter-

calary.

(5) Secondary diplospondyly, ¢. e. fusion of two centra.

4. Large cartilages are present above and below the last two

centra in the adult, which support a dorsal and a ventral series of

procurrent fin-rays. Their great size is probably connected with

the weakness of general ossification.

5. The hypural bones of the adult are formed by a fusion of

hemalarchesand radials. This compound nature of the hypurals

Proc. Zoou. Soc.—1914, No. XVIII. 18

260 MR. A. K. TOTTON ON

may be seen anteriorly in adult Selachians and in the Sturgeon,

but it has not been shown before, as far as I am aware, in

Teleosts.

BIBLIOGRAPHY.

(See O. Hertwig, 1906. Jena. Entwickelungslehre der Wirbeltiere,

Bd. III. Teil 2, Kap. vi. pp. 456-82.)

Srannius. 1854. Handbuch der Anatomie der Wirheltiere,

2 Aufl. 1854. (Siebold u. Stannius, Handbuch d. Zootomie,

Theil ii. Heft i. Berlin, 1854-56.)

A. Kouurker. 1860. Ueber das Ende der Wirbelsaiule der

Ganoide und einiger Teleostier. Leipzig.

Lorz. 1864. Ueber den Bau der Schwanzwirbelsiule der

Salmoniden, etc. Zeitschr. wiss. Zool. Bd. xiv.

Grasst. 1883. (Développement de la colonne vertébrale chez

les poissons osseux. Archiv. Ital. biologie, t. iv. 1883.)

Lo svilluppo della colonna vertebrale ne pesci ossei. Reale

Acead. dei Lincei, ser. 3, Memorie, vol. xv., 1882-83.

Scueet. 1893. Beitriige zur Entwicklungsgeschichte der

Teleostierwirbelsiiule. Morphol. Jahrb. Bd. xx. Leipzig.

vy. Epner. 1896. Ueber die Wirbel der Knochenfische und die

Chorda dorsales der Fischer und Amphibier. Sitzungsber.

d. k. Akad. d. Wiss. Wien, Math.-Nat. K1., Bd. ev.

Ussow. 1900. Zur Anatomie und Entwickelungsgeschichte

der Wirbelsiule der Teleostier. Moskva, Bull. Soc. Nat.,

Ney Seplve ae ullis0))

Auprecut. 1902. Zur Ent. des Achsenskelettes der Teleostier.

Tnaugural-Dissertation d. math. und nat. Fak. d. Kaiser-

Wilhelms-Univ. zur Erlang. d. Doktorwiirde. Strassburg.

A. Acassiz. 1878. On the Young Stages of some Osseous

Fishes. Boston, Mass., Proc. Amer. Acad. Arts Sci.

Ol stusl, 1On WINE

ea elpxennve 8092 %@). Ji. Mess vole wie so.a4 Le

J.A.RypEr. 1884. Washington, D.C., Rep. U.S. Comm. Fish.

SOD.

1a H. Wuitrnouse. 1910. Caudal Fin of Teleostomi. P. Z.S8.

p. 990.

C. T. Regan. 1913. Trans. Roy. Soc. Edinb. xlix.

EXPLANATION OF THE PLATES.

PuateE I.

Fig. 1. Tail end of a specimen of Pleuragramma antarcticum, 8 mm. long. The

specimen was stained, cleaned, and mounted whole in Canada balsam.

The skleroblastic tissue, sk, is represented semidiagrammatically by dots.

Notice the down-bending of the tip of the notochord.

2. Tail end of a specimen of P. antarcticum, °85 em. long. Camera-lucida

sketch from a whole mount. Notice down-bending of the tip of the

notochord, the formation of definite cartilaginous elements, a’, a’’, by the

skleroblastic tissue, and the appearance of lepidotrichia, 7p.

A TELEOSTEAN FISH. 261

. Camera-lucida sketch of the tail of a specimen of P. antarcticum, 1:05 cm.

long. Notice the appearance of skleroblastic tissue, sk’, dorsal to the noto-

chord, accompanied by a slight up-bending of the tip of the notochord.

a, a fusion of cartilages marked a’ and a” in fig.2. 6, a hemal arch.

c, a radial.

. Tail of a specimen of P. antarcticum, 1:3 cm. long. Definite cartilaginous

elements, ep, have been formed dorsal to the notochord, which has now

a marked dorsal flexure. Notice the row of lateral-line sense-organs

extending on each side along the trunk and on to the dorsal fin, marking

its future mid-line. phy, posterior hypural. k, an abnormal piece of

cartilage.

. A whole mount of the tail of a specimen of P. antarctiewm, 1°55 cm. long.

The dorsal flexure of the notochord has reached its maximum. Neural

arches, na, have begun to appear. The figure shows three stages in the

formation of a hypural. Anteriorly there is a heemal arch, behind it is

another hemal arch with a radial, 7, at its distal end. The two elements

have not begun to fuse. Behind these again is a hypural (for description

of which see Stage III).

. A reconstruction from a series of sections of the tail of a specimen of

P. antarcticum, 1°8 cm. long. (Sections posterior to “dorsal hypural”

lost. Tip of notochord and nerve-cord hypothetical.) The cartilaginous

elements are represented as seen in a seetiou near the sagittal plane.

Notice the fusion that has taken place between radial and hemal arch

seen in fig. 5.

. A reconstruction from a series of sections of the tail of a specimen of

P. antarcticum 2°2 cm. long. Notice the anterior peg, p, of the “ ventral

hypural” which passes between the notochord and caudal vein and

artery, fusing with the proximal portion of the hypural in front of it.

Notice also the cartilaginous elements, car, which later on give attachment

to the anterior dorsal and ventral procurrent fin-rays. An asterisk marks

the exit of the caudal artery and vein from the hemal arches.

Puate If.

. A dissection of an “odd tail” of Plewragramma antarcticum, probably

2°8 cm. long. It shows the uniform nature of the “ dorsal hypural,” d,

(the compound nature of which has been seen in previous figures). It

will be noticed that the posterior neural arch, », is expanding at its distal

extremity.

. An antero-lateral view of a dissection of the notochord and cartilaginous

elements of the posterior end of a specimen of P. artarcticum, 42 com.

long. It was drawn under Zeiss binoculars with the aid of a camera

lucida. Note the increase in development of the neural arches from

before backwards. The sketch illustrates the way in which the neural

arches grow over the segmental nerves. sp, backward growing spurs.

g, gap between dorsal and ventral hypurals.

. A latero-dorsal view of the specimen figured above (fiz. 9). It will give a

better idea of the shape of the neural arches. e, f; the two halves of a

neural arch.

. A camera-lucida sketch of a dissection (mounted in balsam) of the noto-

chord and cartilaginous elements of the posterior end of a specimen of

P. antarcticum 45 cm. long. Notice the incipient division of the

posterior neural arch into two. A considerable change in the shape of

the two large hypurals has taken place (see text, p. 257).

A dissection of an adult speeimen (16°5 cm.) of Plewragrammea antareticum.

The tip of the notochord is represented as being naked. The attachment

of the large hypurals is rather broken in this specimen (it had been

devoured by a Seal), so that it is quite possible that the tip of the noto-

chord was ensheathed by thin membranous bone. The drawing was made

with the help of a camera lucida aud Zeiss binoculars. d, posterior or

dorsal hypural. a, anterior or ventral hypural.

. A dissection of a specimen of Trematomus newnesii, 6 cm. long (adult ?).

It is given for comparison with fig. 12. The tip of the notochord is naked

and extends out between the caudal fin-rays.

. A diagram of the caudal skeleton of Trematomus borchgrevinki.

18*

ON TWO NEW SPECIES OF CESTODES.

(or)

17. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea. By Frank E. Bepparp, M.A.,

D.Sc., F.R.S., F.Z.S8., Prosector to the Society.

[Received February 10, 1914: Read April 7, 1914. ]

(Text-figures 1-8.)

XIII. On Two NEw SPECIES BELONGING TO THE GENERA OOCHO-

2ISTICA AND ZLINSTOWLA, WItH REMARKS UPON THOSE

GENERA.

INDEX.

Page

IDMPSAOUOE, (HAVE, SDo We scoocoss. ss05ess0ac0ea0snHD0R9 HDG DR AOOSODDOD ACL)

Oochoristica marmoseé, sp. N. ........ aubueeadade {i ee)

Remarks upon Genera Oochoristica ail Dinstowte |. MASI!

The material upon which the following observations are based

was collected at the Society's Gardens during September and

October of last year, and suitably preserved for microscopical

examination. I was only able to make observations upon the

living worm in the case of Oochoristica marmose. Both worms

were parasitic in the small intestine of their host.

§ LinstowIA AMEIV4, sp. n.

I have examined three examples of a Cestode from the Surinam

Lizard, Ameiva swrinamensis, which I place—atany rate, for the

present—in the genus Linstowia; for I shall call attention on a

subsequent page (p. 281) to the difficulty of distinguishing Lin-

stowia and Oochoristica as they have been defined up to the

present time.

The worm is a small one, measuring up to 25 mm. in length

with a diameter (where it is broadest) of 15 mm. There is

no rostellum and the four suckers are unarmed. The scolex

(when the worm, at any rate, is in a state of contraction) is

not wider than the body which follows; the latter gradually

increases in diameter. A neck is present in which no segmen-

tation occurs; the rudiments of the generative organs appear

almost coincidently with the commencement of segmentation.

The only other external character to be noted is the “alternation

of the genital apertures, which are always anterior in position

in the segment. In transverse sections the cortical layer is seen

to be thick, its diameter in both dorsal and ventral layers being

fully as great as that of the entire medullary layer in the same

section. This is one of the reasons which lead me to refer the

present species to the genus Linstowia as commonly defined.

Such sections also show the disposal of the longitudinal muscle-

fibres into bundles. As will be seen from text-fig. 4 (p. 270)

there is only one row of bundles, which is by no means so marked

as is that in two other species belonging to the same genus or to

264 DR. F. E. BEDDARD ON

Oochoristica, and which are represented in the same figure for

the purpose of a readier comparison. In Linstowia ameive the

bundles contain a varying number of fibres—from as few as two

or even one up to nine or ten. Midway between this inner

longitudinal layer and the subcuticula is an outer layer of longi-

tudinal fibres, which are largely implanted singly or in twos, and

everywhere form a much thinner layer than the inner row.

The water-vascular system is like that of other species of Lin-

stowia; but it is not always quite easy to interpret the facts.

In transverse sections I have seen the smaller dorsal vessel lying

above the ventral either directly or more obliquely ; and this in

quite neighbouring sections, in others of which the two vessels

lay side by side, the dorsal outermost, 7.e. nearer to the nerve-

cord. This is evidently due to the varying degree of contraction

of the proglottids, a fact also emphasized by the zigzag course of

the vessels when viewed in horizontal sections. I believe that

the normal position of these vessels is side by side. The inner

and larger of the two tubes, which I regard as the ventral vessel,

gives off several branches in each segment, which are themselves

branched. There also appear to be connections between the

ventral and dorsal vessels. I cannot, however, give a map of this

network.

The testes lie posteriorly to the vitelline gland, and reach

forward on either side of it; they do not, however, extend

laterally of the ovary. In a given segment the testes were visible

in 18 consecutive sections. The largest number counted in the

middle of the series was 43. I therefore calculate the total

number to be about 200. The testes do not extend laterally

beyond the lateral water-vascular vessels.

The genital ducts of this tapeworm open on to the exterior, as

already mentioned, alternately, with, however, no absolute regu-

larity in the alternation. They both open—the cirrus-sac in

front of the vagina—into a genital cloaca, whose external pore is

close to the anterior margin of the proglottid. This genital cloaca

is separated into two regions, which are obviously of difierent

morphological value. Outside is a funnel-shaped chamber, the

apex of the funnel lying to the inside. In longitudinal sections

through the cirrus-sac it was occasionally noted that the bottom

of the funnel was almost completely closed, leaving but a narrow

slit of communication with the inner chamber of the genital

cloaca. The latter is roughly spherical in form, and receives the

openings of the cirrus-sac and the vagina. Its depth is not very

different from that of the outer funnel-shaped region. In longi-

tudinal sections, it is seen to be marked off from the outer chamber

by a projection on either side which is the expression in such a

section of a circular fold. This fold is muscular and a thick

layer of muscles, of which it is a part, encircles the whole of the

spherical chamber of the genital cloaca. The cirrus-sae lies in

the same straight line with the genital cloaca, and is thus at right

angles with the long axis of the body of the worm. The cirrus-

TWO NEW SPECIES OF CESTODES. 265

sac is somewhat hourglass-shaped through being compressed in

the middle; but the anterior end, 7. e. that nearest the exterior

of the body, is not so wide as the posterior end. The cirrus-sac

has a thick muscular coat, which thins out only posteriorly ;

this end of the cirrus is thin-walled and globuJar in shape.

The cirrus-sac of this Linstowia is large, but not so large as

in L. echidne and L. semoni. In the ripe segments the total

length of the cirrus-sae and the genital cloaca is between j and

$ of the body diameter; more anteriorly, where the cirrus-sac

is not fully formed, but still differentiated from the sperm-duct,

a where the body is narrower, the length is between 7 and

3 of the diameter of the proglottid. The end of the cirrus-sac

lies considerably to the inside of the nerve-cord; it reaches the

level of the innermost of the longitudinal water- vascular trunks.

Text-figure 1.

Linstowia ameive.

Horizontal section of a proglottid in which the uterine cavities are not

yet developed.

g-c. Genital cloaca. O. Ovary between which and testes, ¢, lies vitelline gland,

not lettered.

T note, finally, that the lateral nerve-cord, where it is crossed by

the cirrus-sac, is bent outwards for a space. Soon after issuing

from the cirrus-sac the sperm-duct, at first only sinuous, forms a

coil; this region of the vas deferens is surrounded by laxer

parenchymal tissue than that which pervades the body generally,

but which presents no glandular characters.

The vitelline gland of this species presents the usual characters.

It lies at about the middle of the proglottid antero-posteriorly, as

well as from side to side. It is smaller in horizontal diameter

than the ovary, which lies in front of it, or than the mass of

266 DR. FE. E. BEDDARD ON

testes, which lies behind it. The vitelline gland is not in contact

with hs ovary, and between them lies the shell-gland. Ina

complete series of horizontal sections, it can be obser ell that the

vitelline gland extends further ventrally than any of the gonads

and that it nearly reaches the dorsal limits of the medulla. The

duct of this gland is comparatively wide where it leaves it

anteriorly, but soon narrows. The vitelline gland is later in its

growth than the ovary, for, in earlier proglottids than such as are

veferred to in the above deser iption, the vitelline gland is pro-

portionately considerably smaller than the ovary.

The ovary is single and as nearly as possible in the middle of

the body. Ibis, as is so usual, mainly developed laterally in two

wings. The ovary lies in front of the vitelline gland and of the

neeteee it is very near to the anterior border of the proglottid.

The ovary, as is usual, is not solid and compact, but frayed cut

laterally in a digitiform fashion. The ovary extends laterally

beyond the vitelline glands, but not so far as do the testes.

The female efferent apparatus is simpler than in many tape-

worms. It opens into the genital cloaca by an expanded funnel-

shaped mouth lying behind the entry of the cirrus-sac. The

vagina runs a straight course to about the middle of the proglottid,

where it bends posteriorly and becomes coiled before opening into

the shell-gland. There is no dilatation along its course—nothing

that can be described as a receptaculum seminis. Even in quite

ripe proglottids there is no change in the female duct, except a

slight diminution of the always narrow lumen.

§ Uterus and Embryos.

In this species, as in others referred to the genera Linstowia

and Oochoristica, a uterus exists for a space; but later the

embryos come to be implanted singly in the parenchyma of the

body.

One remarkable feature of the uterus in this species is its

sudden appearance. In one proglottid there is no trace that I

could discover of a uterus. In the immediately following pro-

glottid the uterus was present in a well-developed condition. Its

general appearance under a low power is shown in text-fig. 2

The ripe ova (ova at this stage, not embryos) are found over a

large area of the proglottid, but limited to the medullary region ;

indeed, they do not extend outside of the lateral trunks ofthe

water-vascular system. These scattered ova are found in front

of the ovary as well as behind it; they also occur among the

testes, where the latter begin to thin out ventrally. They are

thus mainly ventral in position, the testes being mainly dorsal.

Although the cavities lodging the ova have, so far, been spoken

of inferentially as a uterus, there is no continuous uterus in this

worm with its own definite walls. On a rapid gion at such a

section as that which is represented in text-fig. 2, the impression

given may be that the ova ave contained in a series of circular or

TWO NEW SPECIES OF CESTODES. 267

oval spaces which intercommunicate. A more careful examination,

however, under higher powers, does not entirely confirm such

an impression. ‘There are undoubtedly cavities of varying dimen-

sions in which one or more eggs often lie; these cavities are, to

some extent, of irregular shapes, and their prolongations may

anastomose. But there is no doubt that there are cavities con-

taining eggs which are entirely isolated from the others. There

is, In fact, nothing that can be described asa branched or retiform

uterus with a continuous cavity. And it is to be observed that

I am here speaking of the earliest proglottids in which the uterus

oceurs—not of later segments where it might have been broken

up into detached cavities. Moreover, a nearer inspection of some

Text-figure 2.

Linstowia ameive.

Portion of a section through a ripe proglottid showing disposition of ova.

e. Ova lying in parenchyma. e.m. Ova lying in definite uterine spaces.

v.g. Remains of vitelline gland. w.v. Water-vascular tubes.

of the egeg-containing cavities shows that they are traversed by

delicate strands of tissue—that, in fact, they are not really all of

them cavities, but are merely looser regions of the delicate

network which chiefly constitutes the medullary parenchyma.

Furthermore, plenty of eggs are placed singly between the

meshes of the parenchyma enclosed in no special cavities and even

lying just outside of such cavities as have been described above.

Some of these eggs have already formed embryos; but perhaps

the majority are still in the unicellular condition. These eggs,

whether lying singly in the parenchyma or a few together in

cavities, extend over much of the proglottid and are quite in

268 DR. F. E. BEDDARD ON

contact with the anteriorly situated ovary. It is impossible to

say where ovary ends and “ uterus” begins. There is nothing in

these facts to forbid the assumption that the eggs leave the ovary

and migrate directly into the parenchyma, not reaching it wid

a uterus.

Towards the end of the body the uterine cavities, if they be

such, have completely disappeared, and the embryos (text-fig. 3)

are more or less evenly scattered through the parenchyma. They

even get to be found in the cortical parenchyma, though by no

means numerously. It is not infrequent among tapeworms for

Text-figure 3.

Linstowia ameive.

Portion of a section through a more fully ripe proglottid than that represented

in text-fig. 2, and also more highly magnified.

A, Outermost membrane of the embryo. B. Middle membrane.

C. Embryo surrounded by delicate innermost membranes.

the ripe embryos to lie also in the cortical parenchyma; and

in an ally of the present species, viz. Linstowia brasiliensis, the

eggs stray thither*. The ripe embryos of the present species

are hexacanth, as is usual; each appears to be wrapped in three

shells—unless the outermost membrane, which is more stainable

by hematoxylin, be regarded as belonging to the maternal tissues.

The spaces of the parenchyma, in one of which each embryo hes,

fit the shells fairly accurately, and show no signs of being

independent cavities lined by an epithelium.

* y. Janicki, Zeitschr. wiss. Zool. lxxxi. 1906, Taf. xx. fig. 2.

TWO NEW SPECIES OF CESTODES. 269

The principal features in the anatomy of this worm may be

thus summed up :—

Linstowia ameive, sp. n.

Length up to 25 mm.; greatest diameter 15 mm. Rostellum

absent ; four suckers unarmed. A neck present ; posterior segments

increased in length, but not longer than wide. Genital pores

alternate in position, close to anterior margin of segment. Cortical

layer thick, about the same diameter as medullary layer; longi-

tudinal muscles in two layers, innermost of small bundles of fibres

not more than 8 or 10 to a bundle, and often less, outermost layers

of fibres implanted singly or in twos. Water-vascular tubes lie side

by side, the smaller dorsal tube being external; there is also a

network of excretory tubes. The testes lie posteriorly to the ovary and

vitelline gland, and extend to the dorsal surface, but do not overlap

female gonads; cirrus-sac moderately large with muscular walls

opening into genital cloaca in front of vagina, cirrus without

spines; sperm-duct coiled, without vesicula seminalis. Ovary

single with lateral wings, lying in front of segment. Vitelline

gland, not so extensive, lies behind. Vagina without dilated recep-

taculum seminis. LHggs lie at first partly within scattered cavities

in the parenchyma and partly between the meshes of the same; later

no cavities are to be seen and the ova are imbedded singly in the

parenchyma ; the eggs are surrounded by three shells.

Hab. Ameiva surinamensis.

OOCHORISTICA MARMOS.£, sp. 0.

Of this new species I have been able to examine but a single

specimen, which was obtained from an American Marsupial,

Marmosa elegans. The specimen was not all in one piece; but,

if the pieces were all of one individual, the length is 84 mm.

Otherwise the length of the largest piece, which included the

scolex, was 54mm. The greatest breadth of a ripe proglottid is

slightly under three millimetres. During life the posterior

segments were extended to a length of rather more than twice

their width. They were quite retracted by alcohol and became

shorter than broad. The unarmed head has the usual four

suckers, which are directed upwards. There is thus nothing

distinctive in the external characters of this species. ‘The genera-

tive pores are not visible, except by the section method; for they

open anteriorly in each proglottid and their orifices are covered

by an overlap of the proglottid in front. These pores alternate

irregularly from side to side of the body as in all other species of

Oochoristica.

In transverse sections through this tapeworm it may be seen

that it agrees with Oochoristica, as opposed to Linstowia, in the

comparative thinness of the cortical layer, which is less in

diameter than the medullary layer. The muscular layers in the

cortex have a characteristic arrangement (text-fig. 4, A), which

270 DR. F. E. BEDDARD ON

- Text-figure 4.

A series of transverse sections through the cortex of various species of Oochoristica

and Linstowia, to illustrate the arrangement of the longitudinal musculature.

A. Oochoristica marmose, B. Linstowia ameive. C. L. echidne or semoni.

Z D. Oochoristica sp.

TWO NEW SPECIES OF CESTODES. 271

may be compared, in the figure cited, with those of other and

allied species (text-fig. 4, B,C, D). In the present species a

delicate layer of transverse fibres forms the innermost layer of

the cortex and separates it off from the medulla. Immediately

outside of this is a layer of longitudinal fibres disposed in stoutish

bundles separated from each other by considerable intervals,

Outside of these again are scattered longitudinal fibres, which are

here and there aggregated into small bundles of two or three

fibres. The large internally situated bundles consist of 15-20

fibres apiece.

Text-fig. 4, B, illustrates a corresponding section through

Linstowia amewee, which I have already described *, and which

shows plain differences from Oochoristica nuarmose.

For the purpose of comparison with these two species and to

show the value as specific marks of the arrangement of the longi-

tudinal muscles, I subjoin corresponding figures of a Linstowia

from Hehidna (text-fig. 4, C) which may be ZL. echidne or L.

semont (I have no means of deciding the point), and of Oochoris-

tica (text-fig. 4, D), which is near to and possibly identical with

O. wageneri, whose general anatomy has been described by myself

lately r. In the former species (text-fig. 4, C) the longitudinal

muscular layer is divisible, as in the other species, into two sheets,

of which, as before, the innermost is the stronger. So much so,

however, that the outer muscular sheath is reduced to a very few

fibres. ‘The inner stronger layer is not by any means so well

developed as in the last two species: the bundles are smaller,

that is, they contain fewer fibres, and they are not by any means

so distinctly marked off from one another as in Oochoristica

marmose; they resemble more Linstowia ameive, a point of

importance in view of possible generic identity.

The last species represented in the figure (text-fig, 4, D) is

quite unlike any of the others, in that there is not a definite two-

layered disposition of the muscular fibres. There is simply one

irregular layer of bundles of varying sizes, that is, containing a

variable number of muscular fibres, which together occupy a good

deal of the space which les between the subcuticular layer and

the transverse muscular layer. It will be observed, however,

that, on the whole, the larger bundles lie to the deeper side of the

cortical layer, those more superficially placed being smaller. On

the whole, the arrangement of the longitudinal muscles in this

species is more like that of Oochoristica marmose than of the

other two species figured, by virtue of the size of the bundles.

Butif we attempt to draw generic definitions from this character,

it might perhaps be urged with equal force that the reduction of

the muscles in Linstowia semoni sets that species apart from all

the rest. In any case, it cannot be doubted that these muscles

furnish very clear specific characters which have not been hitherto

sufficiently represented in figures illustrating these two genera.

* Supra, p. 264.

iy eZee Se LOM 627.

272, DR. F. E. BEDDARD ON

In both transverse and longitudinal sections only two large

longitudinally running trunks of the water-vascular system are

to be seen. These two trunks, as is so often the case, are of quite

different dimensions. The dorsal tube, which is very much

smaller than the ventral, lies above it, sometimes also rather to

the inside, and has thicker walls. This tube is so fine that I have

occasionally been unable to find it in a given section, though it

would appear to be quite a continuous vessel. Below hes the

very much larger ventral tube. Thisis situated at some distance

to the inside of the nerve-cord. In the existence of but two

principal longitudinal water-vascular tubes the present species

agrees with the other two forms which are found in South

American Marsupials, and also with the recently described species

named by Zschokke Oochoristica rostellata *; in the latter case,

however, the two tubes, dorsal and ventral, are of about the same

calibre, as is shown in his figure of a transverse section through

a proglottid of that worm. In longitudinal sections the same two

tubes are always quite visible.

To these two principal longitudinal tubes on each side of the

proglottid a third may be added, which is, however, not comparable

with the additional tubes found in certain species of Oochoristica

from 8S. American Edentates. One does not find in transverse

sections six conspicuous longitudinal vessels such as are so plain

in an Oochoristica from T'amandua, upon which I have recently

reported to the Society. But, on the other hand, the anastomosis

between the branches of the excretory network which pervades

the segments does lead to the formation of at least short longi-

tudinal trunks. I have observed such a one lying outside of the

nerve-cord. The branches of the water-vascular system in this

species are indeed very copiously developed. Four or five, or

even more, large branches from the ventral vessels pass across

the proglottids, and even more are directed towards the lateral

margin passing across the nerve-cord. The whole body is, in fact,

richly supplied by a plexus, of which the individual tubes are

often wide. I have not been able to ascertain whether any of

these branches perforate the cortex and reach the exterior of the

body.

§ Male Gonads and Ducts.

The relative positions of the male and female gonads in this

group are frequently made use of as a generic distinction. The

present genus, Oochoristica, has been partly defined thus by

Ransom + :—“ Testicles numerous, surround female glands pos-

teriorly and on the sides.” The figure given of O. rostellata by

Zschokke t is in accord with this definition; and so are certain

of the figures given by v. Janicki§. Not, however, his illus-

trations of the species O. bivittata and O. didelphydis, which are

* Zeitscbr. wiss. Zool. Bd. Ixxxiii. 1905, pl. i. fig. 2.

+ Bull. U.S. Nat. Mus. No. 69, 1909, p. 85.

Zeitschr. wiss. Zool. Bd. Ixxxiii. 1905, pl. 1. fig. 3.

§ Tbhid. Ba. lxxxi. 1906, pl. xx. figs. 5, 7, pl. xxi. figs. 18, 21.

TWO NEW SPECIES OF CESTODES. 273

for certain reasons (geographical distribution and host) to be

compared particularly with that which forms the subject of the

present communication. In these two species the testes are

entirely posterior to the female gonads. It seems possible that

some differences are to be accounted for by the state of contraction

of the proglottids.

In Oochoristica marmose, in those proglottids where they

are at the height of their development, the testes are mainly

to be found at the sides of the centrally placed ovary and vitelline

gland, which are themselves very large and occupy most of the

central region of the proglottid, both dorso-ventrally and antero-

posteriorly. The testes extend above the female gonads dorsally,

and there is a single row of them posteriorly behind the vitelline

gland—in fact, they occupy pretty well all of the available space

within the proglottid. They are numerous—TI have counted

as many as 50 in a single horizontal section. Inasmuch as a

single proglottid cut horizontally was displayed in 25 sections

(not including sections through the cortex), the number of testes

is obviously large, even though the first and last sections only

show one or two testes. The largest number given by Zschokke

in his review of the genus is 100 for O. rostellata. My species

must considerably exceed that.

The vas deferens opens through the cirrus-sae into a genital

cloaca, The genital cloaca is less conspicuous in this Oochoristica

than it is in some other tapeworms. It is in depth about one-

third of the length of the cirrus-sac, which is itself small, and

has no marked specialization into regions such as occurs, for

example, in Hugonodeum adicnemni*. It is narrow and tubular

in horizontal section, and the cirrus-sac opens into it at its internal

extremity. It is surrounded by muscle-fibres, which doubtless

act as a sphincter. ‘This orifice is in front of that of the vagina.

The cirrus-sac is small and only extends back as far as the nerve-

cord. It is as usual bottle-shaped, being wider posteriorly, where

also the muscular wall is much thinner. I could discover no

spines upon the contained cirrus, which widens out and becomes

very muscular at its outer end. Altogether there is nothing

remarkable about the cirrus-sac and cirrus of this Oochoristice.

But its shape and size and the muscularity of its walls are

important specific marks—for they differentiate it from

O. rostellata.

The vas deferens issues from the cirrus-sac as a straight tube

running parallel with the vagina. Later it forms a loose and

not very extensive coil, and often bends backwards and breaks

up into a number of branches, which again become subdivided

to supply the individual testes. This region of the sperm-duct

is of the nature of a network, for anastomoses exist between

many of the branches. The walls of the finer branches of the

sperm-duct and their communication are quite visible, even

* P.Z.S. 1913, p. 866, text-fig, 144.

274 DR. F. E. BEDDARD ON

under only moderately high powers; and when gorged with

sperm they are not any more conspicuous.

§ Female Gonads and Ducts.

The ovary lies anteriorly in the proglottid, but is prevented

from reaching the actual limit of the proglottid by the transverse

uterus which lies in front of it. It consists of a central portion,

which lies practically in the middle of the proglottid, and of

two lateral wings. ‘The latter are frayed out into numerous

thick digitiform processes, which extend rather beyond the

range of the posteriorly situated vitelline gland. In sections cut

horizontally the digitiform outgrowths of the ovary have the

appearance of circular or oval sacs filled with ova. They present

the very closest resemblance to the sacs of the uterus which le

among them. The vitelline gland is rather smaller than the

ovary, behind which it lies. It is very much of the same form,

being prolonged laterally into blunt processes. The vagina runs

at first a straight or slightly sinuous course behind the vas

deferens; it then bends backwards and becomes dilated to form

a cylindrical receptaculuim seminis, which lies obliquely at an

angle of about 45° with the longitudinal axis of the worm. The

lumen of that part of the vagina which is nearest to the

receptaculum is narrower than that of the outer section of the

vagina; I have observed cilia lining the vagina at a considerable

distance from the external orifice. The receptaculum seminis 18

generally full of sperm. In many cases I have found numerous

ova within it. I have not observed autocopulation in this

species.

§ The Uterus and the embedding of Ova in the Parenchyma.

In the genus Oochoristica the uterus 1s not a prominent feature—

“Die rasche Entwicklung und der ebenso prompte Zerfall des

urspriinglich ventral angelegten Uterus characterisiert, soweil

genauere Untersuchungen ein Urteil erlauben, das ganze

Genus Oochoristica,’ writes Zschokke * in a general survey of

the characters of this genus. No precise figures are given‘as

to the duration of the uterus by Zschokke in the species

(O. rostellata) described in the memoir from which the above

quotation has been made. I find myself that, in the species which

I deseribe in the present paper, the uterus 1s quite obvious in

34 segments, a space of about 10 mm. in length, which is a very

appreciable portion of the entire body-length.

The uterus is seen on a general examination to consist of

numerous closely adpressed circular to oval chambers, which are

more or less filled with eggs. This system of cavities is at first

mainly ventral in position. In a series of horizontal sections

the ventralmost show only uterine cavities, which extend up to

* Zeitschr. wiss. Zool. Ixxxiii. 1905, p. 63.

TWO NEW SPECIES OF CESTODES. 275

the anterior, but not up to the posterior, margin of the segment.

In fact, they completely underlie the ovary, but not always

completely the vitelline gland. In the earliest proglottids which

show a uterus, the latter lies only anteriorly and, of course,

ventrally. The uterus in the first two or three segments in

which it is found consists merely of a transverse tube anteriorly

and ventrally, the two ends of which are dilated into two or

three more or less oval diverticula, which are thus quite lateral

in position. It seems, in fact, to resemble closely the uterus of

Oochoristica rostellata, as represented in Zschokke’s figure *.

Text-figure 5,

ut, ue,

Part of a horizontal section through three consecutive segments of Oochoristica

marmose, to show the extension of the uterus lateral] y-

1. Longitudinal muscles. m. Nerve-cord. wt. Uterus.

In later proglottids (text-fig. 5) the uterine cavities range

through the medulla much more extensively, and also penetrate

into the cortical layer. Finally, the cavities of the uterus are

obliterated by growths of the parenchyma, and the individual eggs

(or rather embryos) come to lie singly, filling up the entire interior

of the proglottid. The development of the uterus, in fact, seems

* Zeitschr. wiss. Zool. Bd. Ixxxiii. 1905, Taf. i. fig. 2, v.

Proc. Zoot. Soc.—1914, No. XIX. 19

276 DR. F. E. BEDDARD ON

to be much as in O. rostellata, though Zschokke does not mention

the invasion of the cortical layer by uterine spaces. It is to be

noted, however, that in my species the extension of the embryos

into the cortical layer mainly takes place laterally, and not

dorsally or ventrally to any great extent, though here and there

I have observed embryos in these regions of the cortical layer.

Text-figure 6.

ui ‘ ts tie me

ieee ee cae oi

e8 %.

SOR

2 oF

Half of a transverse section through a nearly completely mature proglottid of

Oochoristica marmose, showing the embryos scattered through the parenchyma.

m2. Nerve-cord. w.v. Water-vascular tubes.

Particularly is the latter the case with the quite mature

proglottids, where the embryos are surrounded by their shells.

We may note before proceeding further with the description of

the embryos the differences which the uterus shows in the two

species dealt with in the present paper. In Oochoristica marmosce

the uterus consists of a well-marked posteriorly situated transverse

TWO NEW SPECIES OF CESTODES. 277

tube, which gives off ovai diverticula laterally. These cavities

have well-marked walls, which, however, do not seem to be

independent of the surrounding parenchyma; they have no

special lining of their own. Later the cavities are multiplied,

and the eggs from the very first are confined to these cavities

and never lie in the parenchyma between them. The uterus,

moreover, exists throughout a good many proglottids. On the

other hand, in Linstowia ameive the uterine cavities are less

strongly marked off and altogether less definite ; they do not

form a coherent group and exist for a much shorter period.

Moreover, from the very first the ripe ova do not all of them lie

within these cavities ; they are continually to be found imbedded

in the parenchyma between the uterine cavities. The uterus

seems to be degenerating in this species as compared with that

of O. marmose.

Text-figure 7.

A portion of the same mere highly maguiuce.

The membranes surrounding the embryo (e.) are not yet formed.

The disappearance of the uterus in Oochoristica marmose. is

followed by a stage (illustrated in text-fig. 6) in which the eggs,

which have by this time developed into embryos, are uniformly

scattered through the medulla (occasionally invading the cortical

layer, as already mentioned) for a considerable number of pro-

glottids, which I am unable to state exactly; the scattered

embryos possess no definite shell. They lie (text-fig. 7) in

cavities of uniform size separated by meshwork from each other.

These cavities resemble in every way the cavities of the paren-

chyma in various tapeworms, where there is no question of

uterine cavities. It must be admitted, however, that such a

space as there is round an individual embryo is to be looked

278 DR. F. E. BEDDARD ON

upon as the remains of the uterine cavity. Those who have

figured the embryo of Oochoristica have, as a rule, not repre-

sented this stage, which, however, is not the case with Lihe™*,

who has represented such embryos in “ Tenia” megastoma Dies.

The first membrane to be formed is a fine one immediately

surrounding the embryo. Later on a much thicker and deeply

staining outer membrane is formed, which lines the “ capsule”

of the parenchyma in which the embryo lies. Between these

two there is no third membrane. There is thus an important

difference between the present species and Linstowia ameivee (cf.

text-figs. 3, 8), also described in the present communication to

the Society. In Oochoristica marmose (text-fig. 8) the eggs

Text-figure 8.

A later stage than that represented in text-fig. 7.

A. Outer membrane. C. Embryo surrounded by thin inner membrane.

have therefore a much clearer appearance, owing to the larger

empty space which separates the embryo from the outermost

membrane. As to this latter membrane, it is apt to be

hexagonal in transverse section, owing to its separation from

the embryo and close adherence to the parenchymal walls; and

it is not unreasonable to think that it may be a product of the

parenchyma rather than of the embryonic cells. This appears

to be the opinion of Zschokke 7 with regard to the uterine

ova of Linstowia semoni, for he writes: ‘“* Die ganze Markschicht

ist .....vollstiindig angefillt von derbwandigen, rundlichen

Bindegewebskapseln, die sich gegenseitig einengen. Jede

* Arch. f. Natutg. 1895, pl. x1. fig. 15.

+ “Die Cestoden der Marsupialia, etc.,” in Semon’s Zool. Forschungsreise, etc.,

Jan. 1898, p. 368.

TWO NEW SPECIES OF CESTODES. 279

Kapsel beherbergt in der Regel ein einziges Hi.” ‘The structure

figured by Zschokke *, and to which he refers in the above quota-

tion, seems to be of the nature of a membrane and to be therefore

quite like the membrane to which I here refer in Oochoristica

marmose. It does not suggest a cellular layer such as I have

figured in Oochoristica sp. As to the number of shells, whether

thrown off by the embryo or not, which surround the embryo

in the genus Oochoristica, there appear to be differences among

different species. Cohn distinctly represents three shells £ in

O. surinamensis, while Marotel § asserts the presence of only

two in the Oochoristica of the European Badger, O. incisa of

Raillet ||.

§ Affinities of the Species.

I have described this species as an Oochoristica; into the

question of the definition of this genus I propose to enter later.

In the meantime, it is necessary to enquire whether---apart

altogether from the question of genus—it may not be specifically

identical with Linstowia brasiliensis ™ from Didelphys tristriata.

There is, I think, no possibility of confusing the two species, in

spite of many points of general resemblance. In the first place,

the simplicity of the water-vascular system of Dr. v. Janicki’s

Species distinguishes it from mine. Furthermore, the fact that

in Linstowia brasiliensis the ripe eggs of the posterior segments

are limite 1 to the lateral areas of the segments militates against

the identity of the two species now under consideration. ‘These

differences appear to me to be suflicient, without going into a

more detailed comparison between Linstowia brasiliensis and

Oochoristica marmose. The same remarks apply to L. theringi**.

Oochoristica didelphydis is too small a species (it measures only

15 mm. in length) to be confused with the present one. Further-

more, the scolex was absent, which increases the difficulties of

identification. But it agrees with my species in having only

two lateral vessels, of which the dorsal has a very fine lumen. On

the other hand, the transverse tube is quite simple, which is not

the case with that of O. marmose. The obliquely directed

cirrus-sac of O. didelphydis contrasts with the perfectly straight

one of O. marmose. The fact that in the former species

the vagina opens on to the exterior in front of the cirrus-sac

appears to me to be such an important difference, not only

from my species, but also from the species of the genus

Oochoristica, that it leads me to doubt the generic identity of these

forms. And I would again point out that while the vagina

of O. marmose is perfectly straight, that of O. didelphydis

is much coiled. This appears to me to be quite a salient

* Loe. cit. pl. xxiv. fig. 7. + P.Z.S. 1913, p. 875, text-fig. 149.

t Arch. f. Naturg. 1903, p. 65, fig. 9. § C.R. Soc. Biol. 1899, p. 21.

|| Loe. cit. t. cit. p. 23.

| v. Janicki, Zeitschr. wiss. Zool. 1xxxi. p. 507.

** ZLschokke, C.Bl. Parasit. xxxvi. 1904, p. 51.

280 DR. F. E. BEDDARD ON

difference, and adds to the possibility of generic difference.

Indeed, v. Janicki admits the uncertainty of the inclusion of

this species within the genus Oochoristica. No information is

given of the ripe ova; but a small circular space lying

in front of the ovary and marked in his fig. 5 with a ‘“‘?” is

possibly to be regarded as the uterus. In any case, quite apart

from the generic identity of ‘‘ Oochoristica” didelphydis, there

can be no possible comparison between this species and

Oochoristica marmose. The points of difference are too many

and too important.

In comparing Oochoristica marmose with other species of the

genus, habitat comes first into consideration and then the

systematic position of the host. There are only two species of

Oochoristica from South American Marsupials known at the

present time; and these are O. bivittata and O. didelphydis

recently described by v. Janicki*. 0. bivittata is a much more

slender worm than the one described here by myself ; the greatest

diameter is only °9 mm. It agrees generally in the unarmed

scolex, in the fact that the vagina opens behind the cirrus-sac,

and in the imbedding of the ripe eggs singly in the parenchyma,

and in smaller details, which are, nevertheless, of systematic

importance. There is, in fact, no doubt as to the generic identity

of the two worms. The differences, however, are quite of

specific value. The complication of the excretory system

of O. marmose contrasts with the simple transverse vessels of

O. bivittata. It is to be noted, however, that in all three species

from Didelphys (the genus Marmosa but slightly differs from

Didelphys) there are only two main longitudinal trunks—a point

of similarity which is of interest. The generative organs are

quite unlike in the two species in a number of features. The

position of the gonads contrasts, and the testes are very few in

number, in O. bivittata. This may, however, be partly due to the

maturity of the segments, in one of which they are figured as not

exceeding five in number clustered together behind the vitelline

glands. Finally, the ripe eggs, though scattered singly as in

other species of Oochoristica, are limited to, the sides of the

proglottids in O. bivittata.

I conclude with a definition of the new species, which is as

follows :—

Oochoristica marmose, sp. n.

Length at least 54 mm.; diameter 2°38 mm. Scolex unarmed,

with suckers directed forwards, genital pores anteriorly situated

in proglottid, alternate. Cortex not so wide as medulla. Longi-

tudinal muscles forming a layer of bundles consisting of 10

or more fibres, above which are scattered fibres. Water-vascular

system consisting of two longitudinal vessels on each side connected

by a rich plexus of rather large branches. Testes very numerous,

* “Studien an Satigethiercestoden,’ Zeitschr. wiss. Zool. Ixxxi. 1906, p. 505 ;

and a preliminary account in Zool. Anz. xxvii. 1904, p. 770.

TWO NEW SPECIES OF CESTODES. 281

filling all the available space in the proglottid; cirrus-sac small,

opening into a genital cloaca in front of vagina; sperm-duct with

a loose coil dividing up into a meshwork of ductules. Ovary anterior

in position ; vagina with cylindrical receptaculum ovorum. Uterus

consists of a posteriorly situated and transverse tube, from which

arise numerous diverticula, extending ultimately through most of

proglottid ; uterus finally disappears, leaving embryos scattered

singly throughout parenchyma, extending here and -there into

cortec. Hmbryos with two shells.

Hab. American Marsupial, Marmosa elegans.

S$ The Genera Oochoristica and Linstowia.

The fact that Linstowia and Oochoristica are placed in separate

families by systematists at present has tended of itself to

exaggerate the differences which exist between these genera.

Their real propinquity, however, becomes very apparent when

we wise the “ Key to Genera” devised by Mr. B. H. Ransom *.

In this dichotomous table we pursue the two genera side by side

until the very last of the characters made use of ; they are in

this differentiated by the thickness of the cortical layer and the

position of the testes in the proglottid. It should be noted that

this dichotomous table and the subsequent generic definitions f

given by Ransom were published after the information gathered

by Zschokke and v. Janicki had been put forward.

When we come to look into the characters of the known

species, including those deseribed in the present communication,

that have been referred respectively to one or other of the

genera Linstowia and Oochoristica, it does not appear to be at all

an easy matter to draw many hard and fast lines of separation.

The first-described species of Linstowia, not then referred beyond

Tenia by its original describer {, has been investigated in further

detail by Zschokke§. The salient characters of this worm are

the following :—The cortex is thick ||; the longitudinal muscle-

bundles are in two rows, of which the inner is the thicker, but

the bundles have not more than four or five fibres each; the

genital cloaca is deep; the cirrus-sac is very large; the vas

deferens is coiled and dilates posteriorly, just before it begins

to divide, into a kind of vesicula seminalis; the testes extend

through the proglottid dorsally ; the vitelline gland is quite.

posterior in position and nothing lies behind it. The excretory

tubes he side by side, and are formed by a transverse vessel in

each segment as well as by a network.

Zschokke gives in the same work a fuller account of a second

species, viz. Linstowia semoni, an example of which serves to

fill up certain lacune in the generic characters. Of this worm

* Bull. U.S. Nat. Mus. No. 69. 1909, p. 53, &c. + Loc. cit. pp. 65 & 84.

+ D’Arcy Thompson, Journ. Roy. Micr. Soc. 1903.

§ Zschokke in Semon’s Forschungsreise, etc., 1898, pl. xiv. figs. 1 & 2.

|| Thompson, Zoe. cit. pl. v. fig.8. A transverse section is not figured by Zschokke.

282 DR. F. E. BEDDARD ON

there is no transverse section figured to show the relative thick-

ness of the cortex. The water-vascular system, not figured in

L. echidne, has as main trunks two tubes on each side, which

later lie parallel, and not one above the other, which is the case

anteriorly ; of these the dorsal tube lies to the outer side of the

ventral. There is also a transverse trunk to be seen in each

segment. The vagina in this as well as in the last species

possesses a dilatation usually termed the receptaculum seminis.

The uterus “ forms a thin-walled folded tube where the wall is

early lost” *, and the eggs come to be imbedded singly in the

parenchyma. ‘To these descriptions of the two species it should

be added that in both the scolex is unarmed, that the genital pores

alternate, and that there is nothing remarkable in the structure

of other organs which have not been mentioned.

If we contrast with these species certain forms which have

been referred to the genus Oochoristica from Kdentates rather

marked differences at once appear. In such formsas O. wageneri

and the allied form which I have myself + lately described also

from the Lesser Anteater, which may or may not be identical with

it, we find the following assemblage of characters. While they

agree with the members of the genus Linstowia, already referred

to, in the unarmed scolex, the alternate generative pores, and the

imbedding of the ripe eggs singly in the parenchyma, they differ

by the much more complex water-vascular system consisting of

six longitudinal tubes; they have also a small cirrus-sac which

does not extend far into the body, not reaching much if anything

beyond the nerve-cord. If these two groups of species comprised

all that were known, the separation of the two genera would be

quite easy and obvious. But there are forms which render such

a demarcation impossible.

In O. rostellata of Zschokke £ there are but two water-vascular

vessels ; though these vessels are superposed instead of lying side

by side as in Linstowia. On the other hand, in Linstowia theringi

and Z. brasiliensis the cirrus-sac is as small as in Oochoristica !

While in the species described in the present paper as Linstowia

ameive, the cirrus-sac is large (as in the Australian members of

the genus Linstowia), and the testes are as markedly behind

the ovaries as in Janicki’s species Oochoristica bivittata. As for

differences in the relative thickness of the cortex and medulla

insisted upon by many, I can see no difference worth mentioning

in the figures of Oochoristica wagenert and Linstowia brasiliensis

given by v. Janicki§.

It is thus next to impossible to separate the genera if we

accept the present distribution of species among them. Nor is

the matter ameliorated if we make the planes of division some-

what different. It had occurred to me to separate off the

* For the species of Australian Linstowia, see also Zschokke in Zeitschr. f. wiss.

Zool. Bd. lxv. (1899).

+ P.Z.S. 1912, p. 627. t Zeitschr. wiss. Zool. Bd. Ixv.

§ Zeitschr. wiss. Zool. Bd. lxxxi, text-fig. 4, p. 534, & pl. xx. fig. 2

TWO NEW SPECIES OF CESTODES. 283

Australian forms limited to that country and to the two groups

Marsupials and Monotremes. Here geographical range and

the systematic position of the host concur with unusually large

cirrus-sac as an anatomical character. But one of the two

species, viz. Linstowia semoni, has a distinctly larger cirrus-sac

than LZ. echidne ; and in my species LZ. ameive the cirrus-

sac is not much smaller than that of LZ. echidne. One structural

feature occurs to me as being of possible use in better defining

the two genera Linstowia and Oochorisiica. But it is so little

known that it cannot be used for the present and may after all

turn out to be worthless. This concerns the imbedding of the

ova in the parenchyma after the disappearance of the uterus.

I have pointed out *, in describing the ripe eggs of a species of

Oochoristica, that they are encircled by a cellular layer suggestive

of a commencing paruterine organ like that of Davainea, etc.

If it be found that this character also signalizes other South

American species from Hdentates a separation might well be

made. Furthermore, it is quite possible that the condition of

the uterus may serve as a dividing-line, as it certainly appears to

do in the case of two other mutually related genera, viz.

Inermicapsifer and Zschokkeellat. The kind of difference that

is meant by this suggestion is that shown by the two species

described in the present paper, and has been put forward in

detail above, accompanied by illustrations (text-figs. 2 & 5).

In the meantime, it does not seem possible to form a reasonable

definition of the two genera, and [ am strongly of opinion that

there are no grounds at all for placing Oochoristica and Linstowia

in separate families.

* In a paper upon a new genus Wugonodeum in P.Z.8. 1913, p. 875, text-fig. 149.

+ See P. Z.S. 1912, p. 607.

co ed

Hate y bos

ON THE MALAY RACE OF THE INDIAN ELEPHANT. 285

18. The Malay Race of the Indian Elephant, Hlephas

maaimus hirsutus*. By R. LypeKKer, F.R.S., F.Z.S.t

[Received February 24,1914: Read March 17, 1914.1]

(Text-figures 1-3.)

INDEX. Page

Description of the new race .............020.....00806.0..-..... 288

Thanks to the Trustees of the estate of the late Mr. Rowland

Ward, the Natural History Branch of the British Museum has

received the mounted skin of the young Malay Hlephant which

died in the Society’s Gardens during the latter part of last year.

While yet alive, the extraordinary hairiness of this animal

attracted the attention of naturalists; and this feature, coupled

with a pecuharity in the form of the ears, seems so well marked

and so distinctive as to justify the recognition of the Malay

Elephant as a distinct local race of the Asiatic species.

Text-figure 1.

Young Malay Elephant when alive in the Society’s Gardens.

As I learn from Mr. Pocock, this Elephant came from the

Kuala Pila district of the Negri Sembilan province of the Malay

* [The complete account of this new subspecies appears here, but since the name

and a preliminary diagnosis were published in the ‘ Abstract,’ No. 130, 1914, it is

distinguished by being underlined.—Ep1rTor. |

+ Published by permission of the Trustees of the British Museum.

286 MR. R. LYDEKKER ON THE

Peninsula, and is believed to have been about three years old at

the time of its death. As shown in the accompanying photograph

from life (text-fig. 1), it has a somewhat stunted appearance—

suggestive, at first sight, of its belonging to a small race,—the

height of the specimen, as mounted, being about 3 feet 8 inches.

This stunted appearance may, however, be merely due to the

effects of early captivity, for Mr. T. R. Hubback, in his book

on Elephant and Seladang Hunting in the Federated Malay

States *, records that the Elephants of the Negri Sembilan are of

ordinary size, although of late years most of the big bulls have

been killed off. He also mentions that practically all the bulls

are tuskers, and very generally have one tusk much smaller than

the other.

Before proceeding further, it may be mentioned that it is very

difficult to obtain definite information with regard to the hairiness

ov otherwise of Indian Elephant calves—that is to say, in those of

the typical Indian race of the species. Dr. Mobius f has, indeed,

Text-figure 2.

Newly born indian Klephant im the Natural History Museum.

shown that young Elephants may have remnants of a soft hairy

eoat comparable to the under-fur of the Mammoth, and the

young Elephant born—from a presumably Indian mother— in the

* London, 1905, p. 63.

+ Sitzber. k.-preuss. Ak. Wiss., Math.-Nat. 1892, p. 527

MALAY RACE OF THE INDIAN ELEPHANT. 287

Society’s Gardens in 1902, the mounted skin of which (text-fig. 2)

is exhibited in the Natural History Museum, has a considerable

portion of the body clothed with a somewhat sparse coat of rather

short, soft, rufous hair, which also forms a fringe to the margins

of the ears. ‘There likewise appears to be a certain amount of

hair of a similar type shown in the photograph of the young

Indian Elephant born in Copenhagen in 1912.

On the other hand, from the fact that writers like G. P.

Sanderson and W. T. Blanford make no mention of the hairi-

ness of Indian Elephant calves, it seems probable that in many

cases the skin may be more or less nearly bare, although it must

be confessed that information with regard to this matter appears

to be very scant and defective. Still, it may be taken for granted

that if Indian-born Elephant calves exhibited hairiness in any

way comparable to that of the young Negri Sembilan animal, the

fact would have been recorded in textbooks on Indian natural

history and sport.

Text-figure 3.

Outline of a Foetus of a Siamese Elephant.

(From Toldt, Denks. k. Ak. Wiss. Wien, 1913.)

It is, however, very noteworthy that Mr. K. Toldt* has

recently described and figured an advanced fceetus of a Siamese

Elephant (text-fig. 3), in which the sides of the crown of the head,

the terminal half of the trunk, the point of the lower lip, the

under-parts, the greater portion of the limbs, and the hind aspect

of the buttocks are sparsely covered with short bristly hairs, which

would doubtless have attained much greater development after

birth. Nor is this all, for if Mr. Toldt’s figure (text-fig. 3) be

compared with that of the young Indian t Elephant shown in

* Denks. k. Ak. Wiss. Wien, vol. xc. p. 259, 1913.

+ The term ‘* Indian” is here used strictu sensw.

288 ON THE MALAY RACE OF THE INDIAN ELEPHANT,

text-fig. 2, it will be at once evident that there is a marked

difference in the shape of the ears in the two specimens.

To deseribe this difference in words is very difficult, and it

is most easily apprehended by contrasting the figures. It may be

mentioned, however, that in the Indian Hlephant—both young

and adult—the ear is distinctly triangular, owing to the great

development of the descending lobe, and that its longer diameter

is vertical. In the Siamese calf, on the other hand, the descending

lobe is less elongated and the postero-inferior border placed less

obliquely, while there is a greater development of the postero-

superior border. In consequence of this—making due allowance

for the turning-forward of the lower part of the postero-superior

border, which may apparently also occur in Indian calves *,—the

whole lower half of the ear forms, roughly speaking, an obliquely

hung half-square, with somewhat emarginate borders.

Now the ear of the Negri Sembilan calf accords very closely in

general type with that of the Siamese fcetus, the chief differences

being the minor degree of emargination of the borders and the

direction of the antero-inferior border, which is inclined downwards

and backwards in a much more marked degree. The differences

are, however, slight and, coupled with the hairiness of both,

indicate that the two represent either two closely allied races or a

single and slightly variable race. As regards hairiness, the Negri

Sembilan calf has bristly black hairs on most or all of the regions

where these occur in the Siamese foetus; and, in addition, a large

amount of softer and longer black hair on the back and flanks, as

well as on the under surface of the lower jaw.

The tail is considerably longer than in the Indian calf, but

since, according to Sanderson, there is considerable variation in

this respect among Indian Elephants—which may or may not

eventually prove to be of racial value,-—I do not for the present

propose to take any account of this feature. The great difference

in the contour of the ear, coupled with the excessive development

(at least in some instances) of black and in part bristly hair in

the juvenile condition, seems sufficient to justify the separation of

the Elephant of the Malay Peninsula asa distinct race, under the

name of Hlephas maximus hirsutus t. That the Siamese Elephant

is near akin to this race seems certain, but whether it should be

included therein, or should be regarded as a race by itself, may

be left for future consideration.

It should be added that I have no means—except on the ground

of geographical isolation—of distinguishing the Malay Elephant

from HL. maximus swmatranus, the description of that race = not

being comparable with the specimens forming the subject of the

present communication.

* According to Sanderson, ‘ Thirteen Years among the Wild Beasts of India,’ p. 60,

the permanent backward folding of the margin of the ear in Indian Elephants does

not take place till the sixth or seventh year.

+ Abstract P. Z.S. 1914, p. 20 (March 24th).

+ See Sclater, Nat. Hist. Rev. vol. i. p. 72, 1862, and Falconer’s ‘ Paleontological

Memoirs,’ vol. i1. p. 256, 1868.

ON THE ONYCHOPHORA OF W. AUSTRALIA. 289

19. Fauna of Western Australia—I. The Onychophora of

Western Australia. By W. J. Daxiy, D.Sc., F.LS.,

F.Z.8., Professor of Biology, University of Western

Australia.

[Received January 12, 1914: Read March 17, 1914. ]

(Text-figure 1.)

INDEX.

Systematic : Page

Peripatus: Species Of .......ccceeecseesseteetee eee 289

Peripatoides gilesii=P. woodwardi .................. 290

There is a tendency for our knowledge of the species of Peri-

patus in Western Australia to become somewhat confused. Three

species have been described, namely :—

I. Peripatus leuckarti, var. occidentalis.

syn. Peripatoides occidentalis.

II. Peripatoides giles Spencer.

Ill. Peripatoides woodwardi Bouvier.

Of these three, Peripatoides occidentalis was the first to be

made known. ‘The species was described by Fletcher, from speci-

mens collected by a Mr. Lea at Bridgetown, South-Western

Australia (Proc. Linn. Soc. N.S.W. 1895 (2), x.). The diagnosis

reads as follows:—* P. lewckarti Sing, var. occidentalis, var. nov.

With 15 pairs of walking legs; outer jaw-blades without an

accessory tooth.”

Beyond a brief reference to the colour, no further description 1s

given and unfortunately no figures accompany the paper.

Twelve years later, Peripatws was found in another locality

(Armadale) in Western Australia, not far from the capital,

Perth, and specimens collected by Mr. H. M, Giles were sent to

Professor Baldwin Spencer. The latter considered them to belong

to a new species, and, in a short paper read September 1908 (but

not published until March 1909), named the species Peripatoides

gilesii after the collector *.

In the year 1905, however (that is two years before the speci-

mens were sent by Giles to Spencer), the German Expedition of

Michaelsen and Hartmeyer captured a number of Peripatus at

Lion Mill, a place also near Perth. These specimens were sent to

Bouvier, and his description with an account of the anatomy was

published in the Reports of the Expedition in 1909+. Bouvier

recognised these specimens as belonging to a new species, which

he named Peripatoides woodwardt.

It will be noted here that the appearance of the papers of both

* Proc. Royal Soc. Victoria, vol. xxi. (New Series), pt. ii. (1909).

+ ‘Die Fauna Siidwest-Australiens, Band 11. (Jena, 1908-09).

290 PROF. W. J. DAKIN ON THE

Spencer and Bouvier in the same year precluded either from

seeing the publication of the other.

After consulting both these papers, in order to name certain

specimens captured in almost the same district, the probability of

Peripatoides gilesti and Peripatoides woodwardi being one and the

same did not seem very remote. Further investigation has made

the probability an actual fact.

My first specimens, numbering 14 in all, were collected on the

afternoon of July 5th at a spot not far from Mundaring Weir.

They were obtained at distances varying from 3 to 4 yards to

80 yards from a small streamlet 2 or 3 feet in breadth and a few

inches deep, with banks 5 or 6 feet in height. The specimens

occurred lying under small logs, broken branches from the trees,

which were about 1 to 3 feet in length and 2 inches in diameter.

Occasionally they are to be found under larger logs. Further-

more, the animals were not distributed uniformly in the area

examined. They appear to occur on patches of ground the

character of which became familiar after a time, so that one could

speak of a “likely spot.” The ground was not particularly damp _

and was often in brilliant sunlight, there being little shade from

the scattered “blackboys” and eucalypts. In most cases the

surface of the ground was not covered with vegetation, but was

sandy or sand mixed with organic débris. Two, or even three,

specimens might be found under the same log, but usually only one.

These specimens from Mundaring Weir agreed with Bouvier’s

description of Peripatoides woodwardi. In order to be more

certain of their relation to Peripatoides gilesii, I asked Professor

Spencer for the type-specimens of the latter, and he very kindly

had these sent to me, together with a microscopic preparation of

the jaws.

Dimensions. According to Bouvier the length of his specimens

varied from 10°5 mm.to 21mm. This is stated by him to be

almost the size of Peripatoides orientalis and a little larger than

Peripatoides occidentalis.

Spencer's specimens of Peripatoides gilesti measured 22 mm.,

25 mm., and 27 mm., respectively. The discrepancy in size

means nothing, for in Bouvier’s case the individuals were preserved

and contracted. My specimens from Mundaring Weir ranged in

size from 12 mm. to 34 mm. The length of 34 mm. was that of

the largest when outstretched and walking. On being touched

it contracted to 28-30 mm., and on fixation it diminished still

further to about 22 mm. It is obvious therefore that one must

be careful in deducing any differences from dimensions.

Colour. So far as colour is concerned, the specimens collected

at Mundaring can be roughly sorted out into two groups—a group

in which brown-red predominates and a group in which green-

grey of a dull shade is the dominant colour. Spencer’s specimens

vary too in a similar way, and he refers in his paper to greenish-

yellow and reddish-brown specimens. Bouvier’s specimens are

again the same, and he described blue-green to black individuals

and others of a fawn-yellow, sometimes pale, sometimes red.

ONYCHOPHORA OF W. AUSTRALIA, 291

Spencer mentions further a broad lighter band just above the

level of the legs. Bouvier refers to a longitudinal band on each

flank above the bases of the legs, the colour of which is yellow-

red, and poor or even lacking in dark-coloured papille.

The Mandibles. The mandibles are accounted as one of the

chief diagnostic features of the species Peripatoides gilesit.

Text-figure 1.

Inner jaw-blades from the specimens of Peripatoides under consideration.

A. From type-specimens of P. gilesii Spencer.

B. Bouvier’s figure of jaw from P, woodwardi,

C & D. Jaws trom other individuals of same species.

According to Spencer the first jaw is simple in all specimens,

there being no accessory tooth. The second jaw has four clearly

marked and one minute accessory tooth. The above figure (text-

fig. 1) illustrates the jaw from Spencer’s specimen, two of the

jaws from my specimens, and the figure given by Bouvier for the

left mandible of Peripatoides woodwardi.

Proc. Zoot. Soc.—1914, No. XX. 20

292 ON THE ONYCHOPHORA OF W. AUSTRALIA.

Bouvier describes the jaw as follows :—

“‘ Les mandibules (fig. 3) sont dépourvues de dents accessoires

sur leur lame externe, comme dans les Peripatoides suteri, novee-

seylandic et occidentalis, et contrairement a ce que lon observe

dans le Peripatoides orientalis; leur iame interne présente 5 dents

ACCESSOILeS.”

The description and the figures (text-fig. 1, A, B) indicate that

there is no difference between the mandibles described by Spencer

and those deseribed by Bouvier.

There is, however, an interesting variation which seems to have

escaped both Spencer and Bouvier, probably owing to the ex-

amination of Jaws from only one individual. Having mislaid my

first preparation, which agreed with that of Spencer, I mounted

the jaws from a second individual. Jt was rather difficult in this

specimen to make out the fifth tooth, although it was present

(text-fig. 1,C). Wishing to procure a more definite example, I

removed the mandibles from a third individual, the largest one in

my collection. ‘To my surprise there were siz very definite teeth

on the inner lamella of the jaw (text-fig. 1, D). Thus the actual

number of accessory teeth given by Bouvier and Spencer may be

exceeded.

The Feet. 'The number of claw-bearing legsis perhaps the most

important character in the diagnosis of the species Peripatoides

gilesti and of Peripatoides woodwardi. Both Spencer and Bouvier

accentuate the number of legs. which in each case is said to dis-

tinguish their species from all other Australian forms except

Peripatoides sutert. In both species, however, the number is sixteen !

Spencer writes, “The number of legs together with the structure

of the jaws, serves to distinguish this species from all other

Australian species of either of the genera Peripatoides or Ooper-

inatus.”

Bouvier states that ‘‘ Le Peripatoides woodwardi resemble au

Peripatoides suterz et se distingue de tous les autres Peripatopsides

Australasiens par la présence constant de 16 paires de pattes.”

Thus, so far as the supposed characteristic features are con-

cerned, the specimens of Spencer and Bouvier are identical.

The inference is obvious.

There remains the question of priority in nomenclature.

Spencer’s paper is dated March 1909, although read in September

1908. Bouvier’s paper is dated 1909, but no month of issue is

given. In answer to a request for the exact date, Dr. Michaelsen

writes that the paper was publishedin December 1909. Spencer’s

description has therefore several months’ priority. There is, then,

only one species of Peripatus so far discovered in the region near

Perth, and that 1s Peripatoides gilesii Spencer. ‘The name Per-

putoides woodwardi must be rescinded.

It was hoped that specimens of Peripatoides occidentalis would

be collected at Bridgetown for purposes of comparison, but, owing

to heavy rains this year, that district was flooded at the time

when the expedition had been arranged.

P.Z.S.1914DAKIN PLL

we : if A Milf Y

Huth sc.etimp.

PHYLLOPODA FROM WESTERN AUS teens.

PB Z.3S.1914.DAKIN.PLIL,

key

1d.

Ie,

. Huth se, etimp.

PHYLLOPODA FROM WESTERN AUSTRALIA.

ON THE PHYLLOPODA OF W. AUSTRALIA. 293

20. Fauna of Western Australia.—II. The Phyllopoda of

Western Australia. By W.J. Daxin, D.Sc., F.LS.,

F.Z.S., Professor of Biology in the University of

Western Australia.

[ Received January 12, 1914: Read March 17, 1914.]

(Plates I. & II.*)

INDEX. Page

ID RSTRNT TH JNWSTEAIUE, Soogo5 co odcoscosceceosonesobonocamonscaoose AOL

Systematic :

Branchinella australiensis var. occidentalis, nov. ...... 296

1835 (OCTRAMOMBCLISOS, S> Wo coronansoccescocoe ban ssoseredeassoosooveca OS

Hulimnadia cygnorwm, Sp. 0. ...0.. 6c ccc cee eevee eee eeennneee 299

PS MUR E ECS US ASI alent aa iew nae neh ea nee ga ed he Paty ey le OO)

Cyzieus (Estheria) rufa, sp. Me... .....0.c0cccecceeeeee eres 801

Introduction.

The Phyllopoda of Western Australia have, up to date, been

known only from one or two specimens collected in the Murchison

District and examined by Sayce tT; from one or two specimens

worked at by Sars¢; and from those specimens collected by

the Hamburg Expedition to the South-West in 1905§. The

records are anything but extensive. Australia (as Wolf himself

states in his report upon these crustacea) is a country where

the conditions are particularly favourable for the presence of

Phyllopoda.

Western Australia is exceptionally well suited for these

animals, and they are to be found everywhere in large numbers

after the rainy season has commenced, They occur in small

rock-pools far inland, in swampy places with perhaps two or three

inches of water lying in depressions in fields, in temporary lakes

on the goldfields, and even in depressions filled with water—

extended cart ruts—on frequently used country roads.

Future investigation should bring to light many more forms,

for the area to be explored is indeed great. A glance at the

following table will serve to indicate the species discovered up to

date and also their distribution.

Out of the 33 to 35 Phyllopods known from the Australian

Continent previous to this paper, only two of those species found

on the eastern side of the island were known to occur in

Western Australia. Only nine species altogether had been re-

corded from Western Australia, and seven of these were peculiar

to the State.

I have been able to add to the list a number of new forms,

* Wor explanation of the Plates see p. 204.

+ Sayce, Proc. Royal Soc. Victoria, New Series, vol. xv. 1902.

~ Sars, Arch. for Math. og Naturvid. Christiania, xix.

§ Wolf, Phylopoda: ‘Die Fauna Siidwest-Australiens,’ Band iii. (Jena, 1910-

1911).

20%

PROF. W. J. DAKIN ON THE

{ | !

euenee dae eue H “BLIOJOI A .

| |

sod | boobs | citys cm e|ON > Meare |

| | |

| ‘Miay NOG | “[B4svop |

“BLUGUUSE T, | U.lo4oNy pues purpuy |

arse | ne | BALB WIITZAON | “Rav PUR[UT |

| |

| | |

siete | uwozdureypyooyy | °

| |

é po SN iO |= euenood [Pe ke Sed |

UOClOOK es eco = I = SP OOOO } Cry eee see |

| H |

| | curTeygnog | |

Baocou “Balv PUR[UT VY WlWYWON | Oey

SS : sone Sos | “AoupAg awany

HOPS ae eam Cae | aon Aeeoda

| |

“BIUBUISY J, *puejsiaan?t) | “BILOJOTA | *SOTRAA NOG

| MON

“PaPloval Atay SPOUMIS UT

“UILBISNY YNOS

‘qaed uwrey4nog

*]RA} Ua”

‘RITRAGSNW WING

“BI[RGSNW “Jap

“wlTBAysN YW “40a

qsvoy yynog

*RULIPUBXO] VW

eyVy awan

“WOTGaA [BISBOD

‘RITRISNY

Yyqnog

“WUBYION

ITH eutooag,

“UTULUUE T,

AI] LOOB[B YY

"RITRIISUY 489A

“ULVT[}.10 N

‘urddoowume

wy ulyqvioog

‘OIL 100.5[RY]

ANON PST

“RIPBRAGSU VY

ennieoenne FIOM Susojas “iva © iw

oor FIOM sngnhuoja Oty. T9

E ree ee pleg szp2.a snunprday

loon aan ee SES DSEEEEnEe "TO AA SmIUALGS #

eras DOU eeciCicanrcy ae ON supra n.Ab (73

(ea

wy saouedg) srsuarmugsny (sndp) sdorsy,

‘W CIsd014y J,

SIUg d Ldayoun snjppydasojdaigy

‘u'ds ‘sesuawny740U

see" (TOMA) S298 0.00H 107 sf

[ere sess (QahUG) sisuasla «

("tao ‘82709

[-UIPWIO "ABA 23 is

fps e (SloqYoy) seswarv.gsny ypjamyouds.g

“ g0AUG vUvIZIALz YIUWAZDADT

Pec sonal? os (Surly) d veavoud eg

po (Aa KBQ) sisuarpvugsam «

(®DABQ) SepV.ugsny vISIUWAgL

‘WAI dOdiIHONVUG

(on)

in|

PHYLLOPODA OF W. AUSTRALIA.

BOIB ULI NOG

“ULI YJNOG

“Sta

U.L91]J.10 NY

UOT INOS |

UWTOU 4.10 NT

“BOIv

“BVATB

“MS ON

“MSN

“xoupAG Iwan

dM SN

‘KaupAG Iwaut a

Avg Auvjog «vay

*"yaat/) s.aedocs

“ARE TLOATY,

“BITBISDY “Qual

‘BITBAISNY “QUIG

"]RAqZUAD

®W BAIv WIAYQNOG

“UIMIRG 410g

“BITRAYSN-Y “4Ua,)

‘RIpBIYSNW “499

“RILRAYSN YW

‘que ‘Avg IPOATY

“RI[BIYSUY *yUI_

w UIMIeGg WOg

“uo}fassng

“AYIQ lapnog

“UOSTI[I.AN IAT

“AV

yonqaoy lvayy

‘urddoovsang

WY ULGqViloogy

“WRT }.10 NY

‘Yad way

“UOJ LOSSU

" (ee ® taouedg) veuwawa

wee rec een ane wt eee eee (Ape) 109) “e

(SuLy) Vevh'vajanu snaouliry

“H CIGON AT

“ (pareg?) adopsry m2. MISTI

tees Spon d(H ED toured) uokjowp

Sa a ‘u ‘ds “‘pfiut se

eba0000 0o/((f pe Iq) DUADAQN) “

sae ene eee ese etree ese sesons (a0keg) USADS Gh

(sawg) snorgdrj)a pac

(Kpeigq) epunyand (nrwaygspp) snarztiy

ne IH ® laouaedg snawunwg ss

orf ~ TRH » toouadg 29707 ss

eH F teoueds 4 2aaanbs &

“(pareg.) 2 1youig sisdoppuuwuy

dass ~ SuLy nuvliajwn4s DIPDUUA] DLV T

terest) OAK DIDDY “

soostesresssss 99 KRG $28292.009020 ce

Ce iy ucds ‘sisuarlaf 77

Tes sees tr ds Gunwoubhia GG

Fete eee eee eee wee Apeig SISUa]OQIL ce

SULLY wpyp.os “s

SIRS 2YDp VIpHUULR

‘WIIG VNNIYT

296 PROF. W. J. DAKIN ON THE

and, what is perhaps more interesting, to show that some of the

species already known from the other States of the Common-

wealth occur also in Western Australia. The Western State is,

therefore, not so distinct from the other regions as the earlier

investigations might lead one to believe. There are now twelve

species or varieties which are known only from the West, and

there is no doubt that further investigation will reveal several

more new species and very probably a much wider distribution

for the species already known. The new records are from

specimens collected by Professor Woolnough, Mr. Alexander,

and the author. Two of the species discovered by the Hamburg

Expedition have been met with again.

EUPHYLLOPODA.

ANOSTRACA.

Family BRANCHIOPODID.

Genus ARTEMISIA.

ARTEMISIA WESTRALIENSIS (Sayce) *.

This species is the only Artemia known to occur in Western

Australia. Jt was collected in Lake Aurean in the Murchison

District in 1896, but no male specimens have as yet been

obtained.

xenus BRANCHINELLA Sayce, 1903.

This genus was instituted by Sayce to receive two Branchipods

which, though agreeing with Branchipus in very many ways,

differed in the male claspers being without any accessory branch

or spine, the second antenne of the female being long and ribbon-

like, and the penis of the male possessing certain peculiar

characters.

Since the genus was founded, the Hamburg Expedition

discovered one of the two species in Western Australia. This

was previously known only from Central Australia. The

expedition’s collection also included a new species. Up to date

no species of Branchipus has been met with in Australia. We

have rediscovered the species of the German expedition, extended

the distribution of an Hastern form, which, however, occurs here

in the form of a variety, and added still another species to the

list.

BRANCHINELLA AUSTRALIENSIS (Richters).

Variety occidentalis, nov. (PI. I. figs. 1-5.)

The species Branchinella australiensis, a large and stout Bran-

chipod, was first described by Richters fT from specimens from

* [The parentheses around the names of authors placed after scientific names in

this paper are used in accordance with Article 23 of the International Rules of

Nomenclature (Proc. 7th Int. Cong. Boston 1907, p. 44. (1912)).—Epitor. |

+ Richters, Journ. de Muséum Godeffiroy, xii., 1876.

PHYLLOPODA OF W. AUSTRALIA. 297

Queensland. The description was, however, overlooked until

re-discovered by Hall and Sayce. The latter found the species

to be quite common near Melbourne. It occurs in Central

Australia, the coastal area of Southern Australia, and in North

and South Victoria.

In fact, its known distribution was pretty general, and only

Western Australia seemed to be without it.

Two large Branchipods have been investigated in the course of

this work, both unfortunately of the same sex. They agree in

very. many points with the description given by Sayce.

Description of male (fig. 1).— Body stout: trunk of equal length

to tail. Branchial feet with large covering plates. The six

endites are almost the same as in the figure given by Sayce ™.

The position of the flabellum is different, and the relative size

of covering plate (proximal exite) and flabellum is considerably

different. Claspers large and powerful (fig. 3), basal joint with

inner surface clothed with minute teeth, the second joint longer

in proportion to the first than in typical Branchinella australiensis.

The inner surface of the clasper bears curious transverse over-

lapping shelves (fig. 4). These have the appearance of transverse

ridges. They end abruptly along a regular line at each side.

Frontal appendages invisible in specimens. First antenne re-

markably long, filiform, and about three times the length of the

second antenne (fig. 2), and quite different from those four ed by

Sayce for Branchinella australiensis. Caudal rami no longer than

the last four segments of tail. Sheath of penis not notched (penes

were not everted). Length inclusive of caudal rami 35 mm.

The resemblance to Br ranchinella australiensis seems to be great

enough to consider this as a variety of form. The outhatandime

difference is in the first antenne and the reduction of the frontal

appendages.

Locality. Lake Violet (near the centre of Western Australia).

BRANCHINELLA LONGIROSTRIS (Wolf).

This characteristic form was first discoveved in the course ot

the Hamburg Expedition, and had only been recorded this

once from Australia. It has now been collected again (seven

years later) from similar situations in the district where it was

originally found.

Both males and females occur. The characteristic huge frontal

appendage of the male cannot possibly be mistaken.

For complete diagrams see Wolf f.

Locality. Small rain-water ponds in depressions in rocks at

Burracoppin. They occurred with the same companions found

by the Hamburg Expedition, viz. specimens of Hilimnadia

badia.

* Proes. R. Soc. Victoria, N.S. xv. pl. xxx.

+ Wolf, Phyllopoda: ‘ Die Fauna Stidwest-Australiens,’ Band iii. (Jena, 1910-1911).

298 PROF. W. J. DAKIN ON THE

BRANCHINELLA NORTHAMENSIS, sp. n. (PI. I. figs. 6-8.)

Description of female.—No males of this species have been

discovered. The female is an undoubted Branchinella. In general

appearance and size the specimens resemble the females of

Branchinella longirostris, and are much smaller and more delicate

than Branchinella australiensis. The living animal is colourless

and semi-transparent. Eyes very prominent. The head and

limb-bearing region of the trunk are together much longer than

the limbless posterior region (termed the tail by Sayce).

Ist antenne filiform and about 14 times the length of the

second antennee (see fig. 8).

2nd antenne more or less ribbon-like but not flattened very

much. They decrease but little in width until they terminate in

a rounded end with a sharp point which is situated abruptly at

the apex.

The ovisac (fig. 7) is very slender, without lateral expansions

proximally, and extends posteriorly to the middle of the 4th

limbless segment (tail- segment). It possesses the characteristic

ridge on the ventral surface.

The caudal rami are well developed, provided with strong sete

and equal in length to the last four segments of the body

(fig. 6). Length 13mm.

This species is distinct from Branchinella australiensis in

(1) General character, size, and shape.

(2) Form of appendages.

(3) Antenne.

It is distinct from Branchinella eyrensis by

(1) Frontal appendage.

(2) Size and general form.

(3) Ovisac.

It is distinct from Branchinella longirostris by

(1) 2nd antenne.

(2) Ovisac.

BRANCHINELLA EYRENSIS (Sayce).

This species was recorded by the Hamburg Expedition from

Kalgoorlie on the goldfields.

Only two specimens were obtained, and they were in bad

condition. No other specimens have so far been met with.

For diagnosis see Sayce.

CONCHOSTRACA.

Family LiMNADIID#.

Genus HULIMNADIA.

Only one species had been found hitherto in Western Australia,

and that by the Hamburg Expedition in 1905. This was

recognised as a new species by Wolf, so that up to the date of

PHYLLOPODA OF W. AUSTRALIA. 299

the present report, none of the Hulimnadia occurring elsewhere

in Australia had been found in the West.

We have rediscovered Wolf's species and can confirm his

description of the same.

In addition, we have two new species and the first record of an

Eastern form.

EULIMNADIA CYGNORUM, sp.n. (PI. I. figs. 9-13.)

Shell compressed, thin, without trace of umbones (fig. 11).

Lines of growth with great difficulty discernible and only about

three visible.

Dorsal line evenly arched, with highest point anterior to

middle. General shape more resembling Hulimnadia rivolensis,

but with a more obtuse posterior end.

Head with rostral expansion in male, moderately long and

bluntly pointed. In female much shorter and not acute (figs. 9

& 11).

Legs —not possible to make out exact number, but apparently

not exceeding 18 pairs in female.

Male with anterior legs subchelate—developed as claspers

(fig. 13). The hooked termination, which is covered with small

pits, carries a prominent stalked disc. The subapical appendage

is extremely long in this species, much longer than in either

Hulimnadia badia Wolf or Lulimnadia rivolensis Brady.

There are two distinct segments, but it is possible that the last

extremely long one is in reality two, as in Hulimnadia rivolensis.

In any case the segmentation is very much less marked than

in this latter species, specimens of which have been carefully

compared with it. The appendage is longer than in any other

Australian Hulimnadia. There is an angular prominence above

the terminal hook, just as in Hulimnadia badia and Hulimnadia

rivolensis. Whilst, however, it is not so large as in the former

species, it stands away from the claw much more than it does in

Huhmnadia rivolensis. ;

The caudal claws (fig. 12) are slightly bent upwards, the

proximal part very slightly so. The distal end is provided with

a row of sawlike minute prickles. The proximal half bears a few

feathered set.

The lateral plates of the tail are almost straight and bear

about 20 acute denticles. The most posterior one is large and

perfectly straight, differing in this respect from Hulimnadia

rivolensis. All the spines are provided with minute sawlike

points along the sides.

This species is characterised by

(1) Shell and lines of growth.

(2) Anterior legs of male.

(3) Rostrum.

(4) Caudal claws and tail.

Specimens, both male and female, obtained from Cannington,

near the Swan River.

300 PROF. W. J. DAKIN ON THE

EKULIMNADIA FERIENSIS, sp. n. (PI. II. figs. 14-18.)

A considerable number of specimens, both of males and of

females, was obtained from a little patch of flooded bushland at

Northam. In the same pool occurred Branchinella and swarms

of tadpoles.

Shell (fig. 14) very thin and pellucid. Of same appearance in

both sexes. No lines of growth visible. In lateral view almost

equilateral. The dorsal margin slightly vaulted, the greatest

height being only a shade in front of the middle. Shell

compressed,

Head (figs. 15 & 16). No great difference between the sexes.

Rostrum short and bluntly rounded.

Legs 16-18 pairs. Anterior pairs in males subchelate

(fig. 18); not particularly robust. Armed with numerous sete.

No process on inner side of band above thumblike projection

(cf. figure of Hulimnadia cygnorum).

Tail with caudal claws (fig. 17). The proximal part of the

claws bears a few feathered sete. The region, not quite half of

the total length, bears a well-marked sawlike row of minute spines.

The two lateral plates terminate in two long processes, armed

with minute denticles, and the upper edges of the plates in the

female bear about 16 to 18 spines of unequal length. In the

female there is also a tendency for the most posterior of the spies

to be arranged in groups of 4 or 5, the size of spines in each

group diminishing anteriorly (fig. 17). There are only about

12-14 spines in the male. All the spines are armed with minute

denticles.

Size. Length of specimens 5°5 mm.

KULIMNADIA RIVOLENSIS Brady.

This species was described originally by Brady i in 1886. Up to

this present report it was known from Southern Australia,

Central Australia, and Southern Victoria, with the possibility of

an extension into New South Wales. Now it has been found in

Western Australia, and not where it might have been expected,

inland on the goldfields, but down on the South-West Coast in

small ponds near Busselton. Several specimens were obtained

there by Mr. Alexander, and the collection includes both males

and females. There is nothing further to add to the descriptions

previously given by Brady, Spencer and Hall,* and Sayce.

This was the only Hulimnadia obtained by the expedition to

Central Australia. This discovery of its occurrence in Western

Australia is particularly interesting, because it is the first

Kulimnadia of the Kast and Central States to be recorded from

the West.

* Spencer and Hall, Reports Horn Expedition to Central Australia, Part II.

PHYLLOPODA OF W. AUSTRALIA. 301

EULIMNADIA BADIA Wolf.

This species was the only Hulimnadia known from Western

Australia before this report. It was discovered by the Hamburg

Expedition in pools together with Branchinella longirostris. As

mentioned in discussing that species, Hulimnadia badia has

again turned up with its fellow companion. I can confirm the

description of Wolf, but must make a slight alteration, or rather

an addition, to his diagnosis of the species. ‘The specimens were

collected. by Professor Woolnough. They are the largest and

most robust of our Hulimnadia. Fortunately I have been able to

compare these specimens with the type in the Hamburg Collection.

Wolf remarks that never more than five lines of growth can be

observed on the shells. I have recognised six and even seven on

some specimens. Another slight point of difference is that the

shell-gland is stated to occur always inside the first line of growth.

This is frequently the case but not always, and often extends over

the first line. In Wolf’s drawing it is indicated well inside the

lines of growth. One other point is that the head in the region

of the eye does not protrude in such an angular manner as figured

and described by Wolf. There is absolutely no doubt about the

identity of the two sets of specimens. The characteristic features

are all obvious, and it is well to emphasise these slight variations

perhaps due to age or the season, especially since they have been

touched upon and figured as of some hittle diagnostic value.

Both males and females occurred in rock-pools near Burracoppin,

not very far from Boorabbin, where originally discovered.

Genus Cyzicus.

Cyzicus (EsTHERIA) RUFA, sp.n. (PI. IL. figs. 19-21.)

Five species have been recorded from Australia, and out of

this number two were from Western Australia. These were

determined by G. O. Sars and Sayce respectively. No specimens

were met with by the German Expedition. ‘The species of Sarsis

Cyzicus elliptica, and that of Sayce Cyzicus sarsit.

The new species, Cyzicus rufa, was collected in pools at

Lakeside, Boulder City, by Mr. Alexander. Two individuals

were at my disposal, but unfortunately both were females.

Characters. Shell, seen laterally, of a rather quadrangular form

(fig. 19), both ends being rather too obtusely rounded to give the

impression of an ellipse. Anterior-posterior length rather great

for the dorso-ventral depth, and much longer than that of any of

the other Australian species. Umbone prominent and situated

much nearer the anterior end (about one-fourth the total antero-

posterior diameter from the anterior end of the shell). The

ventral margi of the shell is almost straight for some distance.

Both ends are evenly rounded, and the axis of greatest length lies

midway between the dorsal and ventral margins of the shell.

Lines of growth about fourteen, with sete. The sculpture of the

302 PROF. W. J. DAKIN ON THE ~

shell is reticulate and irregular. The valves are red-brown in

colour and of firm consistency.

Rostrum (fig. 21) pointed and moderately long.

The number of legs is less than 20 pairs.

Tail of usual shape (fig. 20). The caudal plates have their

upper margins almost straight. Each plate bears about five equal

denticles and some smaller ones.

The caudal claws bear feathered setee on the proximal half.

Eggs spherical.

Size. Length of adult female, 8°5 mm.

The species Cyzicus rufa differs from Cyzicus lutraria in

(1) Shape of shell—straight ventral margin, convexity of

anterior end. No compression behind middle.

(2) Caudal plates not concave dorsally.

(3) Rostrum of female moderately long.

(4) Much smaller number of appendages.

The species differs from Cyzicus elliptica in

(1) Shell—ridges and shape generally.

(2) Tail.

(3) Size.

(4) Number of appendages.

The species differs from Cyzicus sarsii in

(1) Shape of head.

(2) Shape of shell and number of ridges.

(3) Caudal plates.

The species differs from Cyzicus packardi in

(1) Shell—general shape and lines of growth.

(2) Shell sculpture.

(3) Caudal plates.

Oyzicus dictyon (Spencer & Hall) seems to be synonymous with

Estheria lutraria.

There is a possibility that this might be the female of Cyzicus

sarsit, a species founded by Sayce on one example,a male. The

differences in the shell, etc., are so great that the author considers

it more probably a new species.

Family LyYNCEID&.

Genus Lynceus.

Three species of the genus Lynceus have been recorded from

Australia, and of these none is supposed to occur in Western

Australia.

The record that I have to note therefore shows still further

that the supposed absence of East Australian species of Phyllopoda

PHYLLOPODA OF W. AUSTRALIA. 303

from the Western State is probably due to the small number of

collections that have been made.

Lynceus tatei (Brady).

Several specimens were collected in pools at Busselton, South-

West Australia. They agree in almost all respects with the

description by Sars and the figures by Brady.

There is a slight difference in the shape of the rostrum, but

this is so small that it seems hardly sound to make a new variety

out of the specimens at my disposal. The collection included

both males and females. The size was much smaller than the

figure given by Sayce. Specimens varied between 2°5 mm. and

3°79 mm.

Lynceus tatei is now known from Victoria, New South Wales,

South Australia, and Western Australia.

Family TRiopsi1p#(A PODID!).

Genera Apus and LeprpuRus.

The family Triopside contains but two genera, Apus (7’riops)

and Lepidurus. Both genera occur in Australia, and as a result

of the Hamburg Expedition both were recorded from Western

Australia. Apus itself has been recorded previously from

Western Australia. Both genera appear to be pretty common in

pools scattered about the State, and although specimens have

only been obtained from one or two places, there is evidence that

their occurrence is by no means restricted to these spots.

Genus LEPIDURUS.

Wolf, in his paper on the Phyllopoda of the Hamburg

Expedition, remarks on the great difficulty in diagnosing species

of this genus. The number of the segments which bear no

appendages varies in one and the same species, and in no form is

the number eight exceeded. Large numbers of specimens are

required before one can feel certain of any systematic work. The

result is that five species recorded from Australia are considered

to be all identical—the species being then Lepidurus viridis

Baird. Wolf found, however, that there were some essential

differences in the specimens from different localities, which he

had received for examination, and as a consequence divided his

examples into two new varieties. These are Lepidurus viridis

Baird, var. elongatus Wolf, and Lepidurus viridis Baird, var.

setosus Wolf.

I have met with specimens which are evidently Lepidurus

viridis var. elongatus, and from quite a new locality, namely

Northam.

304 PROF. W. J. DAKIN ON THE

GLEPIDURUS ViIRIDIS Baird, var. ELONGATUS Wolf.

Specimens occur regularly every year about July and August

in very shallow, muddy pools on a road at Northam.

The characteristic carapace covering all the segments of the

body and only allowing a glimpse of the caudal rami enables one

to recognise the similar ity to Wolf’s specimens.

The number of segments not bearing legs is as usual, five. The

length of my specimens averaged about 28 mm. without the

caudal appendages, and these brought the total length to 53 mm.

This size is greater than that of the largest of Wolf's specimens.

LEPIDURUS VIRIDIS.

A large number of specimens of Lepidurus has been obtained

from waters at T'iammin, Western Australia. These individuals

are certainly not Wolf’s variety Lepidurus viridis var. elongatus,

and still more certainly not his variety setosws. They may be

described as follows :—The large examples average about 20 mm.

for length of carapace, with a total length of 40 mm. counting the

long anal furea.

The carapace is broadly rounded in front and somewhat

considerably vaulted behind, with a prominent keel down the

middle lir The posterior notch in the carapace is practically

semicircular.

The long axes of the eyes run at an angle to each other to

meet anteriorly. The anterior and posterior margins of the

erescentic eyes are of similar shape and extent.

The “ Nackenorgan” projecting between the two eyes and

slightly posterior is circular in shape.

lt seems undesirable to name these specimens anything but

the simple Lepidurus viridis. From the difficulty of recognising

varieties it would appear that a comparative study of the genus is

required. This could only be carried out successfully with a very

large collection of individuals from many localities in different

parts of the world.

EXPLANATION OF THE PLATES.

Prats I.

Branchinella australiensis, var. occidentalis, noy.

Fig.1. Male specimen. X 1°5.

2. Head and appendages of male. X 3°5.

3. Antenna or “clasper” of male. X 5.

4. Surface of antenna considerably enlarged.

5. 4th trunk-limb of male. X 6.

Branchinella northamensis, sp. n.

Fic. 6. Posterior segments and caudal rami. % 10.

7. Genital segments bearing ovary. X 10.

8. Head and appendages of female. xX 8.

PHYLLOPODA OF W. AUSTRALIA.

Eulimnadia cygnorum.

Fig. 9. Head of male. X 20.

Vig.

10. Shell. XX 4.

11. Head of female. > 18.

12. T'ail and caudal claws. X 12.

13. Ist trunk-limb of male. > 40.

Pxate IT,

Hulimnadia feriensis, sp. n.

14. Shell. x 5.

15. Head of male. 20.

16. Head of female. x 20.

17. Tail and caudal claws of fema.-

18. Ist trunk-limb of male. x 50.

Cyzicus (Estheria) rufa, sp. n.

g. 19. Shell. x 3°5.

20. Tail. x 10.

21. Head of female. > 20,

x 10.

ON EAST AFRICAN MAMMALS. 307

21. Notes on a Collection of Fast African Mammals pre-

sented to the British Museum by Mr. G. P. Cosens. By

Guy DoLiman *.

[Received February 12, 1914; Read April 7, 1914.]

INDEX.

Systematic :— Page

CEROWILS COSEPB0, EDs We cao ceoscceoss00n0 soo cagceo-no cor dll

DUEPOUS UORBBIy BD We 00000060005 09006008 000 060 050060 312

Epimys walambe amala, subsp. 0..........0:.0 ee 818

Hypimys coucha pallida, subsp. n. ......cseceeee veces 314

Arvicanthis rumruti pallescens, subsp. n. ......... 316

3138

Cephalophus grimmia lutea, subsp. nN. ...... 22.26...

I have the pleasure of furnishing a report on the Mammals

obtained by Mr. Willoughby P. Lowe during the recent East

African Expedition organized by Mr. G. P. Cosens. The entire

collection, some two hundred specimens in all, has been presented

by Mr. Cosens to the British Museum.

Jn working out this collection I have found it necessary to

describe six new forms 7, the most interesting of which, the new

Gerbil (Gerbillus cosensi), I have named in honour of the generous

donor. Another interesting novelty, Z'aterillus lowei, 1 have

ealled after Mr. Willoughby Lowe, the collector.

The East African mammals were obtained in the Naivasha and

Nyanza Provinces. The route taken was from Kijabe to

Mt. Suswa, where the first collecting was done, then westwards

across the Loita Plains and Lemek Valley to the Amala or Mara

River. After ten days’ collecting along the Amala River the

expedition turned south towards the Anglo-German Boundary

and then eastwards, via Leganisho, to Lengototo, the Narossura

and Southern Guaso Nyiro Rivers. Further collecting was done

along the southern Guaso Nyiro, and the party then proceeded

northwards as far as the Mau Escarpment, returning to the

railway at Naivasha.

The Uganda Expedition started from Baringo in December,

1912. and travelled in a north-easterly direction across the Kerio

and Wei Wei Rivers, and then along the Turkwel as far north

as Ngamatak. Turning westwards the party reached the Kozibir

Hills on January 22nd, 1913, and Mt. Maroto on January 28th.

Proceeding in an easterly direction, via the Nakwai Hills and

Lobor Mountains, Kamchuru was reached on February 9th,

and here the greater part of the collecting was done. Other

* Communicated by OLpFIELD Tuomas, F.R.S., F.Z.S., and published by per-

mission of the Trustees of the British Museum.

+ [The complete account of the new forms described in this communication

appears here, but since the names and preliminary diagnoses were published in the

“ Abstract,’ No. 131, 1914, these species are distinguished by the names being under-

lined—Eprror. ]

Proc. Zoot. Soc.—1914, No. XXI. 2

308 MR. GUY DOLLMAN ON

mammals were obtained on the journey from Kamehuru, via

Pader and Falabek, to Nimule.

In the following list the East African and Uganda Collections

are not treated of separately, as it seems more convenient,

the two areas not being faunistically distinct, to consider the

collections together.

1. Pavto rurAx Elliot.

6.73. 12 miles east of the Amala River, British Hast

Africa.

3. 70. Amala River.

3. 74. Hast of Amala River.

The skulls of the two adult males (Nos. 73 and 70) are sur-

prisingly different in general structure, especially as regards the

nasal regions and dental characters. The difference may be of

specific value, as it is not rare to find two nearly allied species

of Baboons existing in the same locality; with such variable

animals, however, I do not feel justified in dealing with this

question until further material is available for examination.

2. CERCOPITHECUS PYGERYTHRUS CENTRALIS Neum.

75. 12 miles east of Amala River.

- 153. Wei Wei River, Rift Valley.

. 127. 20 miles south of Baringo.

. 137. 30 miles N.W. of Baringo.

EPoMoPHORUS MINOR Dobs.

. 111. Southern Guaso Nyiro.

EpomopHorus ANURUS Heuglin.

. 186. Mt. Maroto, Uganda.

LAVIA FRONS FRONS EH. Geoff.

. 40. Amala River.

. 134,138. 30 miles N.W. of Baringo.

. 192. Bakora, Uganda.

. Pader, Uganda.

- 172; 2. 171. Kozibiri River, Uganda.

. 184,188. Mt. Maroto, Uganda.

PIPISTRELLUS DESERTI Thos.

. 185. Mt. Maroto, Uganda.

. 175. Kozibiri River, Uganda.

. 151, 152. Wei Wei River, B.E.A.

OO; Oy LP Gao tororg @ i toy 2 Aartoa ec

7. SCOTOPHILUS NIGRITA Schreb.

3. 140. 30 miles N.W. of Baringo.

179 (in spirit). Mt. Maroto, Uganda.

EAST AFRICAN MAMMALS.

8. SCOTEINUS SCHLIEFENI Pet.

OEGOl Kozibini nnver:

6. 133. 30 miles N.W. of Baringo.

SG. 148, 149, 150, 154. Wei Wei River.

6. 158, 159,166, 167. Turkwel River.

9. TAPHOZOUS MAURITIANUS HK. Geoff.

®. 168. Turkwel River.

10. ELEPHANTULUS DUNDASI Dollm.

do. 145, 12 miles N.W. of the Kerio River, B.E.A.

309

The exact locality where this specimen was obtained 1s not

more than 75 miles N.W. of Baringo, the type locality of the

species.

11. NasrtLio BRACHYRHYNCHUS ALBIVENTER Osg.

3.90; 2. 86. Lengototo, B.H.A.

12. CrocipurA monax Thos.

©. 95. Amala River.

13. CrocipuRA JACKSONI Thos.

6. 50; 2. 63, 66. ~Amala River.

?. 99. Narossura River.

89. Lengototo.

14. FELis CAPENSIS HINDEI Wrought.

3.46. Amala River.

A. fine example of the melanistic phase so frequently met with

in these Serval Cats.

15. GENETTA ERLANGERI Matsch.

3.19. Southern Guaso Nyiro.

@. 33. Amala River.

62345 Q. dil, Ivemelk Walley

@. 80,81. East of Leganisho.

16. Icronyx CAPENSIS ALBESCENS Heller.

¢. 18. Southern Guaso Nyiro.

The type locality of Jctonyx c. albescens is given by Heller as

Mt. Lololokwi, Northern Guaso Nyiro, a long way north of the

Southern Guaso Nyiro; this author also records specimens of

albescens from Nairobi, and it seems probable that this race

extends over a very large part of the Protectorate.

17. MuNnGos ALBICAUDA G. Cuv.

@. 37. Amala River.

18. MunGos sANGUINEUS IBEa Wrought,

3. 102. Narossura River.

21*

310 MR. GUY DOLLMAN ON

19. HELOGALE PERCIVALI Thos.

3. 166. Turkwel River.

The type of Helogale percivali was obtained by Mr. Percival in

the Orr Vailey, Mt. Nyiro. This specimen from the Turkwel

River agrees with the unique type in all respects.

20. HELOGALE UNDULATA RUFULA Thos.

@. 110. Southern Guaso Nyiro.

21. HELOGALE victorina Thos.

SG. 43, 59,62; 2. 42, 44,61. Amala River.

22. CANIS LATERALIS Sclater.

3. 69. Amala River.

2.176. Kozibiri River, Uganda.

23. HELIOSCIURUS MULTICOLOR ELEGANS Thos.

©. 142. Suk Plains, Kerio River.

©. 162. Doroto District, Turkwel River.

3. 181,182. Mt. Maroto, N.E. Karamojo, Central Province,

Uganda.

g. 198. Nakwai Hills, N. of Lake Kirkpatrick, Central

Province, Uganda.

6. 148. N.W. of Suk Plains, on the Wei Wei River.

This series is rather variable in colour, possibly due to

bleaching. The type locality of elegans is Mt. Elgon, so that

this race would appear to extend northwards over a very large

area.

24, PARAXERUS OCHRACEUS ELECTUS Thos.

3. 155. Ngaboro, Wei Wei River, Rift Valley.

¢. 157. Ngeronomi, Turkwel River.

@. 156. Ngabotok, Kilebor, Turkwel River.

These three Squirrels are quite similar to the typical speci-

men of electus, described by Thomas from Elgoyu, British Kast

Africa. Mr. Kemp obtained a series of this race on the Lai-

kipia Plateau; it is worthy of note that all these specimens are

exactly like the type, there apparently beimg no colour variation

at all.

95, XERUS DABAGALA DORSALIS Dollm.

Co laut, Ses Ieraunee.

3g. 126. 40 miles N.W. of Nakuru.

6s 143 O, 144. 9 Suk Blais: 902 miles. NW. sol Kerto

River, B.H.A.

Xerus dabagala dorsal's was founded on a series of specimens

collected by Mr. Kemp at Baringo, so that Nos. 131 and 182 are

topotypes, and agree very closely with Mr. Kemp’s specimens.

EAST AFRICAN MAMMALS, 311

26. GRAPHIURUS BROCKMANI INTERNUS Dollm.

©. 100. Narossura River.

@. 105, 109. Southern Guaso Nyiro,

These Dormice do not appear to differ from the specimens

collected by Mr. Blayney Percival and Mr. Robin Kemp on the

Northern Guaso Nyiro, the type locality of this race.

27. DipopiLLus HARWoopI Thos.

3. 118. Southern Guaso Nyiro.

This diminutive Gerbil is evidently widely distributed over

the southern portion of British Hast Africa. The type locality

of the species is Naivasha, but it would appear to extend much

further south than many Naivasha mammals; Dr. Bayer obtained

a Dipodillus harwoodi from as far south as the Tsavo River.

This specimen, which is in the collection of the Congo Museum,

Tervueren, | have recently had an opportunity of examining,

and there would appear to be no doubt that it represents the

Naivasha species.

28. GERBILLUS COSENST Dollm.

Abstract P. Z.S. 1914, p. 25. (April 14.)

$. 170. Kozibiri River, Ngamatak, Turkwel River. Alti-

tude 1800 feet.

Allied to Gerbillus dunni Thos., the Somali species, but readily

distinguished by its duller colour and smaller size.

General proportions rather less than in dunni, the hind foot

measuring only 25 mm. in length.

General colour of dorsal surface pale buff mixed with slate-

erey, the grey tint most obvious on the shoulders and down the

middle of the back; flanks brownish buff, near ‘‘ warm buff”

(Ridgway, 1912). Head similar in colour to back, the slate-grey

bases of the hairs showing through the yellowish-buff tips,

Sides of face white; pale greyish-buff markings extending from

below the eyes to the ears. Backs of hands and feet white.

Entire ventral surface pure white. Dorsal surface of tail much

darker than in dunni, above dirty brown, below white.

The skull of this unique specimen is unfortunately badly

broken, all the region posterior to the frontals and palate being

missing. Compared with the skull of the Somali species the

general structure would appear to be very much the same, but

rather smaller throughout.

Dimensions of the type (measured in the flesh) :—

Head and body 78 mm.; tail 129; hind foot 25; ear 14.

Skull (posterior part of skull broken): length from fronto-

parietal suture to tip of nasals 19°5 mm.; length of nasals 10;

greatest breadth across nasals 2-4; interorbital constriction 5;

length of anterior palatal foramen 4; length of posterior palatal

cavities 2°2 ; alveolar length of upper molar series 4.

312 MR. GUY DOLLMAN ON

Type. Adult male. B.M. No. 13.10.18. 64. Collected January

Nrfislo, IS),

The donor and collector are to be congratulated on the dis-

covery of the genus Gerbillus on the Turkwel River. The

Somali species would appear to be the nearest ally of this new

form, though doubtless further collecting will bring to light

several other races of East African Gerbillus.

29. TATERILLUS EMINI Thos.

Ge 2022038204206. Dike QS e219 ORe 205 ea A Moe

Kamehuru, Lobor, Uganda.

3S. 225. Falabek, 45 miles east of Nimule.

The general colour of this series is very much as in the type,

a specimen collected by Emin Pasha at Wadelai. The young

specimen are all rather darker than the adults, especially on the

back and hind quarters.

30. TATERILLUS LOWEI Dollm.

Abstract P. Z. 8. 1914, p. 25. (April 14.)

g. 165. 10 miles west of the Negamatak Hills, Turkwel

River. Altitude 1800 feet. é'

A very pale coloured species allied to Taterills nabiles Dollm.

In general proportions this Gerbil is quite similar to mubilius

and the allied forms.

Colour of dorsal surface pale dirty yellow, a mixture of ** vina-

ceous-buff” and “ warm-buff” (Ridgway, 1912); the central

portion of the back is rather darker, owing to the dark hair-

tips being more developed. Flanks purer and yellower in colour.

Head same colour as back; sides of face as in nwbiluws, but rather

paler. Backs of hands and feet creamy white. Mntire ventral

surface pure white. ‘Tail much as in nwbhilus but paler, and with

the cream-white ventral surface more sharply marked off from

the dull upper surface.

Skull very like that of nwdilus, slightly smaller throughout.

Dimensions of the type (measured in the flesh) :—

Head and body 107 mm.; tail 160; hind foot 28; ear 18.

Skull: greatest length 343 mm.; basal length 28°3; zygo-

matic breadth 16°5; length of nasals 13°5: width of brain-case

(across squamosal region) 145; interorbital constriction 6;

length of anterior palatal foramina 5°8; length of posterior

palatal foramina 3-5; alveolar length of upper moiar series 4°8.

Type. Adult male. B.M. No. 13.10.18.66. Collected on

January 15th, 1913.

This species is the palest of all the East African forms, the

only members of the genus at all similar in colour being the

West African species Vaterillus lacustris and Taterillus mgeric.

31. TATERA NIGRICAUDA Pet.

$. 107; 2. 113,114. Southern Guaso Nyiro.

EAST AFRICAN MAMMALS, 313

32. DeyxpRomvs LINEATUS Heller.

¢. 208, 209. Kamechuru, Lobor.

These two specimens show the same variation as regards the

distinctness of the median stripe as is mentioned by Heller in

his description of the species, the type locality of which is

Lado.

33. DENDRomUS AcRmUS Wrought.

go. 45. Amala River.

34. DENDROMUS NIGRIFRONS True.

©. 60. Amala River.

35. Mus Betius Thos.

@, 124. Lake Naivasha.

36. Mus BELLUS GoNDOKOR# Heller.

@. 183. Mt. Maroto, Maroto Stream, Central Province,

Uganda.

37. Mus MUSCULOIDES EMESI Heller.

3g. 227. Falabek, east of Nimule.

38. Mus eratus Thos.

6. 56. Amala River.

39. Mus triton Thos.

ey 8,9. Mau; BELA:

40. Krimys WALAMB& AMAL Dollm.

Abstract P.Z.8. 1914, p. 25. (April 14.)

. 83, 84; 9. 82. Narossura River.

. 38, 58, 72. Amala River.

. 29, 30. Lemek Valley.

. 91; 2. 88. Lengototo,

Closely allied to Hpimys walambe pedester Thos. ; distinguished

by its smaller size, buff-tinted flanks, and much whiter ventral

surface.

Dimensions of bedy and hind foot considerably less than in

pedester.

General colour of dorsal surface quite like that of the Uganda

form. Flanks washed with buff, the cold grey tint so conspicuous

in pedester is here quite hidden by the buff coloration. Backs

of hands and feet yellowish white. Ventral surface pale slate-

grey covered with creamy white, the resulting effect being

distinctly purer and whiter than in pedester, where the belly is

quite grey. ‘ail rather lighter in colour, especially on the dorsal

surface.

Skull smaller, with smaller molars and less inflated auditory

bulle.

Q, +0 & ay

314 MR. GUY DOLLMAN ON

Dimensions of the type (measured in the flesh) :—

Head and body 144 mm.; tail 127; hind foot 26; ear 17.

Skull: greatest length 37°5 mm.; basilar length 31-3; condylo-

incisive length 36; zygomatic breadth 20; interorbital con-

striction 5:5; length of nasals 15:5; palatilar length 18; length

of palatal foramina 9°3; alveolar length of upper molar series 6-7,

Hab. Lemek Valley, between the Amala River and Southern

Guaso Nyiro. Altitude 6500 feet.

Type. Old female. B.M. No. 13.10.18.111. Original number

30. Collected on October 8th, 1912.

This Rat is readily distinguished from the Uganda race by

its smaller size, whiter belly, and buff-coloured flanks. Since

pedester was described more fully adult specimens of this race

have been received. The following dimensions are taken from an

old individual collected by Mr. Robin Kemp at the type locality:

Head and body 196 mm.; tail 151; hind foot 33; ear 23:5.

Skull (broken): palatilar length 20-2 mm.; length of nasals 17;

zygomatic breadth 23°6; alveolar length of upper molar series 7-9.

41. Errmys mepicatus Wrought.

¢. 141. 45 miles N.W. of Baringo.

42, EpIMYys JACKSONI de Wint,

do. 49. Amala River.

43. EPIMYS NIVEIVENTRIS Osg.

3. 97. Narossura River, B.E.A.

A very dark specimen. The large series of this interesting

Mouse collected by Mr. Pereival show that it is subject to very

considerable colour variation.

44, Epimys coucHa PANYA Heller.

6.2. Mt. Suswa.

g. 16,17. Western Slope of Mau Escarpment.

g¢. 35. Amala River.

Heller’s type came from the Athi Plains. The specimens

from Mt. Suswa and Mau are thus nearly topotypes, and appear

to agree very closely with the description and with the topotypes

in the Museum Collection.

45, Kpimys COUCHA PALLIDA Dollm.

Abstract P.Z.S. 1914, p. 25. (April 14.)

6. 199, 210, 211, 216, 220. Kamehuru, Lobor, Central

Province, Uganda.

Allied to Hpimys c. neumant Hell., smaller in size and without

any bufi-coloured suffusion on the ventral surface.

General proportions less than in newmani and effectws Dollm.,

hind foot only 20-22 mm. in length.

Colour of dorsal surface dark brown, between ‘‘olive-brown” and

. S ” . 2 i . ". |. a

st hair-br own (Ridgway, 1912), lined with black and washed over

with pale buff, this latter tint most dominant on the flanks and

EAST AFRICAN MAMMALS, SUS

forming a yellowish line between the dark flanks and pale under-

parts. Backs of hands and feet white. Ventral surface of body

slate-grey washed with white. Tail short, rather paler below

than in newmant.

Skull small and slight in build, very like that of the other

races of coucha.

Dimensions of the type (measured in the flesh) :—

Head and body 87 mm.; tail 100; hind foot 21; ear 19.

Skull: greatest length 28 mm.; basilar length 22; condylo-

incisive length 25:7; zygomatic breadth 13:2; interorbital

constriction 4; breadth of brain-case 12; length of nasals 11:2;

palatilar length 12°6; length of palatal foramina 7; alveolar

length of upper molar series 5.

Type. Adult male. B.M. No. 13.10.18.99. Original number

220. Collected on February 14th, 1913.

While working on this group of multi-mammate mice I have

come to agree with Heller in considering the forms newmanc,

panyd, effectus, and other allies as races of cowcha, and as such I

have described this Kamchuru mouse.

46, 'THAMNOMYS SURDASTER, subsp.

3. 87. Lengototo, B.H.A.

This specimen is so young that it is impossible to say to which

of the many races of surdaster it belongs.

AT, ZELOTOMYS HILDEGARDE® Thos.

@. 76. West of Leganisho, B. EH. A.

So WB. Wem, Jods

6. 5. Mt. Suswa, B.E.A.

The general external characters of this genus are well illustrated

in these three specimens. ‘The light coloured tail (due to the white

skin more than to the colour of the short hairs covering same),

the pale hands and feet, and thick, soft fur readily distinguish

these mice from the members of the allied genus Ypimys.

48, LopHuRomys ZENA Dollm.

OF OAS Maw. HieAs

49, Acomys 1enitus Dollm.

©. 96,98. Narossura River.

¢. 108. Southern Guaso Nyiro.

50. Acomys ABLUTUS Dollm.

g. 164. Near the Ngamatak Hills, Turkwel River.

51. ARVICANTHIS STRIATUS MASSAICUS Pagenst.

@. 51; @. 47, 52,-53, 54. Amala River.

52. ARVICANTHIS ABYSSINICUS NAIROB& Allen.

6. 94; 2. 92. Lengototo, B.E.A.

These two specimens appear very similar in colour to the

Nairobi Arvicanthis.

316 MR. GUY DOLLMAN ON

53. ARVICANTHIS ABYSSINICUS RUBESCENS Wrought.

6. 221, 222. Patong, Uganda.

sg. 190,191; @. 189. Bakoro, Uganda.

The type locality of Arvicanthis a. rubescens is Kibero, Unyoro ;

it appears to be distributed over a very large area, the British

Museum possessing specimens from localities as far apart as

Nimule and Ruwenzori.

54, ARVICANTHIS ABYSSINICUS PRECEPS Wrought.

3g. 130. Baringo.

This Arvicanthis, described from Naivasha, has already been

recorded from Baringo, Mr. Kemp having obtained a large series

of specimens both at Baringo and on the Laikipia Plateau.

55. ARVICANTHIS TESTICULARIS JEBELZ Heller.

g. 200, 213. Kamchuru, Lobor.

Central Province, Uganda.

56. ARVICANTHIS RUMRUTI PALLESCENS Dollm.

Abstract P. Z.S. 1914, p. 25. (April 14.)

6. 20, 23, 24; 2, 21,-22. - Loita Plains.

6.115; @. 117. Southern Guaso Nyiro.

Allied to Arvicanthis rumruti Dollm. ; larger in size and much

paler in colour.

General proportions rather greater than in rwmruti; hind foot

25-28 mm. in length.

Colour of dorsal surface pale olive-grey, lined with brownish

black and washed over with a light yellowish tint. Flanks

dirty white washed with pale yellow, gradually passing ito the

white ventral surface, without any marked junction of the two

areas. Ears clothed with short yellow hairs, almost as brightly

coloured and conspicuous as in the Somali species.

Backs of hands and feet yellowish. Ventral surface of body

white ; hairs longer than in rwmruti, and the general effect very

much whiter, owing to the dark basal portions of the hairs being

hidden by the long white tips. Tail much as in rwmruti, but

rather paler throughout. :

Skull larger than that of the Laikipia form, with larger molars

and more inflated auditory bullee.

Dimensions of the type (measured in the flesh) :—

Head and body 133 mm.; tail 109; hind foot 26; ear 17.

Skull: greatest length 33 mm. ; basilar length 27-7; condylo-

incisive length 31:5; zygomatic breadth 17:8; interorbital

constriction 5; length of nasals 12:7; palatilar length 15 ;

length of palatal foramina 7-4; alveolar length of upper molar

series 675.

Hab. Uoita Plains, B.H.A. Altitude 6300 feet.

Type. Old male. B.M. No. 13.10.18.142. Original number

®§. Collected on October 4th, 1912.

BAST AFRICAN MAMMALS, SIZ

‘The affinities of this Arvicanthis are a little difficult to decide ;

it seems on the whole to be more nearly related to the rwmrwti

species than its near neighbour 4. abyssimicus nairobe. The

Nairobi form is at once distinguished by its larger size and far

richer and darker coloration.

57. PELOMYS FALLAX IRIDESCENS Heller.

65 D2 Os la” Aanailley lsrovere

Heller, in his description of iridescens, points out that it is

distinguished from fallax in possessing a heavier dorsal stripe.

In the male specimen, now under consideration, the dorsal

stripe is well marked ; in the female, No. 71, the stripe is absent

altogether. The type locality of iridescens is Mt. Mbololo, Taita

Mountains, Seyidie Province.

58. TACHYORYCTES NAIVASH# Thos.

Cre eek Sie OF 22 ee Naivasha:

@ yg. 27. Lemek Valley.

59, TACHYORYCTES RUDDI BADIUS Thos.

Go Ws Qe Whe Wilkwi, Issa,

60. HysTRix AFRICH-AUSTRALIS Pet.

Q. 64. Amala River.

61. Lepus vicrort# Thos.

6.1. Mt. Suswa, B.E.A.

©. 41. Amala River.

6. 197. Nakwai Hills, N. of Lake Kirkpatrick, Central

Province, Uganda.

3. 173. Kozibiri River, Uganda.

62. OREOTRAGUS OREOTRAGUS SCHILLINGSI Neum.

©. 230. Narossura River, B.E.A.

3g. 187. Mt. Maroto, Uganda.

In Oreotragus o. schillingst both sexes bear well-developed

horns, a character which readily distinguishes this form from

the closely allied Oreotragus o. aureus Heller; in the female

specimen from the Narossura River the horns are quite as large

as those of the male from Mt. Maroto.

63. OUREBIA OUREBIA COTTONI Thos. et Wrought.

3.193; 9.194. Bakora Plains, Lobor, Uganda.

©. 65. Amala River, B.H.A.

3.77. West of Leganisho, B.H.A.

@. 128. Baringo.

64. RAPHICERUS NEUMANNI Matsch.

3. 93. Lengototo, B.H.A.

65. RHYNCHOTRAGUS GAVENDISHI Thos.

g. 28. Lemek Valley, B.H.A.

318 ON EAST AFRICAN MAMMALS.

66. RauyncoHorracus nasogurratus Lénnberg.

g. 146. 12 miles N. of the Kerio River, B.E.A.

©. 129. Baringo.

3. 147. Wei Wei River, Rift Valley.

Q. 195. H. of Mt. Lobor, Uganda.

67. CERVICAPRA CHANLERI Rothschild.

d. 95. Narossura River, B.E.A.

68. GAZELLA GRANTT RAINEYI Heller.

dg. 196. Bakora Plains, Lobor, Uganda.

3. 136. 30 miles N.W. of Baringo.

69. CEPHALOPHUS GRIMMIA LUTEA Dollm.

Abstract P. Z.S. 1914, p. 26. (April 14.)

3. 178; 9. 180. Mt. Maroto, Maroto Stream, Uganda.

S. 226. Falabek, east of Nimule.

Allied to Cephalophus g. abyssinicus Thos., distinguished by

its far paler colour and larger teeth *

Colour of dorsal surface pale greyish buff, the buff tint most

dominant on the neck, shoulders, and flanks, becoming greyer on

the hind quarters and back; neck and shoulders “ light pinkish

cinnamon ” (Ridgway, 1912) mixed with the dark brown tint of

the hair-bases; hind quarters ‘‘smoke-grey ” speckled with dark

brown. Face and head markings as in the other members of the

group. Ventral surface of the body much as in the allied forms.

Dimensions of the type (uneesimired in the flesh) :—

Head and body 870 mm. ; tail 705 hind foot 225; ear 101.

Skull: greatest length 169 mm. - basal length 150; zygomatic

breadth 77:5 ; oreatest width across orbital region 79: 8; length

of nasals 57: 8 ; greatest breadth across eens 31; palatal

length 84; length of upper cheek-teeth 51-5.

Hab. Mt. Maroto, Maroto Stream, N.E. Karamojo, Central

Province, Uganda. Altitude 3703 feet.

Type. Old female. B.M. No. 13.10.18.164. Original number

180. Collected on January 25th, 1913.

The type is the only fully adult specimen collected; of the

others No. 178, a subadult male, possesses horns lke those of

abyssinicus. This Uganda race is easily distinguished from the

allied forms by the pale greyish-buff colour of the dorsal surface ;

the Duikers most nearly allied are the Abyssinian C. g. aby sstniens

and C. g. nyanse Neum., from the Guaso Negishu District. In

general colour this Uganda Duiker is surpr isingly like the South

‘Afri ican grimmia, the buff on the shoulders and flanks being

rather more dlonmimert in this new race. Cephalophus gq. hinded,

described by Wroughton from specimens collected at Fort

Hall, appears as a bright orange-buff coloured animal when

compared with this Uganda Duiker.

* Tn the Abstract of this paper the teeth were erroneously stated to be “smaller ”

than those of ©. g. abyssinicus.

ON THE LATERAL MUSCLE IN TELEOSTEI. 319

2. On the Nature of the Lateral Muscle in Teleostei. By

Epwarp W. SHANN, B.Sc., Assistant to the Professor

of Natural History and Lecturer in Comparative

Hmbryology in the University of St. Andrews *.

[Received February 2, 1914: Read April 7, 1914. ]

(Text-figures 1-3.)

INDEX. Page

Til EORUSVTROYS LEV GTH(O 7: Sea wet ace aay eck Bander once ance o RUN E ba NeR SMA SIIC)

QEOENISGOuICaI a eters ees ree area we tn nee 320

3.) The Author's Observations) 2.0......0-0-0ecssesse. 328)

(2) Extexnal Conformation 0.2... | 328

(2) PintcernaliS tractureseeae eee eee

Asc ULNA Al Vilas deba aaace eu Teco DN essay SLU Det Sak 336

OS MAGS ALUNey eer edema cae chute gnee dese Web e Ma RN aes BOT

1. IntRopDUCTORY.

During the past two years my research hours have been largely

devoted to a study of the myology of the pectoral girdle and fin

of fishes, and recently the Teleosteans have more particularly

occupied my attention. As the musculature of the limbs is

entirely derived from the so-called lateral muscle, it is desirable,

in the first instance, accurately to comprehend the nature of the

lateral muscle.

That the primitive composition of the lateral muscle, seen only

in the embryonic condition of living fishes, becomes highly

modified in the adult Teleostean is beyond doubt; but it is

ditticult to reconcile the views of various authors, some of whom

interpret it in terms of two superimposed layers and others in

terms of one layer. It would be obviously unwise to advance to

a description of the myology of the limbs while this fundamental

question is left undecided. ‘To uphold the single-layer theory of

thé lateral muscle in Teleostei is, then, the primary object of the

present paper.

The complex series of cranial muscles is also derived from the

lateral muscle, but the secondary modifications which they have

undergone render highly problematical any conclusions based on

their arrangement. For this reason the cranial muscles will only

receive passing consideration, The brunt of the discussion

will be focussed upon that portion of the lateral muscle which

lies behind the pectoral girdle; and, in particular, upon the

caudal region, which is generally accepted as the least modified.

Though not a primary part of my research, the conclusions

advanced below are the outcome of it; and they are offered as a

preface to a section of an original morphological study.

I wish to record my gratitude to the Carnegie Trust, whose

* Communicated by Prof. W. C. McInrosu, F.R.S., C.M.Z.S.

320 MR. E. W. SHANN ON THE

grant in aid of research provided me with an image-erecting

binocular microscope, invaluable in tracing the intricate courses

of muscle-fibres, giving comprehensive stereoscopic views of

the muscle such as are unobtainable by simpler forms of optical

apparatus.

HIsToRICAL.

Owen (8) emphasizes the fact that the lateral muscle is essen-

tially an aggregate structure formed of a series of transverse

muscles (myomeres). The divisions of the lateral muscle which

he cites are based on the directions of the myocommata (‘ ten-

dinous insertions”); but he does not regard these as having any

true morphological value. The carinales, however, are regarded

as entities. I can find nothing in Owen’s writing to show that

he regarded the lateral musele | as divisible into a layers.

Humphr y (4) recognises a true division of the lateral muscle,

by a septum passing inwards beneath the lateral line, into a

dorsal and a ventral moiety. His subsequent divisions of these

moieties are based on the directions of the myocommata; at the

same time he states that when these divisions are traced forwards

to their insertions they become completely severed one from the

other, The red fibres occur superficially on either side of the

lateral line. The latero-ventral portion of the muscle (see table)

is divided by the direction of its component fibres into a super-

ficial layer (obiquus externus) and a deep layer (ob. internus),

but there is no septum between these two layers. In this portion

a third layer was noted in certain Teleosts (Bream, Dace), which,

from its position beneath the ribs, was regarded as a possible

homologue of the transversalis layer of Amphibia. Similar

changes in the direction of the fibres enabled Humphry to dis-

tinguish three areas of the mesio-ventral portion, two superficial

and one deep, but these again were not defined by fascie.

Gegenbaur (8) also recognises a horizontal plane of division at

the lateral line whereby a dorsal is separated from a ventral

moiety. He proceeds to divide these moieties in terms of cones,

complete and incomplete, which are revealed in sections of fishes.

It is desirable here to draw attention to a discrepancy which is.

displayed in the illustration fig. 276, A, which represents the

time-honoured caudal section of a Mackerel, originally drawn

by J. Miller, of which Owen (amongst others) has made use.

Gegenbaur’s "description of this section is perfectly lucid, and

more precise than Owen’s; but, unfortunately, he has added

a diagram of the superficial arrangement of the myocommata

(fig. 576, B), wherein, if we may judge by the lettering, he

represents the upper cones (a) as identical with the cones seen

in section in fig. 276, Aa. In reality the superficial cones are

directed posteriorly and their apices represent the angles which

the uppermost superficial fibres of the true cones make with

fibres of the incomplete cones (which are directed downwards

and backwards), whereas the true cones are directed anteriorly

LATERAL MUSCLE IN TELEOSTEI. 321

and, being deeply seated, are not visible superficially. Moreover,

the apices of the superficial cones are of necessity situated in a

higher horizontal plane than those of the true cones.

The five longitudinal portions of the lateral muscle which

MecMurrich (7) derived from his investigation of Amiwrus are

equivalent to various areas described by previous authors. This

is indicated by the positions which they take in the above table.

‘he author implies that the classification is only of empirical

value. His divisions are based upon the superficial appearance

of the muscles, and upon the extent of their origins. He does

not recognise a clear division beneath the lateral line, nor is

there any suggestion of stratification in any portion of the lateral

muscle.

Maurer (6) divides the lateral muscle into two portions, a

dorsal and a ventral, separated by the lateral line. The dorsal

portion is outside the sphere of his investigation, the ventral he

proceeds to spht into layers.

If the red fibres * immediately ventral to the lateral line be

removed, a stout muscular layer is observed whose fibres run

obliquely, from forwards and upwards to backwards and down-

wards (7. e. in theSame direction as those of the obliquus externus

of Amphibia). The outermost layer of this muscle runs from

septum to septum, the inner from rib to rib; since, however, the

direction of the fibres does not change, the two layers are regarded

as comprising a single muscle, the obliquus externus 7.

Ventrally the origins of the fibres of this muscle change so as

to resemble those of the obliquus internus of Urodeles t, while

the fibres which meet in the mid-ventral line their fellows of the

opposite side go to form a rectus.

Returning to the area of the ventral muscle immediately below

the lateral line, where the fibres take the direction of the obliquus

externus of Amphibia, if the superficial fibres be removed a deep

layer is found whose fibres slant in the opposite direction (Ge. @s

from forwards and downwards to backwards and upwards, like

those of the obliquus internus of Amphibia), These fibres form

a thin deep stratum running from rib to rib§. There is no

mention, however, of a fascia separating the deep from the super-

ficial stratum.

Beneath the above layers is yet a third, in Chondrostoma, at

least ; this is spoken of as a thick one covering the peritoneum jj.

The rectus is not, he says, developed as a costal muscle, rather

as a ventral part of the primary belly-musculature 4.

* The occurrence of these red fibres is more fully described on p. 326.

+ The equivalent of Humphry’s latero-ventral (surface) portion.

+ This and the superficial layer of the next form the “‘ pectoralis ” of Humphry.

§ Equivalent in all respects to Humphry’s “ obliquus internus.”

|| This is apparently equivalent to the transversus stratum which Humphry

observed in the Bream and Dace. Nevertheless, Maurer states that the transversus

properly so called is absent in all fishes.

4 From which I gather that this muscle is regarded as being continuous dorsally

both with the obliquus externus and with the obliquus internus. It thus included

the “rectus ”’ and ventral portion of “‘ pectoralis” of Humpliry.

322 MR. E. W. SHANN ON THE

Here we have an exposition of the two-layer theory of the

lateral muscle of Teleostei, differing in few points from that

advanced by Humphry.

Wiedersheim (9) describes the completeness of the division of

the dorsal and lateral moieties of the lateral muscle by a con-

nective tissue septum extending from the axial skeleton to the

integument at the region of the lateral line. Only one layer of

muscle is recognised.

With Knauer (5) we are brought into contact with a new view

of the lateral muscle. It is divided longitudinally into two main

portions which are defined according as they arise from (@) the

head (Riickenmuskulatur) or (6) the neck (Bauchmuskulatur).

The division at the lateral line, though complete, is only regarded

as being of secondary importance. The oblique line in the table

severing the back musculature from the belly musculature indi-

cates the author’s contention that the former gradually pushes

its way over the latter as we proceed in an antero-posterior

direction. Finally, in the posterior abdominal region the back

musculature comes completely to enclose the diminishing belly

musculature ; and in this area a true stratification of the body-

wall is realized. His lateral line muscle is syronymous beyond

doubt with the “ red fibres ” of other authors.

Text-figure 1.

Diagram to illustrate the apparent (ABCDE) and the real (ABHC’H’Dk)

construction of a myomere. (After R. Chevrel.)

Dietz (2) bases his division of a Teleostean myomere, and hence

of the lateral muscle, on the transverse septum extending beneath

the lateral line. His subsequent divisions are due to the direc-

tion of the myocommata. He proceeds to investigate myomeres

from various parts of the body, and shows how the pleuro-dorsal

portion gradually becomes emancipated from the pleuro-ventral

as we proceed antero-posteriorly; with this is correlated the

LATERAL MUSCLE IN TELEOSTEI. 323

change of a single half myomere, dorsal or ventral, from a single

bend to a double bend *.

Chevrel (1) has given a lucid description of the conical, or

pyramidical, structure of the myomeres forming the lateral

muscle. His diagram illustrative of the apparent and the real

structure of a single typical Teleostean myomere is reproduced

here (text-fig. 1). “At Bis seen the dorsal posterior pyramid, at H

the anterior dorsal pyramid, at H' and D the ventral pyramids,

while C indicates the lateral line. It is only necessary to add that

in the abdominal region the dorsal moiety (portion épiaxiale) alone

exhibits this double evagination ; the ventral (portion hypoaxiale)

is ribbon-like. Chevrel recognises a horizontal septum dividing

completely a dorsal from a ventral moiety; but there isno mention

of stratification in any area of the lateral muscle. The carinales

(m. gréles) are the homologues behind the pelvis of the rectus

which is found in the thoracic region.

Tue AUTHOR'S OBSERVATIONS.

Having now stated the principal views extant as to the

divisions of the lateral muscle in Teleostei, the conflicting nature

of these becomes patent. I shall next proceed to formulate a

classification of the parts of the lateral muscle, retaining such

features as are agreed upon by the majority of the investigators

whose work I have studied, and discarding those which I find to

be incapable of general application. In this classification no new

terms will be introduced, preference being accorded to those which

are deemed most descriptive. I wish it clearly to be understood

that I regard any such classification, in the light of our present

limited knowledge, as essentially empirical anne tentative.

That the so-called lateral muscle of fishes is, more strictly

speaking, a sequence of serially homologous transverse muscles

(myomeres) is the fundamental starting point upon which all

investigators are agreed. Setting this fact in the background,

but never letting it completely out of sight, let us proceed first

to map out the superficial divisions of this muscle in a Teleostean,

and then to prove by dissection the morphological value of these

divisions.

No specified type has been taken; but the description, and

herein lies what merit it possesses, is so generalised as to apply

to the majority of Teleostean fishes. Exceptional cases will be

cited wherever practicable.

(1) Haternal Conformation.

The most obvious longitudinal division giving rise to a dorsal

and a ventral moiety is formed by the passage of the lateral line.

Upon this point there is but one voice.

* This process can be very clearly followed in Knauer’s drawings of Salmon

sections, op. cit. pl. 11. figs. 15-19.

Proc. Zoou. Soc.—1914, No. XXIT. 22

MR. E. W. SHANN ON THE

Tabulated Summary of the

GEGEN- McMorricu

Owen (8). Houmpury (4). BAUR (8). Maurer (6).

Superficial. : Deep. Superficial. Deep.

I 1

1 1

Supra- 1 Ist Portion. !

carinalis. : Incomplete |) i

DorsaL ‘ 3 1 Cones 1

Dorsal Mesio-dorsal 1 (directed | | ;

Morery. | Section. Portion. backwards). | |

| .

I Entire r 2nd Portion. | Dera 1

Deal a I ntir { ortion. |

Middle Latero-dorsal 1 Cones | | I

Cae Portion. J (directed ;

: ' forwards). | | i

1 1

I ; 1

Wateral Red fibres. 1 Red fibres.

inves ahi Manistee 7 ra Leta ae US Ae ee ca Te om

Red fibres. ! Red fibres.

1 Entire 1

Ventral LLatero-ventral | Sui E laa Cones s a Obliquus 1 Obliquus

Middle Portion. 1 Sy (directed 3rd. Portion. externus. ! internus.

Venrrat| Section. | Obliquus externus. ‘- | forwards).

I 1

Mesio- ( Lat. dorsi; 1

MGI Merits ventral | Rectus. Foeorplete Diagonal fibres!

c nyty ra ly aya sy

Portion Pectoralis., (Gecied Ae RoPakiorn: Gee as

Iheal went) . internus).

1 Rec'tus.

Infra- ! :

carinalis. Carinales. ; 5th Portion. ;

With regard to the dorsal moiety, of the writers quoted above,

MeMurrich, Maurer, Wiedersheim, and Knauer see no reason for

a further subdivision.

The remainder, however, favour a division,

but are divided as to the nature of that division, Owen and

Humphry using the external appearance as a basis, while Gegen-

baur, Dietz, and Chevrel refer to it in terms of internal cones

(the difference will subsequently be shown to be of less degree

than at first sight appears). The myocommata of the dorsal

moiety in the trunk region are V-shaped, the apex of the V being

directed posteriorly. The inclination of the V varies from an

acute to an obtuse angle. Between two successive myocommata

the muscle-fibres have often been said to run parallel to the long

axis of the body. This is the case in the middle region, but, in

the arms of the V, the fibres gradually become more or less

inclined in the same direction as the arms; that is to say, if

produced, the fibres in the upper and lower arms would meet

behind the V. With Humphry, we will denote that portion of

the dorsal moiety which contains the upper backwardly directed

arms of the V as the mesio-dorsal portion, and that which

LATERAL MUSCLE IN TELEOSTEI.

Views of the Authors cited.

325

WIEDERSHEIM (9). KNAUER (5). DiETz (2). CHEVREL (1).

Superficial. | Deep

{

5) ( Posterior, external,

Im, Dorsal proper. or superficial

f I Pyramids.

| Straight Dorsal

| Sa eae

r Dorsal Portion. BS GFe ime ag

I Portion.

| | Back I Anterior, internal,

(Riicken) Pleuro-Dorsal. or deep Pyra-

| > Musculature. | mids.

B I

Lat. line muscle. Red fibres.

Se loaaepeoooeocead

| | Lat. line muscle. Red fibres.

Oblique I ( Anterior, internal,

) ( Analogue J Back-muscle. ieee Pleuro-Ventral. or deep Pyra-

| | of Obliquus. i mids.

Ventral eee Ventral

L ; 1 Oblique

Portion. | Analogue Belly ' Belly-muscles. Portion.

of Rectus. (Bauch) Posterior, external,

Musculature. | Ventral Proper. or superficial

(Recitus). Pyramids.

Straight Belly-muscles.

Muscles gréles.

contains the lower backwardly directed arms as the latero-dorsal

portion (see text-fig. 2, I7.D., L.D. p. 331). In the trunk region

there is no distinction between the mesio-dorsal and the latero-

dorsal portions except in the direction of their fibres, and, even

then, the differently directed fibres are connected by a transitional

area. ‘Tracing these portions forwards to the back of the head

the two are found gradually to be severed from one another ;

strong fasciz separate them; their transverse divisions begin

successively to fade and disappear; and, finally, they are inserted

on the skull, the mesio-dorsal portion on its roof (supraoccipital),

the latero-dorsal portion on its side (exoccipital and otic recess)

and on the post-temporal; the last named gives off a superficial

branch to the supraclavicle (Llennius pholis) or to the cleithrum

itself (Cottws scorpius, where it is attached to the posterior border

of the backwardly directed spine in which that bone ends),

In the little gobiiform fish, Periophthalmus, the mesio-dorsal

portion is attached to the upper border of the post-temporal as

well as to the supraoccipital ; but this so far as my observations

go is an exceptional occurrence.

22*

326 MR. E. W. SHANN ON THE

In the majority of Teleosteans a small muscle is cut off from

the latero-dorsal portion immediately above the lateral line.

From the distinctive coloration which it frequently presents it

has become known as the ‘‘red muscle” or “red fibres.” It is

not merely differentiated from the latero-dorsal portion by the

colour of its fibres, or even by their direction, which is parallel

to the long axis of the body (whereas those of the main muscle

are directed slightly upwards and backwards); but a distinct

fascia separates it in each myomere (see text-fig. 3, rf. p. 334).

A similar “ red muscle” is cut off from the latero-ventral portion

immediately below the lateral line. The “red muscle” is

mentioned by Chevrel; we have seen that it was also recognised

by Maurer. Chevrel states in another part of his text that this

muscle is constant neither in presence, nor in form, nor in tint,

nor even in its relations with the lateral nerve. Further, in

Humphry’s paper (op. cit. p. 294) the following words in reference

to fishes appear in a footnote :—“..... and the lateral furrows

are commonly occupied by muscular fibres which bear the trans-

verse septa, but which are more closely connected with the skin,

and peel off with it more easily than the rest of the lateral muscle.

These fibres are more vascular than ordinary muscular fibres ;

and in a piece which I examined from a Dace they contained

more oil than the other muscles. Stannius (Handbuch der

Zootomie (2), 112) says that they, in addition, present micro-

scopically the appearance of tissue in process of conversion into

muscle. I did not find that to be the case. With the exception

of the excess of oil, they represented the usual microscopical

characters of striped muscle.” Such observations as I have

made go to support Humphry’s statements. The “red muscle”

ovadually disappears anteriorly, so that it never reaches the

shoulder-girdle.

taraine next to the ventral moiety, we find, in the trunk

region, an exact repetition of the condition that has been

described for the dorsal moiety. Following Humphry’s nomen-

clature, which we have used above, let us designate the area

immediately below the lateral line, comprising the backwardly

and downwardly directed arms of the V- shaped myocommata, as

the latero-ventral portion, and the area between this and the

mid-ventral line (or the infracarinales muscles, where such are

present) as the mesio-ventral portion (see text- fig. 2, Z.V. and

MV.). It will be apparent, on turing back to ‘the tabulated

summary of the views of the various authors, that this division

of the ventral moiety of the lateral muscle meets with more

general recognition than does a corresponding division of the

dorsal moiety. Tracing these portions forward to the shoulder-

girdle, we find that thet apices of the backwardly and downwardly

directed V- shaped myocommata rapidly approach the lateral line:

in other words, the latero-ventral portion rapidly disappears.

MeMurrich states that this portion (his “3rd portion”) dis-

appears in Amiurews before reaching the shoulder-girdle; such is

LATERAL MUSCLE IN TELEOSTEI. 327

the case in some other Teleosteans, but in the majority which I

have examined a small bundle of fibres, undoubtedly the con-

tinuations of the latero-ventral portion, are attached to the upper

extremity of the cleithrum. At its point of attachment and for

a short distance behind, the latero-ventral portion is actually

severed from the mesio-ventral; though, further back, the two

are continuous, just as are the mesio-dorsal and latero- dorsal

portions. With the decrease in width of the latero-ventral

portion, the mesio-ventral portion increases so that it becomes

attached to the whole length of the cleithrum, except at the

dorsal extremity of that bone, and to the coracoid in part. There

exists, however, a triangular area, composed of connective-tissue

traversed by a few muscle fibres, beneath the pectoral fin ; this

triangle has its base on the cleithrum, consequently there appear

to be two main insertions of the mesio-ventral portion on the

cleithrum.

On the ventral surface in the anterior region the fibres of the

mesio-ventral surface meet their fellows of the opposite side ;

the fibres are here arranged parallel to the long axis of the body,

which has led some authors to regard them as homologous

(Humphry and Maurer), or, at least, analogous (Wiedersheim)

to the rectus abdominis of the higher vertebrates. In Teleosteans

whose pelvic fins are thoracic or jugular in position the rami-

fications of this portion of the lateral muscle show remarkable

variety. To describe these ramifications in detail would only

serve to confuse the issue of the investigation in hand; but

it is worthy of note that, m every example which I have had

occasion to examine, a bundle of fibres from the mesio-ventral

portion of the lateral muscle runs forward without interruption

to be inserted on the hypohyal.

Knauer (5) figures the lateral muscle of the Gurnard im the

area just behind the shoulder-girdle (Pl. ui. fig. 20). In this

illustration the mesio-ventral portion (Bauchmuskel) is depicted

as gradually disappearing towards its posterior extremity, and in

the text he says that in the anal region this muscle, with its

straight-running fibres, is reduced to a small strip on either side ;

further, that these strips fuse with the latero-ventral portions

(Riickenmuskulatur) behind the anus. This is not indicated in

the figure, where the mesio-ventral portion appears to die out

in the anal region; Jrigla, however, with its flattened ventral

surface is not, perhaps (with deference to the author), the best

type of Teleostean fish for representing in side view the entire

extent of this portion of the lateral muscle. A little distance

behind the anus the mesio-ventral portion entirely disappears,

and the latero-ventral portions (schrigen Riickenmuskel) of

either side are separated by the anal fin. I had also used

Gurnards* in my investigation of the lateral muscle. Since

reading Knauer’s paper, I have examined them again most

* Trigla gurnardus, whereas Knauer’s specimen was 7’, hirundo.

328 MR. E. W. SHANN ON THE

carefully, and with the result that I still maintain that the

mesio-ventral portion is continued to the extremity of the tail

in that species just as has been described above for Teleosteans

in general.

There remain yet to be considered the small cylindrical muscles

known as carinales. These may be absent, and, when present,

exhibit considerable variety in their extent. The upper pair,

supracarinales, run parallel to one another along the mid-dorsal

line, separating the mesio-dorsal portions of either side of the

main lateral muscle-masses. They take origin on the occiput in

the Cod (Owen), whence they run to the first dorsal, and reappear

again in the interspaces between the dorsal fins; in Periophthal-

mus they arise slightly behind the skull from the neural spines,

and traverse the interspaces between the dorsal fins ; in Amiwrus

they only appear behind the first dorsal fin, and MeMurrich

describes them as “‘formed by the union of slips arising by tendons

from the spinous processes.” They are separated by strong fasciz

from the main masses of the lateral muscles, but retain their

original metameric segmentation. Functionally they act, accord-

ing to their position, as elevators or depressors of the dorsal

fin-rays. They may be regarded as specialised modifications of

the lateral muscle, just as are the other fin-muscles.

The lower pair, infracarinales, are similar in nature to the

foregoing muscies, and, when present, divide the mesio-ventral

portions of either side of the lateral muscle. Both Owen and

MeMurrich, who have described them, agree that they may be

divided into an anterior and a posterior section. The latter

extends from the posterior end of the anal fin to the base of the

caudal, and between the two anal fins where such occur. The

former extends from the anterior end of the anal fin to be~

inserted on the posterior face of the pelvic bone (ischium, Owen) ;

hence they describe it as the “ retractor ischii,” and Owen goes

so far as to describe in the Perch the “protractor ischii” as a

still further anterior prolongation of the same muscle.

A difficulty now arises. In many Teleosteans, especially in

forms whose pelvic girdles are situated far forward, the “ retractor

ischii” a short distance behind its origin on the posterior aspect

of the pelvic bone becomes inseparably fused with the mesio-

ventral portion of the lateral muscle ; or, in other words, part of

the mesio-ventral portion forms in many cases the “ retractor

ischii.” It would seem that the latter condition is more primitive

than that where the “retractor ischii ” is composed of a carinalis

muscle, which has been noted as a secondary modification of the

lateral muscle. Yet the ‘“vetractor ischii” composed of an

infracarinalis occurs for the most part in Teleosteans with

abdominal pelvic fins, that is to say, in the more primitive

forms; and vice versa. the (presumably) more primitive condition

of the ‘‘ retractor ischii” occurs in the more specialised 'Teleosteans.

This interesting anomaly is beyond the scope of the present

investigation ; it recalls frequent analogous occurrences in the

LATERAL MUSCLE IN TELEOSTEI. 329

evolution of animals, and is merely noted here as a bye-way

which might repay exploration.

Summary of the external divisions of the lateral muscle :—

. Supracarinalis.

. Dorsal moiety.

(a) Mesio-dorsal portion.

(6) Latero-dorsal portion.

3. “ Red Muscle.”

(Lateral Line.)

*“ Red Muscle.”

. Ventral moiety.

(a) Latero-ventral portion.

(6) Mesio-ventral portion.

6. Infracarinalis.

Js) |

(2) Internal Structure.

In the foregoing section we have named the outstanding

superficial divisions of the lateral muscle in Teleosteans. Let us

now proceed to prove by dissection to what extent these external

markings are evidence of internal structure. Enough has already

been said of the carinales and of the “red muscles ” to show that

these are specialised offshoots from the lateral muscle ; and, since

all the authors who have mentioned them are agreed as to their

morphological value, it would be superfluous to treat of them at

greater length in the present paper ; suffice it to say that there

is no question of their division into more than one stratum,

nor are they themselves regarded in the light of strata, deep or

superficial, of the lateral muscle. Let us concentrate our attention

upon the dorsal and ventral moieties of the main mass of the

lateral muscle.

With the exception of Owen and McMurrich, all the authors

whom I have quoted maintain the completeness of the division at

the lateral line. Dissection shows a well-marked septum running

continuously from beneath the lateral nerve to the bodies of the

vertebre. The lateral line, then, must be regarded as the

external evidence of a true horizontal division of the lateral

muscle.

Having established this fundamental division, let us enquire

to what extent the dorsal moiety may legitimately be subdivided,

It has been shown that it comprises superficially, («) a mesio-

dorsal portion, in which the fibres of each myomere are directed

from above anteriorly to below posteriorly, and (6) a latero-dorsal

portion in which the fibres take the opposite direction. On

stripping the superficial fibres from the latero-dorsal portion of a

typical myomere such as occurs in the caudal region, it is found

that the direction of the underlying fibres gradually changes so

as to become, first, parallel with the long axis of the body, and

then, very near to the vertebral column, from above anteriorly

330 MR. E. W. SHANN ON THE

to below posteriorly (like those of the superficial area of the

mesio-dorsal portion).

It is this change in direction of the fibres which led Humphry

and Maurer to distinguish two layers in the ventral moiety of

the lateral muscle. If Chevrel’s account of the internal structure

of a myomere has been followed, the reason for this change in

direction of the fibres will be perfectly clear. Since the ventral

half of the dorsal moiety of the myomere takes the form of a

pyramid with its apex directed anteriorly, of the fibres running

from the apex the uppermost ones may be expected to take a

direction from below anteriorly to above posteriorly, and the

lower ones from above anteriorly to below posteriorly ; and this

is indeed the case. Moreover, since the apex is directed inwards

and downwards, only the outer upper faces of the pyramids on

which the fibres are directed from below anteriorly to above

posteriorly, is visible externally, and has been designated the

“latero-dorsal portion” in our superficial examination. The

lower face, on the other hand, in which the fibres are directed

from above anteriorly to below posteriorly, is hidden beneath the

surface.

Similarly, the portion which has been designated “ mesio-

dorsal” in our superficial examination represents the outer face

of a backwardly directed pyramid.

It has already been mentioned that, just behind the head, near

the insertions of the mesio-dorsal and latero-dorsal portions of

the lateral muscle, these two masses are divided. The division

is not merely superficial, but extends usually throughout their

depth ; though in some forms (Cotius scorpius) a certain number

of strands from their deeper contiguous faces serve to connect

them almost up to the point of their insertions.

The above description of the arrangement of the dorsal moiety

of a myomere of the caudal region applies equally well to the

ventral moiety in the same region, only the direction of the

apices of the pyramids is reversed, the dorsal one being directed

anteriorly, the ventral posteriorly. Here, again, only the outer

face, but in this case the lower, of the anteriorly directed

pyramid, with its fibres running from above anteriorly to below

posteriorly, is visible externally, where it is recognised as the

“Jatero-ventral portion” of our superficial examination; while

beneath it lies the upper face, in which the fibres run in the

opposite direction, namely, from below anteriorly to above

posteriorly. Hence the explanation of the description in this

portion of the lateral muscle of two superimposed strata, equi-

valent respectively to the obliquus externus and obliquus internus

of higher vertebrates, advanced by Humphry and Maurer.

In the abdominal region, where the walls of the ventral moiety

are of necessity thin, the conical structure of the myomere

is lost. The latero-ventral portion throughout its depth con-

tains fibres which are directed from above anteriorly to below

posteriorly, at least in its most anterior area. It acquires its

LATERAL MUSCLE IN TELEOSTEI. 331

characteristic conical structure, however, considerably further

forward than does the mesio-ventral portion. The latter contains

fibres which are directed from below anteriorly to above posteriorly

on the lower parts of the flanks ; but, as noticed above, the fibres

of the ventral surface change in direction the nearer they

approach to the mid-ventral line so as to be disposed parallel to

the long axis of the body. I have failed in every instance to

find a septum, such as is described by Knauer, dividing the

mesio-ventral portion into two superimposed layers.

It has been stated above that a strand of the mesio-ventral

portion is prolonged anteriorly to be inserted on the hypo-hyal.

This strand, together with the hyo-cleithrafe *, constitutes the

neck musculature. Knauer (5) recognised these parts of the neck

musculature, and adds the following note (op. cit. p. 11):—

“Die Angabe Schneiders, dass sich der Sternohyoideus * aus

zwei der Linge nach zusammengewachsenen Muskeln (Hyo-

dorsalis und Hyoventralis) zusammensetzt, die als Fortsetzung

des Riickenmuskels und des Rectus zu betrachten sind, ist dadurch

zu verstehen, dass Schneider offenbar schon die oben erwihnte

Fortsetzung der inneren Schichte (von ihm als Riickenmuskulatur

aufgefasst) in die Halsmuskulatur bemerkte.”

Text-figure 2.

Transverse section through the caudal region of a typical Teleostean with the

contiguous. body-surface, to show the relations to one another of six con-

secutive myomeres. The right side of the figure represents the anterior

portion of the section. (Generalised, mainly from Cottus scompius:)

M.D., Mesio-dorsal portion; 1Z.V., Mesio-ventral portion ; Z.D., Latero-

dorsal portion; Z.V., Latero-ventral portion; 1-6, consecutive myomeres,

the first is anterior; f.s., horizontal septum ; /./., lateral line (with “red

fibres”? on either side of it); @ftm., anal fin-muscle; d.fm., dorsal fin-

muscle.

T, too, have noted this double condition of the hyo-cleithrale in

certain Teleosteans (Lophius, Trigla, Zeus). My observations

* A yevised term more accurately descriptive of the muscle hitherto known as

hyo-clavicularis (synon. Stenohyoideus, Cervicalis profundus, etc.).

aon MR. E. W. SHANN ON THE

tend to indicate that the division line is horizontal rather than

tangential; that is to say, they support Schneider’s view.

Chevrel’s diagram of the pyramidical disposition of a caudal

myomere has been reproduced on p. 322 (text-fig. 1); the

accompanying illustration (text-fig. 2) will serve to convey a more

realistic view of the manner in which the pyramids fit one into

the other. Let us follow in turn the course of six myomeres

represented in the text-figure.

First myomere. —This is seen at the surface on the right of the

text-figure(1). In the mesio-dorsal portion (J/.D.) its fibres are

directed from above anteriorly to below posteriorly, in the

latero-dorsal portion (Z.D.) the fibres are directed from below

anteriorly to above posteriorly ; in the ventral moiety (Z.V. and

M.V.) this sequence is repeated. The dorsal moiety is separated

from the ventral by the lateral line (J./.) and by the “red fibres ”

on either side of it. Looking into the section, we find the first

myomere (1) in the form of an acute angled triangle with

its base abutting on the vertical septum, in which run dorsally

the neural spines and ventrally the hemal spines; it presents a

similar appearance both in the dorsal and ventral moieties.

Second myomere.—Seen at the surface immediately behind the

foregoing, it takes a similar course; the posteriorly directed

apices of its V-shaped bends are broken, however, by the section.

Following its course into the section we find that it presents a

V-shaped mass which bestrides the triangular section of the first

myomere.

Third, fourth, and fifth myomeres.—These are seen externally,

each at four points, as a series of gradually vanishing segments

of the V-shaped arms (the fifth myomere, however, does not

appear superficially in the central region). In section each

presents four quadrangular faces, two above and two below the

horizontal septum. Near the middle line the faces become

curved, so that they present a convex surface to the exterior

and a concave surface to the interior.

Siath myomere.—This does not appear superficially, but is seen

as a roughly triangular mass on either side of the horizontal

septum. The base of the dorsal triangular mass rests against

the body of the vertebra, that of the ventral one against the

hemal arch. The outer angles of these triangles, unlike those of

the sections of the first myomere, are rounded off, and fit into the

concave inner surfaces of the fifth myomere.

Having obtained a sequence of views of a series of consecutive

myomeres, let us proceed to reconstruct from them the conform-

ation of a single mvomere. As the ventral moiety of a myomere

is the counterpart of the dorsal, it will be sufficient to confine

our remarks to the latter.

It is evident from the foregoing observations that a myomere

is not visible superficially in its most anterior region, We have

seen it (6) as a triangular area in the angle between the centrum

and the horizontal septum. By dissection it is easy to demonstrate

LATERAL MUSCLE IN TELEOSTEI. 333

that this triangle represents the base of a pyramid which con-

verges to an apex on the centrum*. ‘Tracing the sixth myomere

backwards through the width of a myomere, we should find it to

show the appearance of No. 5 in the text-figure ; it has then been

thrust from contact with the vertebral column by the appearance

of a new pyramid: moreover, it has made its anterior superficial

appearance dorsally. Backwards again through the width of

another myomere it has the appearance of No. 4; here a consider-

able portion of it is visible externally, both dorsally and just

above the “‘red fibres”; in this and in its next stage, No. 3, it

‘appears in section as two quadrangular faces which are approaching

one another. In No. 2 we are able to see how the two faces,

whose superficial continuations are now recognisable as the mesio-

dorsal (J7.D.) and latero-dorsal (Z.D.) portions respectively,

become confluent. Passing from this point through the width

of another myomere the two faces are found fused into one

triangular area (1, in section), which is invisible externally owing

to the convergence of the faces of the myomere next in front.

By dissection it can be demonstrated that this triangle forms

the base of a pyramid whose apex is attached to the vertical

septum.

Thus it is seen that the dorsal moiety of a myomere is com-

posed of two opposed hollow pyramids, a dorsal one directed

inwards and backwards, and a ventral one directed inwards and

forwards. They are not divided by a septum but share a common

face, which may be seen in the text-figure at the areas marked

2, 3, 4, and, in part, 5 in the Z.D. portion of the section f+.

The ventral moiety of the myomere takes a course symmetrical

with the dorsal, which it resembles in all essential points of

structure.

Before leaving this figure it would be well to observe that it

serves admirably to indicate why there is but one change in

direction of the external fibres, correlated with the change which

the conical structure involves, in each moiety of the myomere,

instead of the two which one would be led to expect. Following

the central course of myomere No. 4, it is seen that only the

fibres above the mesial plane of the Z.D. portion, and below that

of the Z.V. portion, are visible at the surface. The fibres below

the mesial plane of the Z.D. portion and above that of the Z.V.

portion, which take the opposite directions respectively to those

named in the preceding sentence, are only found in myomeres 5

and 6; and these do not appear on the surface.

I have also followed the course of the myomeres in the dorsal

moiety of a young Salmon through over a hundred consecutive

sections from the pelvic region. Two such sections are repro-

duced here (text-fig. 3) to indicate the directions of growth and

diminution in seven consecutive myomeres. As corroborative

* This will be corroborated in the remarks on sections of a young Salmon, p. 334.

+ It is evident that this area may be regarded as the forward continuation of a

pyramid similar to 1, or as the backward continuation of a pyramid similar to 6.

334 MR. E. W. SHANN ON THE

evidence of the above observations the figures speak for them-

selves ; but the essential features which I wish to emphasize are

these :-—

First, that the horizontal septum (f.s.) running from the

lateral line to the vertebral centra is continuous ; also that the

“red fibres” (r,f.) remain constant in position and extent,

thereby showing that they form no part of the system of

pyramids or cones.

Text-figure 3.

Two transverse sections through the dorsal moiety of the Jateral muscle of a

young Salmon (34°5 mm.) taken from the pelvic region.

A represents a plane ‘064 mm. anterior to B; 1-7, series of myomeres, identical

in both sections; h.s., horizontal septum ; J.7., lateral line; 7.f., “red fibres”

~~, arbitrary internal division-plane between the mesio-dorsal and latero-

dorsal portions.

Secondly, that the numbers are not merely written down

symmetrically on either side of an arbitrary starting point.

Where any two parts of myomeres are numbered as the same

myomere, this has actually been proved to be the case by tracing

their passage through a sufficient number of sections until they

become confluent one with the other; to take an example, the

separate parts denoted by the number 5 in A, become confluent

when traced backwards to their position in B 5.

Thirdly, that the ventral, forwardly directed pyramid is seen

at its origin at A 1, and again, considerably increased in bulk,

when traced backwards to its position at B 1; conversely, that

the dorsal posteriorly-directed pyramid is seen decreasing from

A7 to BZ.

LATERAL MUSCLE IN TELEOSTET. 339

Fourthly, that at the outermost extremity of myomere B6 (at

the point where it is cut by the index line of B7) we find, very

nearly *, the apex of one of the external backwardly-directed

V-shaped myocommata. It has been pointed out that an

imaginary line joining such apices forms the division between

the mesio-dorsal and latero-dorsal portions of the surface area.

lt is now apparent that any deep continuation of this superficial

division must also be guided by expediency; for, internally to

our superficial guide-point, we find a <-shaped area (the section

of a hollow pyramid), which may be regarded in its entirety as

part of a dorsal, posteriorly-directed pyramid, and so of the mesio-

dorsal portion of the muscle; or, with equal justice, its lower

arm may be regarded as part of the latero-dorsal portion, from

the fact that its fibres follow the same direction as those in that

portion.

The question to be decided is this: granted that such a division

of the lateral muscle is in either case an arbitrary one, is it

expedient to frame it in terms of pyramidal masses or in terms

of direction of fibres ?

Before answering this question let us examine the fissure

which divides the mesio-dorsal and the latero-dorsal portions

near their anterior origins on the skull. It is found here that

all the fibres above the cleft throughout its depth run in the

same direction, namely, from above anteriorly to below posteriorly,

and that all those below it take the opposite direction. That is

to say, when they split apart, the two portions are not divided

according to their pyramidal structure, but according to the

direction of their fibres ; and this fact indicates that the division

of the dorsal moiety of the lateral muscle into a mesio-dorsal

longitudinal mass and a latero-dorsal longitudinal mass can most

naturally be framed in terms of the direction of its component

fibres.

In text-fig. 3, B it has been pointed out that the index-line to

the 7th myomere cuts the skin approximately at a point in the

superficial line of division of the mesio-dorsal and latero-dorsal

portions. From this point inwards a zigzag line has been drawn

to indicate the arbitrary internal division- plane between the

mesio-dorsal and latero-dorsal portions; above this plane the

fibres are inclined upwards and forwards, below it they are

inclined downwards and forwards. In A the superficial division

can be found by bisecting the line of skin between the two

portions of the myocomma which encloses the 6th myomere ;

from this point the internal division-plane can be traced as

min. 1B

It will now be realised that by somewhat devious paths we

have reached Gegenbaur’s position with regard to the structure

of the dorsal moiety of the lateral muscle (of. p. 320). Our

arbitrary division leaves us with a series of incomplete cones, or

* Tn the section next in front of this the apex of the sixth myomere is actually m

the surface.

336 MR. E. W. SHANN ON THE

rather pyramids, whose apices are directed backwards, and, below

them, a series of complete pyramids whose apices are directed

forwards.

The incomplete pyramids correspond externally with Humphry’s

mesio-dorsal portion, and the complete pyramids with his latero-

dorsal portion. For descriptive purposes this division of the

dorsal moiety (which is equally applicable to the ventral moiety)

has its uses. At the same time one cannot insist too strongly

on the fact that its morphological basis is only slightly more

secure than that on which Humphry and Maurer framed their

two-layer hypothesis ; both views depend upon the direction of

muscle-fibres, but the former derives certain additional support

from an investigation of the muscles in their most anterior

region.

The apices of the ventrally-directed pyramids do not. always

rest against the vertebral centra, but in some ‘Teleosteans

(Scomber, Trigla) they originate in the midst of the muscle-mass.

In this case the pyramids become converted into true cones, and

in section they appear as a series of concentric circles. In rarer

instances (Conger) the apices of the dorsal backwardly- directed

pyramids are also similarly situated, and in a section of the

dorsal moiety of the lateral muscle two superimposed series of

concentric circles appear.

SUMMARY.

1. The body-wall of a Teleostean fish is composed on either

side of a series of transverse muscles (myomeres) divided from

one another by fascie of connective-tissue (myocommata).

For practical purposes these may be regarded collectively as

forming a single lateral muscle.

2. The lateral muscle is composed of a single layer, which

varies considerably in thickness in different parts of the body.

3. In its primary condition, such as obtains in the caudal

region, it is divided into two symmetrical longitudinal moieties,

which are separated by a horizontal septum passing from beneath

the lateral line to the vertebral column.

4. The dorsal moiety is marked externally by a series of back-

wardly directed >-shaped bands. By joining the apices of these

>’s an imaginary line is formed dividing the dorsal moiety into

a (dorsal) mesio-dorsal portion and a ‘(ventr -al) latero-dorsal portion.

In the mesio-dorsal portion the muscle-fibres run from above

anteriorly to below posteriorly ; in the latero-dorsal ay they

take the opposite inclination.

Internally the mesio-dorsal portion is recognised as a series of

incomplete hollow pyramids, or cones, whose apices are directed

posteriorly ; while the latero-dorsal por tion is composed of a series

of complete hollow pyramids, or cones, whose apices are directed

forwards.

The ventral moiety is likewise divided into a (dorsal) latero-

LATERAL MUSCLE IN TELEOSTET. 337

ventral portion and a (ventral) mesio-ventral portion. The fibres

of the latter, at first oblique, as they approach the middle line

become straight, and resemble a rectus.

5. The above divisions of the dorsal and ventral moieties

suffice for descriptive purposes; but, pending further evidence,

they inust be regarded as essentially empirical in nature.

6. Small cylindrical longitudinal muscles are frequently cut off

from the mesio-dorsal portion, along the mid-dorsal line in the

interspaces of the dorsal fins. ‘These are the supracarinales of

Owens

Similar muscles are frequently cut off from the mesio-ventral

portion along the mid-ventral line. ‘These are the infracarinales

of Owen.

7. A small muscle, whose fibres are usually distinguished by

their red colour, is cut off from the latero-dorsal portion and

occupies the slope of the furrow in which the lateral nerve is

situated.

A corresponding muscle is cut off from the latero-ventral

portion immediately below the lateral line.

LITERATURE.

1. Curyret, R.—“ Essai sur la morphologie et la physiologie

du muscle latéral chez les Poissons osseux.” Arch. Zool.

Expér., Tome 52, fasc. 8, 1913, p. 478.

2. Dinrz, P. A.—‘Over den vorm der Myotomen bij de

Teleostei.” Tijdschr. Ned. Dierk. Vereen. ser. 2, vol. ii.

1910, Appendix, pp. xxv—xxvil.

3. GEGENBAUR, C.— Elements of Comparative Anatomy.

Translated by Mr. Jeffrey Bell and revised by Sir Ray

Lankester, 1878, p. 493.

4. Humpury, P.—‘‘ On the Disposition of Muscles in Vertebrate

Animals.” Journ. Anat. and Phys. vol. vi. 1871-72,

p. 293.

5. Knauer, K.—“ Die Bauchmuskulatur der Fische.” Arb.

Zool. Inst. Univ. Wien, vol. xvi, No. 3, 1909-10, p. 207,

3 plates.

6. Maurer, F.—“Der Aufbau u. die Entwickelung der ven-

tralen Rumpfmuskulatur bei den urodelen Amphib.

deren Beziehung zu den gleichen Musk. der Selach.

Teleost.” Morph. Jahrb. xvii. 1892.

7. McMurricu, J. P.—‘* Myology of Amiurus catus.” Proc.

Canad. Inst. vol. ii. No. 3, 1884, p. 60.

8. OwEN, R. — Comparative Anatomy and Physiology of

Vertebrates, vol. 1. 1886.

9. WiEDERSHEIM, R. — Comparative Anatomy of Vertebrates.

Adapted from the German by Prof. W. N. Parker. 3rd

English edition, 1907.

i eae ¥: fees ISI Re : °

PE 2 PENT secs ae path waite

(ees eae i Ariat ete rel a

rcAT pS edi Paaaicts

\ Hl: Bix ty AF NE RE ge

Mate coy ee hee NLRs #) tio peer

a ee eee hele aye

Fats iT elo eee ee es ty “UF OE RE. a6 ir: altel VER soil

a “ih a

EMD + Caw he SAKE Awe mm TW

: ap

ae y rea \ haat wit

ON SOME FISHES FROM NEW GUINEA, 339

23. Note on Aristeus goldiei Macleay, and on some other

Fishes from New Guinea. By C. Tare Reaan, M.A.

[Received March 17, 1914: Read April 21, 1914. |

(Text-figures 1 & 2.)

INDEX.

Page

Apisteus Goldiet 1.0... 6.2.0.0 v vee eee ener p eet tet enn ees 339

Nematocentris rubrostriatus ...... 20.00 cece ee 339

ETD GETE STURT S eo OE iter eu Gua aeie ea yea)

Mr. A. R. McCulloch, of the Australian Museum, has very

kindly sent me some notes on the types of Aristeus goldiei Macleay,

accompanied by some drawings, which are here reproduced.

These leave no room for doubt that this is the species that I have

recently described as Rhombosoma nove-guinee Rams. & Ogilb.

(Regan, Trans. Zool. Soc. xx. p. 983, 1914); as Macleay’s name

has precedence the species should be named Rhombosoma goldiet.

According to Mr. McCulloch the types, from the Goldie River,

New Guinea, are ten in number and measure 44 to 86 mm. in

total length. The dorsal rays number V-VI, I 13-14, the anal

rays I 21-24 and the scales in a longitudinal series 34 or 3D.

Text-figure I,

Rhombosoma goldiei : mouth, four times natiral size

The depth of the body varies from 34 in the length in the largest

specimen to 3? in the smallest. Allare marked alike and appear

to have been silvery, with the back darker, a broad dark band

from pectoral to caudal, and a black band from shoulder to eye.

Mr. McCulloch tells me that the types of Vematocentris nove-

guinee are mislaid and that the type of V. rubrostriatus Rams. &

Ogilb. is damaged about the mouth-parts ; a figure of the upper

Proc. Zoou. Soc.—1914, No. XXIII. 23

340 ON SOME FISHES FROM NEW GUINEA.

jaw is sufficient to indicate that the examples I have recently

described as belonging to this species are correctly determined.

Text-figure 2.

Rhombosoma goldiei: one ot the types, natural size.

Mr. Douglas Ogilby has written calling my attention to Dules

nitens (Rams. & Ogilb. Proc. Linn. Soc. N.S. Wales, xii. 1887,

p. 4) from New Guinea; there can be little doubt that this

is a synonym of Kuhlia humilis De Vis (Regan, Proc. Zool. Soe.

1913, p. 380), already known from Queensland and the Fiji

Islands.

P Z.$.1914. Stebbing. Pl.I.

Cambridge University Press.

T.R.R.Stebbing del.

TANAIS OHLINI, (Stebbing).

PZ S 1914 Steboing iil

as}

‘

VARS

TES

AR SOS

APNE NERS

T.RR.Stebbing, del. Cambridge University Press.

EXOSPHAFROMA CALCAREUS (Dana).

PZ.S.1914. Stebbing. Pl. I.

a“

ial

je ~ oe RCE TEN

Deen ato

Cambridge University Press.

T.R.R.Stebbing, del.

TRYPHOSITES CHEVREUXI, (Stebbing).

P Z.S 1914. Stebbing. PL. IV.

areal

epitome

x Mige

Cambridge University Press.

TR R Stebbing, del.

PARIPHIMEDIA NORMANI (Cunningham).

: ie mee

A ee or i peo eee

i oe - ee

Se S a ON

A= =

ie)

oe) es |

S aoe fen ge 7 e

SH edt 2 ” é

4 oP ~ iO oe

: Pe wee

: as ae SSeS

Ni eee

Ay ea Sree Sel SIs

eee Ss

Coen

eerie: Sea ore A 3

Cambridge University Press,

T.R.R.Stebbing , del.

PARIPHIMEDIA NORMANI (Cunningham).

PZ.S.1914. Stebbing. Pl VI.

T.R.R.Stebbing , del. Cambridge University Press.

MONOCULOPSIS VALLENTINI ,(Stebbing).

P Z.S.1914. Stebbing. PLVII.

3 ee

pet

ptt

= ~~

Deke Pilbests eee. alae aks Po o pe

} oh eee

Prp. 2.

ca ee nt aan goes

A ea = ao

Cambridge University Press.

T.R.R Stebbing ,del.

MONOCULOPSIS VALLENTINI,(Stebbing).

PZ. S.1914. Stebbing. PLVI.

Cambridge University Press.

T.R.R.Stebbing, del.

BOVALLIA REGIS, (Stebbing).

P Z.S.1914. Stebbing Pl.IX.

\ |

VAY

T.R.R.Stebbing ,del.

Cambridge University Press.

LEMBOS FUEGIENSIS (Dana).

ON GRUSTACEA FROM THE FALKLAND ISLANDS. 341

24. Crustacea from the Falkland Islands collected by

Mr. Rupert Vallentin, F.L.8.—Part I]. By the Rev.

Taomwas R. R. Stespine, M.A., F.R.S., F L.S., F.Z.8.

[Received February 24, 1914: Read April 21, 1914.]

(Plates I.-IX.*)

: INDEX.

Systematic :-— Page

kistiotspecies dealtawathy) re acne es eases) One

Dee GIS OUODCS 05 Win” vaderopeaeeodecapacser eee cos ace ues daeads pauses | 4este)

LCHUCATE DUG EXO Na 806, season sade nasdetase soo eNBetsEOs eoneoreescaseen OL!

Tryphosites chevrewxt, Sp. MW. 1.0.0 cece eet cec eee cette eee BOD

Monoculopsis vallentini, Sp. W. 00... 2.. cee ceeeee eee eee eee. 360

JEOCOUIE, FRAGT BBo Ws sea aqnonn saseoo esonewsadasnoasbausaanseasads | GOL

Paradexamine Manus, SP. De... .ecccc ccc eeeeee cee ceeeae recesses 366

Iphimedia normani transterred to Pariphimedia ......... 359

The record of which this is a continuation was published in the

Proceedings of the Zoological Society about fourteen years ago.

In the interval Mr. Vallentin has continued his researches during

more or less prolonged visits to the Falklands, with the result

that very extensive additions have been made to the series of

specimens left undescribed in my earlier report,

When Samuel Johnson, in 1771, published his entertaining

but politically-minded history of the Falkland Islands, there was

naturally no forecast in it that the restless “ barren ocean ” which

breaks on the shores of those wind-swept outposts of civilization

would eventually become a happy hunting-ground for students

of marine zoology. Nevertheless, as explained in my formev

paper, the nineteenth century found those waters fruitful in

interest. In the present century, while Mr. Vallentin has been

waiting with friendliest patience for my further account of his

unwearied and still unexhausted researches, the rush to the

Antarctic has incidentally brought the island fauna into renewed

prominence. As the following discussion will show, it has

engaged the attention of numerous eminent carcinolog’sts,

such as Chilton, Hansen, Ohlin, Ortmann, Thomas Scott,

Tattersall, and Thiele. The present paper proposes one new

generic name and five new species ?; but Mr. Vallentin’s collection

has made possible a reconsideration of various forms already

known by name, though very imperfectly known by nature, If

some useful light has been thrown upon these obscurities, it ma y

perhaps be welcomed as compensation for shortness in the list of

novelties, at an epoch when the discovery and display of new

species has been almost overwhelmingly rapid.

* Wor explanation of the Plates see p. 376.

+ (‘The complete account of the fiye new species described in this communicazion

appears here, but since the names and prelimjaary diagnoses were published in the

“Abstract” No. 182, 1914, these species are distinguished by the names bemg

underlined.—Ep1Tor. | nig Rapa RFR

Zee

342 THE REY. T. R. R. STEBBING ON

The former report dealt with Peltarion spinosulus White,

Halicarcinus planatus (Fabricius) *, Hurypodius latreillit Guérin,

Paralomis granulosus (Jacquinot), Hupagurus comptus (White),

Kuphausia vallentini Stebbing, Thysanoessa macrurus Sars, [ais

pubescens (Dana), Hxospheroma gigas (Leach), Cassidina emar-

ginatus Guérin-Méneville, Zrichoniscus magellanicus (Dana).

Of these the Hupagurus reappears under a different specific name,

and the species of Cassidina has in the interval suffered a generic

transfer. Of the specimens not included either in the past list or

the present, some are well known. They are reserved, along with

others of less obvious character, on the chance that detailed

examination, should time and opportunity permit, may yield

material for useful comment. In the meantime the following

identifications are offered.

Isopoda anomala

MALACOSTRACA.

or Apseudacea.

Brachyura. Fam. TANAIDA.

Tribe CyCLOMETOPA. Gen. Tanais Audouin & M.-Edwards.

Fam. ACANTHOCYCLIDA. Tanais ohlint Stebbing.

Gen. Acanthocyclus M.-Edw. & Lucas. |

Acanthocyclus albatrossis Rathbun. | Isopoda genuina.

Tribe FLABELLIFERA.

Tribe CATOMETOPA. | ifwa, Steeamaoniip a.

Fam. GRAPSID. Gen. Hxospheroma Stebbing.

Gen. Planes Bowdich. Hrospheroma calcareus (Dana).

Planes nuinutus Linn.) Gen. Cassidinopsis Hansen.

Cassidinopsis emarginatus (Guérin-

Méneville).

Vallentinia, gen. n.

Vallentinia darwinii (Cunningham).

Macrura anomala.

Tribe PAGURIDBEA.

Fam, PAGURIDZ.

Gen. Ewpagurus Brandt.

Hwpagurus forceps (Milne-Edwards). Wnlge WA Gy gine A

Fam. ASTACILLID&.

Tribe GALATHEIDEA. Gen. Astacilla Cordiner.

Fam. GALATHEIDA. Astacilla falclandicus Ohlin.

Gen. Munida Leach.

Munida gregarius (Fabricius).

Munida subrugosus (White).

Fam. IpoTEIDz.

Gen. EHdotia Guérin-Méneville.

Edotia tuberculatus Guérin-Méneville.

Macrura genuina. Gen. Macrochiridothea Ohlin.

Tribe CA RIDEA. Macrochiridothea stebbingi Ohlin.

Fam. HIppoLyTipa”. Tribe ASELLOTA,

Gen Nauticaris Bate. Fam. JANIRIDZ. -

Nauticaris magellanicus (A. Milne- | Gen. Notasellus Pfeffer.

Edwards). | Notasellus sarsii Pfeffer.

<lnne parentheses around the names of authors placed after scientific names in

this paper are used in accordance with Article 23 of the International Rules of

Nomenclature (Proc. 7th Int. Cong. Boston, 1907, p. 44 (1912)).—Eprror. |

CRUSTACEA FROM THE FALKLAND ISLANDS.

Amphipoda.

Tribe GAMMARIDEA.

Fam. LYSIANASSIDA. *

Gen. Tryphosites Sars.

Tryphosites chevreuat Stebbing.

Gen. Acontiostoma Stebbing.

Acontiostoma marionis Stebbing.

Fam. AMPELISCIDA.

Gen. Aimpelisca Kroyer.

Ampelisca macrocephalus Liljeborg.

Fam. PHOxXOCEPHALID®.

Gen. Pontharpinia Stebbing.

Pontharpinia rostratus (Dana).

Fam. MEeTorip2.

Gen. Metopella Sars.

Metopella ovatus (Stebbing).

Fam. ACANTHONOTOZOMATIDZ.

Gen. Iphimedia H. Rathke.

Iphimedia nodosus Dana.

Gen. Pariphimedia Chevreux.

Pariphimedia normani (Cunning-

ham).

Fam. (iDICEROTIDA.

Gen. Monoculopsis Sars.

Monoculopsis vallentimi Stebbing.

Fam. CALLIOPIID.

Gen. Halirages Boeck.

Halirages hualeyanus (Bate).

Fam. PONTOGENEIIDA.

Gen. Bovallia Pfeffer.

Bovallia regis Stebbing.

Gen. Pontogeneia Boeck.

Pontogeneia antarcticus Chevyreux.

Gen. Atyloides Stebbing.

Atyloides magellanicus (Stebbing).

Gen. Paramera Miers.

Paramera austrinus (Bate).

Fam, GAMMARIDA.

Gen. Melita Leach.

Melita inequistylis Dana.

Fam, DEXAMINID.

Gen. Paradexamine Stebbing.

Paradexamine nanus Stebbing.

343

Fam. TALITRIDA.

Gen. Talorchestia Dana.

Talorchestia scutigerwlws (Dana).

Gen. Hyalella Smith.

Hyalella patagonicus (Cunningham).

Fam. AORIDA.

Gen. Lembos Bate.

Lembos fuegiensis (Dana).

Fam. PHOTIDA.

Gen. Haplocheira Haswell.

Haplocheira barbimanus (Thomson).

Fam. AMPITHOIDA.

Gen. Ampithoe Leach.

Ampithoe brevipes (Dana).

Fam. JASsIDA.

8 Gen. Jassa Leach.

Jassa falcatus (Montagu).

Fam. COROPHIIDA.

Gen. Corophiwm Latreille.

Corophiwm cylindricus (Say).

Fam. PODOCERID.

Gen. Podocerws Leach.

Podocerus brasiliensis (Dana).

Tribe CYAMIDEA.

Fam. CAPRELLIDA.

Gen. Caprella Lamarck.

Caprella penantis Leach.

Tribe PHRONIMIDEA.

Fam. HYPERIID.

Gen. Hyperia Latreille.

Hyperia gaudichaudii Milne-

Edwards.

LEPTOSTRACA.

Fam. NEBALIIDA.

Gen. Nebalia Leach

Nebalia bipes (O. Fabricius)

THYROSTRACA

Fam. LEPADIDA.

Gen. Lepas Linn.

Lepas wustralis Darwin.

Fam BALANrIDsé,

Gen. Elminius Leach.

Elminius kingii Gray.

344 THE REY. T. R. R. STEBBING ON

MALACOSTRACA.

BRACHYURA.

Tribe CYCLOMETOPA.

Fam. ACANTHOCYCLIDZ.

1852. Cyclinea Dana, U.S. Expl. Exp. vol. xiii. p. 294.

1886. Cyclinea Miers, Rep. Voy. ‘ Challenger,’ vol. xvii. pt. 49,

p. 208.

1898. Acanthocyclide Rathbun, Pr. U.S. Mus. vol. xxi. p. 597.

1899. Acanthocycline Alcock, J. Asiat. Soc. Bengal, vol. xvii.

pt. 2; p. 96.

Alcock accepts Dana’s legion as a subfamily of the Cancride,

with the definition, ‘‘ Carapace subcireular: front ending in a

triangular point. Hpistome short, sunken, completely concealed

by the external maxillipeds which also completely cover the buccal

orifice. Antennal flagella absent.” The second character must

be modified for the specimens referred to A. albatrossis, as in them

the front is not triangular.

Gen. AcantHocycius M.-Edwards & Lucas.

1844. Acanthocyclus Milne-Edwards & Lucas, D’Orbigny’s Voy.

Amer, Merid. vol. vi. pt. 1, p. 30.

1849. Acanthocyclus Nicolet, Gay’s Hist. Chile, Zool. vol. ii. p. 176.

1898. Acanthocyclus Rathbun, Pr. U.S. Mus. vol. xxi. p. 597

(with further synonymy).

ACANTHOCYCLUS ALBATROSSIS Rathbun.

1861. Acanthocyclus gayi Strahl, Monats. Ak. Wiss. Berlin, July

25, 1861 (1862), p. 713, pl.

» Targioni-Tozzetti, R. Ist. Stud. super.

Firenze, vol. i., Crost. della Magenta,

p- 95, pl. 7. fig. 1, a-f.

1898. A. albatrossis Rathbun, Pr. U.S. Mus. vol. xxi. p. 599.

Miss Rathbun distinguishes three species of the genus—the

original A. gayi M.-Edwards & Lucas, 1843, renamed 4. villosus

by Strahl in 1861, Strahl’s A. gay, for which preoccupied name

A. albatrossis 1s substituted, and A. hassleri, discovered by

Dr. Faxon, but by his wish described and named by Miss Rathbun,

who remarks that “the general appearance of the three species

is much the same,” but that the differences are constant. Of

these eleven are tabulated, and, granting in each case the con-

stancy of the combination, the specific distinction may be justified.

Taken separately, many, even most if not all, of the differences

relied on, have a rather untrustworthy appearance. Thus the

front is entire in gayi and hassleri, faintly bilobed in albatrossis ;

the dactyli of ambulatory legs are long, little curved in the last,

short, much curved in the other two; both carapace and legs are

very hairy in gayi, less hairy in both the others, which again have

the carapace tuberculate, whereas in gayi it is almost smooth ;

1877. "

CRUSTACEA FROM THE FALKLAND ISLANDS. 345

the pleon of the male is narrow in gay2, wide in albatrossis, inter-

mediate in hassleri. Without having specimens from various

localities for comparison, it would be presumptuous for me to

contest Miss Rathbun’s judgment on the matter. I accept the

name albatrossis for the Falkland Islands specimens, inasmuch

as they have the front faintly bilobed, the dactyli long, with a

curvature which may be relatively less in a long finger than in a

short one, the carapace not very hairy (but at the same time

generally smooth except in front), and the fourth joint of the

third maxilliped with the outer margin diverging slightly from

that of the preceding joint.,

The carapace is stated to be in A. gayi “narrow; width 1-05

to 1-08 times length”; in A. albatrossis “ width mtermediate,

1-08 to 1°12 times length”; in hasslert “wide; width 1:16 times

length.” In measuring Falkland Islands specimens, I was certainly

prejudiced in favour of the name albatrossis by finding the

measurements respectively, width 1:12 times length of the first

example and 1-08 of the second. These were females laden with

egos. Then a male gave width 1:15 times length, with the pleon

decidedly narrow. The females laden with eggs vary greatly in

size, One specimen measuring 13°5 mm. long, 15 mm. broad,

another 23 mim. long, 26 mm. broad. The right cheliped appears

to be usually, but not invariably the larger. The ambulatory

legs have a dense fringe of hairs, but whether this leaves them

less hairy than those of 4. gayi 1s matter for comparison.

Locality. Roy Cove, Nov. 12, 1909, and June 16—24, 1910.

Tribe CATOMETOPA.

Fam. GRAPSID&.

1900. Grapside Alcock, J. Asiat. Soc. Bengal, vol. xix. pt. 2,

p- 283.

Gen. PLANES Bowdich.

1825. Planes Bowdich, Excursions in Madeira and Porto Santo,

Da Lomesse2 210.

1910. Planes Stebbing, Ann. 8. Afr. Mus. vol. vi. p. 320.

PLANES MINUTUS (Linn.).

1758. Cancer minutus Linn. Syst. Nat. ed. 10, vol. 1. p. 625.

It is not surprising that this little wanderer has been taken by

Mr. Vallentin at the Falkland Islands, its distribution being very

extensive.

MACRURA ANOMALA.

Tribe PAGURIDEA.

Fam. PAGURID 4.

Gen. EKupracurus Brandt.

1851. Hupagurus Brandt, Middendorff's Sibirische Reise, Zool.

pt. I, p. 105.

346 THE REY. T. R. R. STEBBING ON

‘Kupacurus Forceps (Milne-Edwards),.

1836. Pagurus forceps Milne-Edwards, Ann. Sci. Nat., Zool.

Ser, 2) vol. wis ps 202) poled: ties:

1837. - » Milne-Edwards, Hist. Nat. Crust. vol. 1.

p. 221.

1847. Pagurus comptus White, Pr. Zool. Soc. London, vol. xv.

p. 122.

1858. Hupagurus comptus Stimpson, Pr. Ac. Philad. p. 237 (75).

1871. Pagurus forceps? Cunningham, Tr. Linn. Soc. London,

vol. xxvii. p. 495.

1881. Hupagurus comptus Miers, Pr. Zool. Soc. London, p. 72.

1900. bs 59 Stebbing, Pr. Zool. Soc. London, p. 535.

In now identifying White’s species with that described by

Milne-Edwards eleven years earlier, I am bound to offer some

grounds for my change of opinion. As years pass on, the form

known as /. comptus is repeatedly collected in the southern parts

of South America, and this form agrees well with the description

and figure given by Milne-Edwards for his E. forceps, with the

remarkable exception of the smaller cheliped. This he describes

and figures as having the palm extremely short, the fingers slender,

long and pointed, the movable finger almost filiform and straight,

or even sinuous. There is something so unusual in this character

of the smaller cheliped, that, as it does not appear ever to have

been observed again, one may be excused for regarding it as an

abnormality. It is not at all certain that Milne-Edwards had

more than one specimen, though he gives an alternative for the

form of the movable finger. He figures it with two slight curves

but base and apex in line one with the other. That the species

is hable to abnormality may be judged from the figure of the

larger cheliped in Zool. ‘Erebus’ and ‘Terror,’ Crustacea, pl. 2.

figs. 5, 5 « (1874), where the movable finger is evidently stunted.

The variability of the species is further shown by the fact that

Miers thought it necessary to name a var. latimanus, and Hen-

derson a var. jugosa.

Mr. Vallentin obtained specimens from Macrocystis.

Tribe GALATHEIDEA.

Fam. GALATHEID &.

Gen. MunipA Leach.

1820. Munida Leach, Dict. Sci. Nat. vol. xviii. p. 52.

ROMO estas Stebbing, Ann. 8. Afr. Mus. vol. vi. p. 364.

MUNIDA GREGARIUS (Fabricius).

1793. Galathea gregaria Fabricius, Ent. Syst. vol. ii. p. 473.

1891. Munida gregaria Mocquard, Miss. Cap Horn, Crustacés,

p. 32, pl. 2. figs. 1, 1 a-c.

CRUSTACEA FROM THE FALKLAND ISLANDS, 347

1902. Munida gregaria Benedict, Pr. U.S. Mus. vol. xxvi. p. 308,

figs. 45, 46 (with synonymy).

III - 5. Ortmann, Princeton Univ. Exp. Pata-

gonia, p. 659.

Specimens of various sizes which I refer to this species were

taken by Mr. Vallentin Jan. 28, 1899 (at the surface), Dec. 4,

1901, in Roy Cove, Feb. 1, 1910, and in Whales Bay, March 11

of the same year.

Munna susrucosus (White).

1847. Galathea subruyosa White, List of Crust. Brit. Mus. p. 66.

1852-5. Munida subrugosa Dana, U.S. Expl. Exp. vol. xiii. p. 479,

pl. 30. fig. 7 a-e.

1891. a 5 Mocquard, Miss. Cap Horn, Crustacés,

p- 36, pl. 2. figs. 2, 2. a-c.

1909. “3 i Chilton, Subantarctic Is. of N.

Zealand, p. 612 (with synonymy).

eo ele a h Ortmann, Princeton Univ. Exp.

Patagonia, p. 659.

Dr. Chilton discusses the question of the specific identity of

M. subrugosus and M. gregarius, 11 which he definitely inclines to

believe, though still adopting the later name for the species. As

to the generic name, Grimothea Leach, 1820, seems to have page

precedence over J/unida, but I forbear the dangerous task of

arbitrating between the rival nymphs.

Mr. Vallentin took MW. subrugosus at a depth of 4 fathoms in

Whales Bay, 6 fathoms in Roy Cove, and Dec. 12, 1909, in

““8 fathoms creek.” All these specimens were adult, and all but

one of rather large size.

MACRURA GENUINA.

Tribe CARIDEA.

Fam. HiprPoLtyTipD&.

1888. Hippolytide Bate, Rep. Voy. ‘Challenger,’ vol. xxiv.

p. 976.

Gen. Nauricaris Bate.

1888. Nauticaris Bate, Rep. Voy. ‘Challenger,’ vol. xxiv. p. 577.

Having already noticed this genus in another (as yet un-

published) paper, I refrain from further discussion here.

NAUTICARIS MAGELLANICUS (A, Milne-Edwards).

1891. Hippolyte magellanicus A. Milne-Edwards, Miss. Cap

Horn, Crustacés, p. 46, pl. 5. figs. 2, 2 a-7.

This species was taken by Mr. Vallentin from root of JMJacro-

348 : THE REY. T. R. R. STEBBING ON

cystis on Dec. 29, 1910, and at various other dates, by hand-net

from bed of Macrocystis in Roy Cove, from a depth of between

3 and 4 fathoms, and in the same locality two specimens from a

depth of 6 fathoms, on which he notes that one was banded red

and chocolate in colour, and the other cream-coloured with

chocolate markings.

ISOPODA ANOMALA

(or Apseudacea).

1902. Zsopoda anomala Stebbing, 8. African Crustacea, pt. 2,

p- 48.

GAO} i Fe Stebbing, Ann. 8. Afr. Mus. vol. vi.

pp. 413, 576.

Fam. TANAID&.

1905. Tanaide H. Richardson, Bull. U.S. Mus. no. 54, p. 3.

1905. Ry Stebbing, Herdman’s Pearl Fish. Rep., no. 23,

p- 2.

LOWS: A Nierstrasz, Siboga-Expeditie, Mon. 32 a, p. 20.

The last of these references supplies an ample bibliography of

the Apseudacea. In my own treatise above mentioned, on p. 4,

T attribute to Sars the statement that in his genus Heterotanas

the palp of the first maxilla is terminated by a single seta, a

mistake for which I cannot account, as he distinctly states that

there are two sete. The distinction which he does in fact draw

is that in Tunais the palp is biarticulate and tipped with several

sete, but in Heterotanais uniarticulate with the sete only two.

These minute features have been so seldom attended to in

descriptions, that they are difficult to use for the settlement of

genera. It might be convenient to withdraw from Vanais those

species which have six separate pleon segments instead of only

five. But even on this point authors are not always as definite

as could be desired. ‘The species about to be described belongs

clearly to the group in which the pleon has six separate segments.

Asin Tanais, it has only three pairs of pleopods. Its form is

robust like that of J’. robustus Moore, but while that species has

seven setze on the palp of the first maxilla, this has only two. It

shows points of agreement with the much smaller 7’. sewrate

Nobili, 1906-1907, but there the second joint of the second

antenne is much shorter than the first, here the reverse is the

case.

The late Dr. Nobiliin 1907 gives the family name as Tanaidide.

As, however, the genitive case of Vanais, in Latin, is the same

as the nominative, it can scarcely be necessary to alter the

accustomed form.

CRUSTACEA FROM THE FALKLAND ISLANDS. 349

Gen. Tanats Andouin & M.-Edwards.

1829. Tanais Audouin & Milne-Edwards, Précis d’Entomologie,

vol. i. p. 46, pl. 29. fig. 1.

TANAIS OHLINI Stebbing. (PI. I.)

Abstract P.Z.S. 1914, p. 30. (April 28.)

The segments of the pleon successively decrease in length to

the sixth, which is longest of all, the curve of its hind margin

slightly extended at the middle; the fourth and fifth segments

much the shortest.

The eyes are irregularly ovoid, bluntly narrowed forward, the

front margin of the head between them not clearly made out.

First antenne with third joint a little shorter than second, and

less than one third as long as the first; flagellum consisting of a

minute joint, broader than long, with fascicle of sete. Second

antenne more slender and a little shorter than the first; first,

joint shorter than third, which is a little over half the fifth, fifth

rather shorter than fourth, fourth than second ; flagellum one

little joint with fascicle of sete. One of the mandibles has a

small tooth-hke accessory plate. Whether the palp of the first:

maxille is divided into two joints or not, could not be made out 5

one of its apical sete is very much longer than the other. The

maxillipeds have an unguis-like spine or fifth joint at the apex.

The large first gnathopod is of the ordinary type, the movable

finger rather longer than the thumb, the apical points of the two

curving one towards the other ; inner margin of the thumb thin,

not continuous with that of the apex, and flanked with setules.

The slender second gnathopods have the antepenultimate joint

not distally widened and without spines, the next joint very

much longer, with needle-like finger half its length. The two

following pairs of pereopods have the antepenultimate joint

distally widened, only a little shorter than the following joint,

and distally fringed with small spines, the hand and finger similar

to those of the second gnathopod but shorter. The three

succeeding pairs of pereeopods have the second joint robust, the

hand not longer than the wrist, slightly curved, the finger sickle-

shaped, with four little spines or teeth on the concave margin

near the apex. The marsupium on the penultimate segment of

the perzeon was crowded with rather large eggs. The pleopods

are as usual strongly setose. The uropods are six-jointed, the

largest joint constituting the peduncle, the five small joints of the

single ramus being, except the first, plentifully furnished with

sete.

Length estimated at 4°5 mm., supposing the body to be

flattened out.

Locality. Roy Cove at low spring tide.

The specific name is given in memory of the late Dr. Axel

Ohlin, whose valuable researches in the Falkland Islands have

only been in part reported on, death having interfered with the

fulfilment of his plans.

(JU)

(‘>

THE REY. T. R. R. STEBBING ON

ISOPODA GENUINA.

Tribe FLABELLIFERA.

Fam. SPH HZROMIDZ.

1847. Spheromide White, List of Crustacea in Brit. Mus.,

p. 102.

1910. 43 Stebbing, Ann. 8. Afr. Mus. vol. vi. p. 426.

Gen. ExospH#RomA Stebbing.

1900. Hxospheroma Stebbing, Pr. Zool. Soc. London, p. 553.

1905. be Hansen, Quart. J. Microsc. Sci. vol. xlix.

PpalOs. ils:

ESOS e Stebbing, Ann. 8. Afr. Mus. vol. vi. p. 428.

This genus is placed by Hansen in the section Spheromini of

his group Spherominz hemibranchiate,

EXxosPHAROMA CALCAREUS (Dana). (PI. IT.)

1853-55. Spheroma calearea Dana, U.S, Expl. Exp. vol. xiii.

p..776, pl. 52, fig. 2 a-c.

1891. Spheroma - Dollfus, Miss. du Cap Horn, Crust.

p: 64, pl. 8a. figs. 7, 7a, 7 0.

1913. Hxospheroma coatsii Tattersall, Tr, R. Soc. Edinb. vol. xlix.

p. 885, figs, 3, 4.

This is one of the species which are now in rapidly increas-

ing number perplexing the systematist by their variability. A

comparison of the figure supplied by Dana in 1855 with

Dr. Tattersall’s in 1913 would scarcely suggest a suspicion of

specific identity. As it is, probably some allowance must be

made for a little want of detail in Dana’s sketch. But Dollfus,

who had at command several specimens, explains that the

granules and tubercles on the general surface and the double

crest on the pleon sometimes disappear, leaving a smooth form

such as Dana represents. Dr. Tattersall, describing and figuring

an adult female and a young form, from Dr. Bruce’s Scottish

Antarctic Expedition, noted the comparative infrequency of

tubercles in the young, with other differences, but he had no

mature male to test for sexual difference. This deficiency I have

been able to supply from Mr. Vallentin’s collections. The

specimen figured was 13 mm. long by 7 mm. broad. A female,

16 mm. long, containing a great number of eggs, was taken by

Mr. Vallentin at low ebb of a spring tide in Stanley Harbour,

Nov. 12, 1901. Other specimens, taken at Rapid Point, low

water, Jan. 30, 1911, comprise a male 19 mm. long by 10 mm.

broad at the sixth pereon segment. This capture corroborates

the statement by Dollfus that he had observed males which were

strongly granular and others almost smooth. Variation also

affects the colour, at least to judge by preserved specimens,

CRUSTACEA FROM THE FALKLAND ISLANDS. 351

some being a uniform brown, while others long retain signs of

a brilliant marbling, such as that suggested by Dana of purple

patches on a yellow ground.

Gen. Casstprnopsts Hansen.

1905. Cassidinopsis Hansen, Quart. J. Microsc. Sci. vol. xlix.

pp. 77, 82, 87, 90, 94, 106, 109, 128, 130.

Hansen places this genus in his group Spheromine eubranchiate,

with the definition, ‘“‘ Head small, narrow in proportion to largest

breadth of thorax. Basal joint of antennule without process

from the distal posterior angle. End of abdomen feebly

emarginate. Uropoda similar in both sexes; endopod laterally

expanded, very much broader and a little longer than exopod.

Both sexes similar, without processes ; female with normal

mouth-parts and the brood in internal pouches.” He states that

“the type is Cassidina emarginata (Guér.), which in many

important points—structure of plp.’ and plp.’, shape of epistome,

mandibles, fifth joint of mavxillipeds, end of abdomen—differs

strongly from the type for the genus Cassidina, C. typa (M.-Kdw.).”

CASssIDINOPSIS EMARGINATUS (Guérin-Méneville).

1843. Cassidina emarginata Guérin - Méneville, Icon. Regne

Animal, Crust., texte, p. 31.

1900. 3 Stebbing, Proc. Zool. Soc. London,

p- 562 (with synonymy).

1905. Cassidinopsis ,, Hansen, Quart. J. Microse. Sct.

WOM, aeiixee yon CWle

1910. * e Hodgson, Nat. Antarct. Exp.

vol. v. p. 4.

1911. Cassidina Ortmann, Princeton Univ. Exp.

Patagonia, vol. 111. p. 650.

The pigmented portion of the eyes has a conical shape, the

narrow end foremost. Mr. Vallentin at various dates obtained

specimens of different sizes, especially at Roy Cove, from fronds

of Macrocystis between 2 and 4 fathoms.

VALLENTINIA, gen. NOV.

A member of the Spheromine eubranchiate, near to Paracercets

Hansen, 1905, but distinguished by not having the basal joint

of the first antenne produced into an acute process, the mandibles

of the female not coalesced with the head, the exopod of the

uropods much shorter and narrower than the endopod, first

gnathopod prehensile in the male.

VALLENTINIA DARWINIE (Cunningham).

1871. Cymodocea darwintt RB. O. Cunningham, Tr. Linn. Soe.

London, vol. xxvii. p. 499, pl. 59. figs. Il, Ul ae 1,

352 THE REY. T. R. R. STEBBING ON

- 1881. Dynamene darwinii Miers, P. Z.5. Lond. p. 79.

U-1884. Cymodocea darwin Studer, Abh. K. Ak. Wiss. Berlin,

1883, p. 18, pl. 2. figs. 6, 6.4, 66.

1886. is darwint Beddard, Rep. Voy. ‘Challenger,’

vol. xvii. pt. 48, p. 150.

- 1891. a darwintt Dollfus, Miss. Cap Horn, Crustacés,

p. 65, pl. 8. figs. 8, 8a, 8b.

EOI 55 darwiu Ortmann, Princeton Univ. Exp.

Patagonia, p. 649.

Hansen in his treatise on the Spheromide is evidently alluding

to this species when he says (p. 125), “According te kind

information from Dr. Calman, D. Darwiniit (Cunningham) has

exopod of plp.* divided by an articulation; the species must, in

my opinion, be established as a new genus near Paracerceis.”

On this recommendation I have acted, naming the genus after

Mr. Vallentin, te whose researches I owe the opportunity of

examining the species. An interesting feature is the dilatation

at the fifth pereeon segment, well marked in Cunningham’s figure,

and noticed by Dollfus but scarcely appreciable in his coloured

drawing of an example 19 mm. long. The specimen I have had

under observation measured only 9mm. The apical emargination

of the telsonic segment is squared at the base. The epistome is

not like the figure given by Dollfus; it widens much more

abruptly backwards, and then narrows before forming the divergent

arms which clasp the upper lip. The mouth-organs are much as

in Cymodoce. The first antenne have a very large first joint

followed by a short one, to which succeeds one that is long and

slender. ‘The first gnathopods are rather robust, but as Dollfus

notices, the large tooth produced from the base of the hand is no

doubt a male eharacter, giving to that sex im this genus a pair of

prehensile hands.

In his eubranchiate group Hansen makes an infermal separation

between the genera which: have and those which have not an

articulation of the exoped in the third pleopod. In the present

species the articulation is very conspicueus, through the strong

incurving of the inner margin of each joint at the junction; the

exopod itself is unusually narrow. The fourth and fifth pleopods,

in accord with their systematic position, have beth rami strongly

pleated. There are five of the denticulate bosses on the end of

the exopod in the fifth pair; the exoped ef the feurth pair is

clearly two-jointed.

Locality. Stanley Harbour, low water.

‘Tribe VALVIFERA.

Fam. ASTACILLIDS.

1897. Astacillide Sars, Crustacea of Norway, vol. ii. p. 88.

1901. SA Ohlin, Svenska Exp. Magellanslind. vol. ii.

p. 265,

CRUSTACEA FROM THE FALKLAND ISLANDS, S18}

Gen. AstacttiA Cordiner.

1795. Astacilla Cordiner, Remarkable Rivers, and Nat. Hist.,

Section “ Astacille.”

1905. - Stebbing, Herdman’s Pearl Fish. Rep., Suppl.

Rep. 23, p. 46.

ASTACILLA FALCLANDICUS Ohlin.

1901. Astacilla falelandica Ohlin, Svenska Exp. Magellanslind.

vol. 11. p. 266, pl. 20. fig. 1.

I have very little doubt that Ohlin’s Astacilla magellanicus is

a synonym of this species. Mr. Vallentin’s specimens were

obtained in the Falklands from hulks at low water. The first

antenne have a few filaments in an apical group. The first

pereon segment is completely coalesced with the head, and the

rather compact little first gnathopods are so attached that they

can scarcely have any function but that of mouth-organs. The

slender second gnathopods and first two pairs of pereeopods have

the natatory sete not at all densely crowded; they havea minute

hooked spine as representative of the seventh joint. The hind

pereopods are robustly uncinate. The length of the body is

between 4 and 5 mm.

Fam. IDOTEID &.

1852. Idoteide Dana, Amer. Journ. Sci. ser. 2, vol. xiv. p. 300.

1911. 3 Tattersall, Nordisches Plankton, vol. iii. p. 216,

Gen. Epott1a Guérin-Méneville.

1843. Hdotia Guérin-Méneville, Icon. Régne Animal, p. 34.

1901. Hdotia Ohlin, Svenska Exp. Magellansliind. vol. 11. p. 292.

EporiA TUBERCULATUS Guérin-Méneville.

1843. Hdotia tuberculata Guérvin-Méneville, Icon. Régne Animal,

p- 04.

TO OME 55 35 55 Ohlin, Svenska Exp. Magellanslind.

vol. ii. p 292, pl. 23. figs. 10, 10 4-c, ete.

The synonymy, characters, and distribution of this species are

well discussed by the late Dr. Axel Ohlin. More recently it is

noted by Ortmann and Hodgson. Mr. Vallentin took specimens

in Roy Cove, from a depth of between 3 and 4 fathoms.

Gen. MACROCHIRIDOTHEA Ohlin.

1901. Macrochiridothea Oblin, Svenska Exp. Magellansliind.

vol. ii. pp. 282, 286.

The great development of the first gnathopods in both sexes

354 THE REY. T. R. R. STEBBING ON

is referred to in the name of the genus, which also alludes to its

alliance in various other respects with Chiridotea Harger. As in

that genus, the so-called palp of the maxillipeds is three-jointed,

but alike in C. cecus (Say) and C. tuftsii (Stimpson) the first

joint of the palp is much the shortest, whereas in the two species

of Ohlin’s genus that proportion belongs to the third joint.

MACROCHIRIDOTHEA STEBBINGI Ohlin.

1901. Macrochiridothea stebbingi Ohlin, Svenska Exp. Magel-

lanslind. vol. 11. p. 289, fig. 9.

The species has been amply described and figured by Dr. Ohlin

from a female specimen, 7 mm. long. Mr. Vallentin obtained a

specimen 15 mm. in length, another 14 mm., both 6-5 in breadth,

and a third of nearly the same length as the second. As these

all happened to be females, there was no opportunity of com-

paring the male appendix with that of Ohlin’s other species,

M, michaelsenii, of which he gives the measurements as “ length

of males 11°5 mm.; breadth 5°55 mm. Female smaller.”

Locality. Port Harriet, low-water mark spring-tide.

Tribe ASELLOTA.

Fam. JANIRID&.

1897. Janiride Sars, Crustacea of Norway, vol. i. p. 98.

1901. Janiride Richardson, Pr. U.S. Mus. vol. xxii. pp. 497,

550, 553.

Stebbing, Herdman’s Pearl Fish. Rep., Suppl.

Rep. 23, p. 48.

1905.

Gen. Notasetuus Pfeffer.

1887. Notasellus Pfeffer, Jahrb. wiss. Anstalten Hamburg, vol. iv.

p. 85.

1902. & Hodgson, Nat. Hist. Southern Cross Exp. p. 251.

1905. am Stebbing, Herdman’s Pearl Fish. Rep., Suppl.

Rep. 23, p. 53.

1910. My Richardson, Pr. U.S. Mus. vol. xxxvu. p. 649.

1913, A Richardson, Deuxiéme Exp. Antarct. francaise,

Isop. p. 17.

NorasELLus sarsi Pfeffer.

1887. Wotasellus sarsii Pfeffer, Jahrb. wiss. Anstalten Hamburg,

vol. iv. p. 85, pl. 7. figs. 5-28.

This species has been very fully described and illustrated by

Dr. Pfeffer. Specimens were taken by Mr. Vallentin at Rapid

Point, low water, Jan. 30, 1911.

CRUSTACEA FROM THE FALKLAND ISLANDS. 355

AMPHIPODA.

Tribe GAMMARIDEA.

Fam. LySraNASSID&.

1874. Lysianasside Buchholz, Zweite D. Nordpolarf. vol. ii.

p. 299.

1913. bighs Chevreux, Deuxiéme Exp. Antarct. francaise,

p- 87.

Gen. TRYPHOSITES Sars.

1891. Tryphosites Sars, Crust. Norway, vol. i. p. 81.

1906. Pi Stebbing, Das Tierreich, vol. xxi. p. 77.

SNe _ Sexton, Ann. Nat. Hist. ser. 8, vol. vii. p. 510.

1912. 54 Chilton, Tr. Roy. Soc. Edinb. vol. xlvili. pt. 2,

p- 469.

To receive the new species here referred to this genus, its

definition must be a little modified, by withdrawal of the state-

ment that the postero-lateral angles of the third pleon segment

are acutely upturned, nor does the shape of the hand in the

second gnathopod precisely conform with that in the type species,

Also the inner ramus of the second uropod is not constricted.

TRYPHOSITES CHEVREUXI Stebbing. (Pl. IIT.)

Abstract P. Z.S. 1914, p. 30. (April 28.)

The third pleon segment, instead of having the postero-lateral

corners upturned with a smooth concave margin above, has the

lower half of the postero-lateral margin convex and cut into

a serration of nine little teeth. Thus the species is sharply dis-

tinguished both from 7”. longipes (Bate & Westwood) and from

Hoplonysx stebbingi Walker, 1903, which Chilton in 1912 transfered

to V'ryphosites, with the remark, among others, that it “ appears

to be very close to 7’. longipes of northern seas, differing chiefly

in having the perzopoda shorter and stouter and the eyes in-

distinct.” Walker lays some stress upon “the absence of a

depression ” dorsally in the fourth pleon segment. Such a de-

pression is sometimes masked by the telescoping of the segment.

In the new species the depression is very marked.

Eyes obscure or absent. Both pairs of antennee strongly

resembling those of 7’. Jongipes. Accessory flagellum in first pair

of the male 7-jointed, principal with 16 joints, some of which

carry small calceoli. Flagellum of second pair not so long as the

body, with 32 joints, several of the alternate ones carrying cal-

ceoli, decreasing in size on the distal portion. Mouth-organs in

close agreement with those of 7’. longipes.

The gnathopods differ from those of the two earlier species in

scarcely anything but the hand of the second pair, which is not

Proc. Zoou. Soc.—1914, No. XXIV. 24

356 THE REY. T. R. R. STEBBING ON

quite half as long as the wrist. The pereopods are distinguished

chiefly by the strong denticulation of the hind margin in the

second joint of the fifth pair; in this and the two preceding

pairs the terminal joints are not so long and slender as in

7. longipes ; the fourth joint is rather narrower in the fifth pair

than in the two preceding pairs. The branchial vesicles show

various proximal folds.

The telson is divided nearly to the base, each division having

three submarginal spines, and three apical, of which the central

is the longest, with a setule between it and the very small outer

spine.

Length of male9 mm. A second specimen, with flagellum of

second antenne broken, the remainder of seven joints carrying no

calceoli and suggestive of a short termination, is probably the

female. It measures 6 mm., and lke the male is very narrow,

with the curved process of the epistome conspicuous.

Locality. Roy Cove, from the depth of 8 fathoms. Specimens

from Whales Bay, observed after the above description was

written, show the second antenne a little longer than the first.

The species is named in honour of my friend, M. Edouard

Chevreux, a brilliant student of the Amphipoda.

Gen. AcontiostoMA Stebbing.

1888. Acontiostoma Stebbing, Rep. Voy. ‘Challenger,’ vol. xxix.

3 (0S)

1906. bi Stebbing, Das Tierreich, vol. xxi. pp. 9, 15.

ACONTIOSTOMA MARIONIS Stebbing.

1888. Acontiostoma marionis Stebbing, Rep. Voy. ‘ Challenger,’

WOl, Sx, Jo, FOS), jolly BO),

1893. B Ke Della Valle, F. & Fl. Neapel, vol. xx.

p. 786.

1906. ss ” Stebbing, Das Tierreich, vol. xxi.

p. 15, text-fig. 4.

1912: A F Chilton, Tr. R.Soc. Edinb. vol. xlviu.

p- 462.

Mr. Vallentin obtained a specimen from roots of Macrocystis

on Jan. 14, 1902, and another, 7 mm. long, much more recently

at Rapid Point, low water of spring tide. The finding of these

specimens in the Falkland Islands increases the probability, with

which Professor Della Valle naturally agrees, that my Acontio-

stoma magellanicus is merely a young form of A. marionis.

Fam. AMPELISCID2.

1882. Ampeliscide Sars, Forh. Selsk. Christian. no. 18, p. 29.

1906. Ms Stebbing, Das Tierreich, vol. xxi. pp. 6,

Sie 2).

CRUSTACEA FROM THE FALKLAND ISLANDS. a

“I

Gen. AmpEtiscA Kroyer.

1842. Ampelisca Kroyer, Naturh. Tidsskr. vol. iv. p. 154.

1906. Ps Stebbing, Das Tierreich, vol. xxi. pp. 98, 721.

AMPELISCA MACROCEPHALUS Liljeborg.

1852. Ampelisca macrocephala Liljeborg, Ofv. Ak. Férh, vol. ix.

We Ue

1903. af Ms Walker, J. Linn. Soc. London,

a vol. xxix. p. 53, pl. 9. figs. 58-61.

1905. ‘ an Holmes, Bull. U.S. Bureau Fish.

vol. xxiv. p. 479, text-figs.

1905. br iF Paulmier, Bull. New York Mus.,

Bull. 91, Zool. 12, p. 158, fig. 26.

1906. a. Ps Stebbing, Das Tierreich, vol. xxi.

ojo SO, VOI.

1907. . is Walker, Nat. Antarct. Exp. vol. tii.

We Wee

It may seem extraordinary that this northern and even arctic

species should reappear, as Mr. A. O. Walker has determined, in

antarctic waters. It has been taken by Mr. Vallentin at low

water at spring tides on a sandy beach in Shallow Bay,

Falkland Islands, Jan. 15,1911. The bright red pigment of the

eyes lasts long in preservative fluid.

Fam. PHOXOCEPHALIDA.

1891. Phoxocephalide Sars, Crustacea of Norway, vol. i. p. 142.

1906. F Stebbing, Das Tierreich, vol. xxi. pp. 6,

USB}, (233.

Gen. PonTHARPINIA Stebbing.

1899. Pontharpinia Stebbing, Tr. Linn. Soc. London, ser. 2

vol. vii. p. 32.

1906. . Stebbing, Das Tierreich, vol. xxi. p. 146.

OMS 5 Chevreux, Deuxiéme Exp. Antarct. fran-

caise, Amph. p. 101.

PONTHARPINIA ROSTRATUS (Dana).

1853-55. Urothoe rostratus Dana, U.S. Expl. Exp. vol. xiii. p. 921,

pl. 62. fig. 5 ap.

1906. Pontharpinia rostrata Stebbing, Das Tierreich, vol. xxi.

p. 146.

Chevreux’s Pontharpinia uncinatus is distinguished by the

shorter wrist of the second gnathopods and the upturned postero-

lateral angles of the third pleon segment, but in many respects,

as the eminent French author observes, isa near neighbour of the

present species.

Locality. Falkland Islands, low water of spring tide.

24*

358 ; THE REY. T. R. R. STEBBING ON

Fam. METOPIDS.

1899. Metopide Stebbing, Ann. Nat. Hist. ser. 7, vol. iv. p. 210.

1906. i Stebbing, Das Tierreich, vol. xxi. pp. 7, 171, 724.

Gen. Mrropreua, Sars.

1892. Metopella Sars, Crustacea of Norway, vol. i. p. 274.

METOPELLA ovatus (Stebbing).

1888. Metopa ovata Stebbing, Rep. Voy. ‘Challenger,’ vol. xxix.

p. 764, pl. 44.

1893. Metopoides ovatus Della Valle, F. & Fl. Neapel, pp. 645, 907,

938. iP

1906. Metopella ovata Stebbing, Das Tierreich, vol. xxi. p. 183,

figs. 47, 48.

rs » Chilton, Trans. R. Soc. Edinb. vol. xlviil.

p. 481.

A female specimen, containing five large eggs, measured in its

folded posture less than 1 mm. in length.

Locality. Stanley Harbour, on seaweed, at low water of spring

tide.

1912,

Fam. ACANTHONOTOZOMATIDAE.

1906. Acanthonotozomatide Stebbing, Das Tierreich, vol. xxi.

joe Mi, Zl;

Gen. IputmepiA H. Rathke.

1843. Iphimedia Rathke, N. Acta Ac. Leop. vol. xx. p. 85.

; He Stebbing, Das Tierreich, vol. xxi. p. 214.

1907. 3 Walker, Nat. Antarct. Exp. vol. iil. p. 37.

1910. 5 Stebbing, Mem. Australian Mus. vol. iv. pp. 584,

637.

IPHIMEDIA NoDOsus Dana.

1852. Iphimedia nodosa Dana, P. Amer. Ac. vol. ii. p. 217.

1906. 5 » Stebbing, Das Tierreich, vol. xxi.

pp. 214, 216.

The identification and fuller description of Dana’s species

supplied in 1906 were made possible by the specimens which

Mr. Vallentin obtained at low water of a spring tide in Stanley

Harbour. More recently, Dec. 29, 1910, he obtained a specimen

from the root of Macrocystis.

Gen. ParirHiMEpDIA Chevreux.

1906. Pariphimedia Chevreux, Bull. Soc. Zool. France, vol. xxxi.

NOs 2Vpsooree

1906. 5 Chevreux, Exp. Antarct. francaise, Amphip.

p- 38.

CRUSTACEA FROM THE FALKLAND ISLANDS. 359

1910. Pariphimedia Stebbing, Mem. Australian Mus. vol. iv.

pt. 2, p. 584.

1912. a Chilton, Tr. Roy. Soc. Edinb. vol. xlviii.

p- 487.

H. Rathke’s [phimedia and G. M. Thomson’s Panoplea,

according to Chevreux, are distinguished from this genus chiefly

by the following characters :—the cutting-edge of the mandibles

not denticulate, the principal lobes of the lower lip emarginate

on the inner edge, the palp of the first maxille two-jointed, the

inner lobe of the second maxille carrying a single series of

sete, the second gnathopod not completely chelate, and the

telson apically emarginate.

It would, I think, be inconvenient to press the first of these

characters as essential to either of the genera mentioned ; but

both are rather sharply separated from Pariphimedia by the two-

jointed palp of the first maxilla, in strong contrast with the feeble

single-jointed structure in Chevreux’s genus, which makes an

approach to that found in Odius Lilljeborg.

In adding a second species to the genus, I feel fairly sure that

it is identical with the scantily-described Jphimedia normant

Cunningham, which has so long remained obscure. But the

addition tends to weaken the original definition, inasmuch as the

telson has a neat little convex emargination which helps to dis-

tinguish it from P. integricauda, in which, as the specific name

declares, the telson is unincised. By a curious contrariety the

upper lip, which is there slightly emarginate, is here simply

convex. The mandibles in the two species essentially agree, the

trunk tapering to a fine point, the distal part of the margin

minutely denticulate, the molar represented by a projection with

no triturating surface, the palp well developed, its second joint

much the longest, the third curved, setose. Neither mandible in

P. norman has a secondary plate, so far confirming the sugges-

tion which I have earlier made, that the same is the case in

P. integricauda. While it may be said that the distal lobes of

the lower lip are in both species undivided, it will be seen that in

P. norman there is a marked tendency towards apical division.

PARIPHIMEDIA NORMANI (Cunningham). (Pls. LV. & V.)

1871. Iphimedia normanit Cunningham, Tr. Linn. Soc. London,

vol. xxvii. p. 498, pl. 59. fig. 7.

1906. 5 a Stebbing, Das Tierreich, vel. xxi. p. 217.

This species, by the notable features of its mouth-organs and

enathopods, clearly belongs to the family Acanthonotozomatide.

But of spine-like processes on the back, which so many members

of that family exhibit, it is singularly devoid. Only the third

segment of the pleon makes a show of relationship in this respect

by an upturned postero-lateral angle and high up on the side

a still stronger upturned tooth. The two preceding segments

have the postero-lateral angle produced into acute points, and

360 THE REY. 't RB. Rk. STEBBING ON

the sides angled. ‘The side-plates of the person have no acute

points except that which forms the boundary of the emargination

in the large fourth pair. The fifth pair are bilobed and not pro-

duced backwards as in the congeneric species.

The eyes have numerous small components. The flagellum of

the first antenne shows fifteen joints, that of the second twenty-

nine, in each case the first joint being much the longest, the

second flagellum about a fifth of its length longer than the first.

The mandibles and maxille are in: close agreement with those

described by Chevreux, but the maxillipeds differ by the greater

length of both the inner and the outer plates, the latter being

nearly as long as the palp; a faint transverse line gives them the

appearance of being jointed.

The gnathopods, perzeopods, and uropods also differ but little -

from those of the companion species, but the second joint of the

first gnathopod is here sinuous, not straight, and the second

joint of the third perzeopod is here broader, with the hind margin

convex.

The specimen, a female with a few large ova, measured about

9 mm., in near agreement with Dr. Cunningham’s specimen,

4 lines long, but much less than the specimen of P. integricauda,

described by Chevreux as 15 mm. in length. The colour as

preserved was marbled red.

Locality. Whales Bay, Falkland Islands, May 17, 1910.

Cunningham states that his specimen was dredged off Elizabeth

Island in February 1867.

Panoplea joubini Chevreux, 1912, strikingly distinguished

from the present species by numerous spiniform processes,

curiously resembles it in the unemarginate upper lip, long plates

of the maxillipeds, emarginate telson, and in the gnathopods.

Fam. Gi DICEROTID &.

1906. @dicerotide Stebbing, Das Tierreich, vol. xxi. p. 235.

Gen. MonocuLopsis Sais.

1892. Monoculopsis Sars, Crust. Norway, vol. i. p. 310.

In many respects this genus agrees with J/onoculodes Stimpson.

Distinguishing features are the considerable size of the fourth

and fifth side-plates, the relatively greater length of the third

joint of the peduncle in the first antenne, and the somewhat

tapering form of the long sixth joint in the second gnathopods.

Monocuuorsis VALLENTINI Stebbing. (Pls. VI. & VII.)

Abstract P. Z.S. 1914, p. 30. (April 28.)

From Monoculopsis longicornis (Boeck), the type of the genus,

the present species is distinguished chiefly by characters of the

gnathopods. In the first pair the process of the wrist or fifth

CRUSTACEA FROM THE FALKLAND ISLANDS, 361

joint, though well pronounced, is very slender and does not

reach the palm. In the second pair the process of the same

joint, instead of being very long and extending beyond the inner

corner of the palm, is very short and quite distant from the palm.

Moreover, the first antenna in the female is decidedly shorter

instead of a little longer than the second, and it has a flagellum

as long as the peduncle instead of one only a little longer than

the peduncle’s third joint. Here, it may be theught, are materials

for establishing a new genus, but that may wait, since the

discovery of intermediate forms might easily make it unnecessary.

The organ of vision on the short rostrum is white in the pre-

served specimens. ‘The first antenne have the first joint as long

as the second and much stouter, the third joint little more than

two thirds as long as the second, the flagellum of sixteen joints.

In the second antenne the last joint of the peduncle is longer

than the stouter penultimate; the gland-cone of the second joint

is blunt-ended ; the flagellum is composed of twenty-two joints,

but fewer in a smaller specimen.

The trunk of the mandibles has the cutting-edge not strongly

dentate, the molar not very prominent, the third joint of the

palp much shorter than the setose second, each curved but in

opposite directions. ‘The spine-row consists of five spines. The

inner plates of the lower lip are distinctly developed. The inner

plate of the first maxille is tipped with three small sete; the

elongate second joint of the palp has several sete along the

outer margin and six spines on the distal part of the inner. The

maxillipeds are like those in the type species.

The oblique palm is longer than the hind margin of the hand

in the first gnathopod, but considerably shorter than it in the

second. The first and second perzopods are alike. The third

and fourth differ from them in the greater expansion of the

second joint. The fourth differs from the third by the greater

size of its second, fifth, and sixth joints. In all four pairs the

sixth joint is notable for the dense clothing of sete along the

back or convex margin of the sixth joint. The small finger is

unarmed except for a microscopic unguis. The long fifth perzo-

pods do not appear to be distinctive.

The pleopods have two minute coupling-hooks on the inner

distal corner of the peduncle, and five coupling-spines on the

first joint of the inner ramus, which is very slightly shorter than

the outer. The uropods have their long peduncles successively

shorter, in each case longer than their respective rami, which

are also long, in the first and second pairs the inner ramus

slightly longer than the outer, the spine armature throughout

rather slight. The telson scarcely longer than broad, with a

minute spinule at each rounded corner of the truncate distal

margin.

The female specimen measured 9 mm. across the curve from

the rostrum to the end of the third pleon segment, so that if

straightened out the full length to the end of the telson might

362 THE REY. T. R. R. STEBBING ON

have been 12 mm., but it is difficult to say what allowance should

be made for the telescoping of the segments,

Locality. Top of Roy Cove, low water, Aug. 1, 1910.

Fam. CALLIOPIID!.

1893. Calliopriide Sars, Crust. Norway, vol. i. p. 431.

1906. c Stebbing, Das Tierreich, vol. xxi. pp. 285, 727.

Gen. Hatiraces Boeck.

1871. Halirages Boeck. Forh. Selsk. Christian. 1870, p. 194.

1906. ac Stebbing, Das Tierreich, vol. xxi. pp. 285, 290.

HALIRAGES HUXLEYANUS (Bate).

1862. Atylus huxleyanus Bate, Catal. Amph. Brit. Mus. p. 135,

pl. 25. fig. 4.

1888. Halirages huxleyanus Stebbing, Rep. Voy. ‘ Challenger,’

WO Loa [OS COA, ols 33

1906. * % Stebbing, Das Tierreich, vol. xxi.

Dao

Mr. Vallentin obtained this species from a nest on Macrocystis,

Dec. 24, 1898. He also records it from Stanley Harbour, taken

by tow-net; he took it from a rock-pool, June 10, 1910, having

previously on March 11 of that year found the sea teeming

with it.

Fam. PONTOGENEIID4.

1906. Pontogeneride Stebbing, Das Tierreich, vol. xxi. p. 356.

OMS 4 Chevreux, Deuxiéme Exp. Antarct. francaise,

Amph. p. 167.

Gen. BovatuiA Pfeffer.

1888. Bovallia Pfeffer, Jahrb. Hamburg. Anst. vol. v. p. 95.

1906. ye Stebbing, Das Tierreich, vol. xxi. p. 357.

1909. 53 Chilton, Subantarct. Is. of N. Zealand, p. 622.

IT, 4 Chilton, Tr. R. Soc. Edinb. vol. xlviii. p. 494.

ils. eC Chevreux, Deuxieme Exp. Antarct. francaise,

Amph, p. 168.

In Chilton’s paper, 1912, Walker’s Husiroides orchomenipes,

1904, is accidentally cited as 1’. archomenopsis.

BovaLiiA REGIS Stebbing. (Pl. VIIT.)

Abstract P.Z.S. 1914, p. 30. (April 28.)

In the medio-dorsal structure the new species strongly

resembles L. gigantews Pfeffer, and b. walkeri (Stebbing), first

described by Walker as Atylus antarcticus. But it hasa character

not attributed to these, in that the lower borders of all the

CRUSTACEA FROM THE FALKLAND ISLANDS. 363

pereeon segments are extended laterally outwards over the side-

plates. The latter agree with those figured by Walker and

Chevreux for B. walkeri. The subacute “medio-dorsal extension

of the last pereeon segment and the first two pleon segments is

very pronounced, as is that of the third pleon segment, but this

last is distinguished from the others by its obtuseness. The

second and thir d pleon segments have the postero-lateral angles

minutely acute. The telson has an extremely short division

between the subacute apical lobes, which reach a little beyond

the peduncle of the third uropod. In both specimens examined

the lobes were slightly unsymmetrical.

The eyes are round, not crescentic as in B. giganteus, nor large

and reniform as in B. walkeri. The first antenne agree with the

former only, in having no accessory flagellum; the principal

flagellum showed short filaments on the first, second, fourth, and

seventh articulations, and so on at each successive third to the

twenty-second or twenty-eighth, the total in one specimen being

30, in another 33. The longer second antennz show a flagellum

of 46 joints, the proximal group very short, those towards the

end rather long, the whole flagellum longer by half than that of

the first pair. Hach mandible has an accessory plate, that on the

left forming five little teeth, that on the right having only two,

which are longer and apical instead of serial; the third joint

of the palp is shorter than the second. The lower lip appears

to be without inner lobes. The first maxille have four plumose

sete on the apical margin of the rather broad inner plate

second joint of the palp long.

The first and second gnathopods are extremely similar in the

female, the hand oval, narrowest at the finger-hinge, the palm

making a continuous curve with the hind margin, its limit

defined by spines which the tip of the curved finger reaches ;

hand and finger slightly larger in the first gnathopod than in the

second. The fifth perseopod has the hind margin of the second

joint sinuous, the greatest width of the joint being near its base.

The first ur opods have a peduncle much longer “than the inner

ramus, which is longer than the outer, bis shorter than the

inner ramus of the second pair, that ramus exceeding its peduncle

in length. The third uropods have the rami subequal, much

longer than their peduncle and somewhat longer than the

telson.

Length of one specimen 12 mm., that of the specimen figured

9 mm. in its bent posture, probably about 12 mm. if extended ;

it contained numerous eggs.

Locality. Low spring tide at Roy Cove, the specific name

alluding to that of the place so diligently examined by Mr. Val-

lentin.

Gen. PontoGENEIA Boeck.

1871. Pontogeneia Boeck, Forh. Selsk. Christian. 1870, p. 193.

1906. 5 Stebbing, Das Tierreich, vol. xxi. p. B59,

364 THE REY. T. R. R. STEBBING ON

PoNTOGENEIA ANTARCTICUS Chevreux.

1906. Pontogeneia antarctica Chevreux, Bull. Soc. Zool. France,

vol. xx. p. 79, text-fig. 2 aA—K.

1906. 6 os Chevreux, Exp. Antarct. francaise,

Amphip. p. 69, text-figs. 40,

4] A-K.

1909. a # Chilton, Subantarct. Is. of N. Zea-

land, Art. 26, p. 624.

1912. i Ms Chilton, dhe Rey sec. skidimib

vol. xlviii. p. 796.

1913. i a Chevreux, Deuxieme Exp. frangaise,

Amphip. p. 177, text-fig. 59 a—c.

This species by its smooth, compressed, and not dentate body

is strikingly distinguished from Bovallia regis, which in many

other points it nearly resembles.

The specimen which I am here assigning to Chevreux’s species

agrees admirably in most respects with the French author’s

figures and description, especially with the figures which he has

recently given of the male gnathopods. Of these, the first are

larger than the second, the hands in both pairs oval, with the

palm scarcely distinguished from the hind margin except by

the extent of the respective fingers. The unarmed telson, with

short division between the rounded apical lobes, is also in pre-

cise agreement. ‘The inner plate, however, of the first maxille

has only three terminal sete, instead of the four shown in

Chevreux’s figure and five mentioned in his text. Also the third

uropods have few spines instead of many, and the flagellum of

the first antenne after the first two joints has the filament-

bearing joints separated from two to two, not three to three.

These differences may well be attributed to an earlier stage in

the development, but if so, the last of them would throw doubt

on the importance which has been attributed to these intervals

in the flagellum of various specimens. In Dana’s /phimedia

simplex (from Hermite Island), which evidently belongs to this

family, the first two filament-bearing joints are the third and

sixth, but the following are the eleventh, sixteenth, and so on for

each successive fifth joint.

Locality. Stanley Harbour, among seaweed at low water of

spring tide.

Gen. AryLorpEs Stebbing.

1888. Adyloides (part) Stebbing, Rep. Voy. ‘Challenger,’ vol. xxix.

lla

1906. as Stebbing, Das Tierreich, vol. xxi. pp. 356, 362.

1909. 5 Chilton, Subantarct. Is. New Zealand, p. 627.

1912. <3 Chilton, Tr. Roy. Soc. Edinb. vol. xlviii, pt. 2.

p. 496.

Chilton, in 1909, inclines to identify this genus with Paramera

CRUSTACEA FROM THE FALKLAND ISLANDS. 365

Miers, 1875, but in 1912 he still retains it, and endows it with

a new species, A. calceolatus.

ATYLOIDES MAGELLANICUS (Stebbing).

1888. Atylopsis magellanicus Stebbing, Rep. Voy. ‘ Challenger,’

vol, xxix. p. 925, pl. 79.

1906. Pontogeneia magellanica Stebbing, Das Tierreich, vol. xxi.

p. 360.

1906. 2 a Chevreux, Exp. Antarct. francaise,

Amph. p. 64, figs. 37-39.

IDO. A ‘, Walker, Nat Antarct. Exp. vol. iii.

p. 33, pl. 12. fig. 20.

1909. Atyloides magellanica Chilton, Subantarct. Is. of N. Zealand,

p. 627.

1912. * ; Chilton, Tr. R. Soc. Edinb. vol. xlviu.

p- 496, pl. 1. fig. 18.

UG)US%, is magellanicus Chevreux, Deuxieme Exp. Antarct.

francaise, Amph. p. 178.

The transference of this well-distributed species from genus to

genus is at least some testimony that the genera concerned

belong to a single family. There is general agreement as to the

variability of the telson, to which Mr. Vallentin’s collection bears

further witness.

Locality. Whales Bay, May 17, 1910.

Gen. PARAMGRA Miers.

1875. Paramera Miers, Ann. Nat. Hist. ser. 4, vol. xvi. p. 75.

1906. 9 Stebbing, Das Tierreich, vol. xxi. p. 363.

ISIS i Chilton, Tr. R. Soc. Edinb. vol. xlviii. p. 498.

PARAMG@RA AUSTRINUS (Bate).

1862. Atylus austrinus Bate, Catal. Amph. Brit. Mus. p. 137,

pl. 26. fig. 4.

1906. Paramera austrina Stebbing, Das Tierreich, vol. xxi.

p. 363.

1909. ny re Chilton, Subantarct. Is. of N. Zea-

land, p. 625.

1912, he % Chilton, Tr. R. Soc. Edinb. vol. xlviu.

p- 498.

Among the specimens which I refer to this species, one had

calceoli on fourteen consecutive joints of the flagellum in one of

the first antenne, while in the other the calceoli were only on

alternate joints. In another specimen, calceoli were present on

alternate joints of both members of the second pair of antenne,

while on the one remaining member of the first they were, if

present, very inconspicuous.

Locality. Low water at top of Roy Cove creek, Aug. 1, 1910.

366 THE REV. T. R. R. STEBBING ON

, Fam. GAMMARID&.

1814. Gammaride Leach, Kdinb. Encycl. vol. vii. p. 432.

Gen. Meurra Leach.

1813. Melita Leach, Edinb. Encycl. vol. vii. p. 403.

1906. ,, Stebbing, Das Tierreich, vol. xxi. pp. 366, 421, 732.

MELITA IN%QUISTYLIS Dana.

1852. Amphaitoé (Melita) inequistylis (2) and A. (M.) tenwicornis

(a) Dana, P. Amer: "Acivol: it pp) 2140 il.

1906. Melita inequistylis Stebbing, Das Tierreich, vol. xxi. pp.429,

732. .

WOODS 5 Chilton, Subantarct. Is. of N. Zealand,

p. 630.

This species has been rediseussed by Dr. Chilton, who unites

with it Melita zeylanica Stebbing, 1904, from Ceylon. Mr. Val-

lentin procured a male specimen at low water in Rapid Point,

deta, AAO UII

Fam. DEXAMINIDA.

1888. Dexaminide Stebbing, Rep. Voy. ‘ Challenger,’ vol. xxix.

p. 573.

Gen. PARADEXAMINE Stebbing.

1899. Paradexamine, Stebbing, Ann. Nat. Hist. ser. 7, vol. iv.

pare:

1906. 5 Chevreux, Exp. Antarct. francaise, Amphip.

‘ [Ds 1885

1909. a Chilton, Subantarct. Is. of New Zealand,

p. 632.

WOUOr ie Stebbing, Mem. Australian Mus. vol. iv.

p- 602.

I) he Chilton, Tr. R. Soc. Edinb. vol. xlviii. pt. 2,

p. 501.

OMe # Chevreux, Deuxiéme Exp. Antarct. fran-

caise, Amphip. p. 181.

Professor Chilton proposes to make Chevreux’s P. jissicauda

a synonym of Thomson’s P. pacificus, although in the latter

species the telson is not divided to the base as it is in the former.

In 1913 Chevreux observes that his P. fissicauda is separated

from all the other known forms of the family Dexaminide by the

second maxille, the inner plate of which carries a series of spines

on the inner margin.

PARADEXAMINE NANUS Stebbing.

Abstract P. Z. 8. 1914, p. 30. (April 28.)

In point of size this species is comparable with P. flindersi,

from which it is distinguished by the differently-shaped telson

and by the greater length of the palp of the maxillipeds. The

CRUSTACEA FROM THE FALKLAND ISLANDS. 367

body is much less conspicuously dentate than in any other

described species of the genus, having a medio-dorsal tooth

extended backwards only on the second, third, and fourth pleon

segments, this tooth being flanked on the third segment by a

very small pair of additional teeth, which may be present also on

the second segment but were not perceived. The mouth-organs,

both gnathopods, uropods, and telson, are in close agreement

with those described and figured by Chevreux for P. fissicauda,

15 mm. in length. In the first maxille, however, there is only

one seta on the narrowly oval inner plate, and few sete on the

single-jointed palp. The inner margin of the inner plate of the

second maxille could not be made out. The fifth joint in the

first gnathopods is not longer than the sixth, but in the other

species the difference in length appears to be very slight. In the

fourth pereeopods the second joint has a convex hind margin, not

a sinuous one as in the species compared. The telson does not

reach the end of the third uropods, and each of its long narrow

lobes has three or four spines along its outer margin with two

unequal spinules at the apex. The flagellum of the first antennz

is composed of fourteen joints, that of the second is more slender

with nine joints; in both pairs the joints in general being con-

siderably longer than broad. Hach of the two specimens mea-

sured 2°5mm. The one dissected contained numerous eges, and,

whatever allowance is made for variability, I think it would be

scarcely reasonable to regard this matron, a tenth of an inch long,

as belonging to the same species as a congener over thirty times

her bulk.

Locality. Stanley Harbour, in seaweed at low water of spring

tide.

Fam. TALITRID&.

1906. Valitride Stebbing, Das Tierreich, vol. xxi. pp. 8, 523, 735.

1913. a G. M. Thomson, Tr. N. Zealand Inst. vol. xlv.

p. 243.

Thomson is ‘inclined to reduce Zalitrus, Talitroides, Orches-

toidea, Talorchestia, and Parorchestia to Orchestia.” But to play

the part of Saturn swallowing his children, he should have chosen

Talitrus in preference to Orchestia. Calman in 1912 agrees with

him in questioning the independence of Vulitroides.

Gen. TAuorcHestrA Dana.

1852. Talorchestia Dana, Amer. J. Sci. ser. 2, vol. xiv. p. 310.

1906. . Stebbing, Das Tierreich, vol. xxi. p. 543.

1907. ee Chevreux, Mém. Soc. Zool. France, vol. xx.

p- 495.

TALORCHESTIA SCUTIGERULUS (Dana).

1853-5. Orchestia scutigerula Dana, U.S. Expl. Exp. vol. xiii.

p- 863, pl. 58. fig. 2.

1862. es Me _ Bate, Catal. Amph. Brit. Mus.

p- 26, pl. 4. fig. 7.

368 THE REV. T. R. R. STEBBING ON

1906. Zalorchestia scutigerula Stebbing, Das Tierreich, vol. xxi.

p. 544.

1912. r 5 Chilton, Tr. Roy. Soc. Edinb.

vol. xlviil. p. 508.

Mr. Vallentin reports this species as very common along the

shore at Stanley Harbour, Nov. 20, 1898, and he obtained

numbers of smaller specimens from cast up Macrocystis on

March 21, 1902. One of the largest of these latter measured

13°5 mm. in length, and is in good correspondence with Dana’s

figure from a Tierra del Fuego specimen. But a larger specimen

from Mr. Vallentin’s earlier find measured 15 mm., and agrees

with Bate’s figure, showing the shield at the back of the second

joint of the fifth pereeopod rising above the animal’s back, over

which the two bucklers meet. ‘The difference in appearance

between the two forms is very considerable, but further com-

parison shows that it is due to the maturing of the single feature

to which it is confined.

Gen. Hyaewua S. I. Smith.

1874. Hyalella 8. I. Smith, Rep. U.S. Fish Comm. vol. ii. p. 645.

1906. 94 Stebbing, Das ‘Tierreich, vol. xxi. p. 574.

1907. ais Weckel, Pr. U.S. Mus. vol. xxxu. p. 54.

1906. re Chevreux, Lacs des hauts plat. d’Amer. du Sud,

p- | (extrait, 1907).

1910. oa Weckel, Pr. U.S. Mus. vol. xxxviil. p. 623.

1911. s4 Ortmann, Princeton Univ. Exp. Patagonia, vol. 111.

p. 650.

HYALELLA PATAGONICUS (Cunningham) Ortmann.

1871. Allorchestes patagonicus Cunningham, Tr. Linn. Soe.

London, vol. xvii. p. 498, pl. 59. fig. 4.

1888. Hyalella patagonicus Stebbing, Rep. Voy. ‘Challenger,’

vol, xxix. p. 404.

1911. Hyalella patagonico Ortmann, Princeton Univ. Exp.

Patagonia, vol. iii. p. 650, pl. 48. fig. 3 ah.

If the above identification could be proved to be erroneous by

comparison with Cunningham’s original specimen, a change in the

recent specific name would become necessary on the ground of

preoccupation. As it stands the species has been amply described

as new by Dr. Ortmann, who also mentions its near relationship

to other species of the genus. In our specimens the sixth joint

of the fifth pereeopod is relatively longer than in Ortmann’s

figure, and the third uropods agree not with his figure, but with

his text. Mr. Vallentin reports the “ colour when alive very dark

brown, almost black.”

Localities. ‘‘In sand ground near old house, Port North,

10 Aug., 1910.” Along with it were numerous specimens of the

little Copepod Boeckella michaelseni (Mrazek), of which Dr. Thomas

Scott, F.L.S., reports in the Ann. Nat. Hist. ser. 8, vol. viii.

p. 3, 1914, “This species occurred in at least eight gatherings ”

CRUSTACEA FROM THE FALKLAND ISLANDS. 369

(of Mr. Vallentin’s collection). The Hyalella was also “found in

a freshwater stream some miles distant from Stanley,” where

‘this species appeared to be fairly common.”

Fam. AORID&.

_1899. Aoride Stebbing, Ann. Nat. Hist. ser. 7, vol. iv. p. 211.

Gen. Lemsos Bate.

1857. Lembos Bate, Ann. Nat. Hist. ser. 2, vol. xix. p. 142.

1906. » Stebbing, Das Tierreich, vol. xxi. pp. 594, 737.

1909. », Walker, Tr. Linn. Soc. London, vol. xii. p. 337.

GOOF » Chilton, Subantarctic Is. of N. Zealand, p. 646.

LeMBOS FUEGIENSIS (Dana). (PI. IX.)

1853-55. Gammarus fuegiensis Dana, U.S. Expl. Exp. vol. xiii.

p-. 954, pl. 65. fig. 8 a—-h.

1862. Mera fuegiensis and M. fuegeensis Bate, Catal. Amph.

Brit. Mus. p. 194, pl. 35. fig. 4.

1906. Lembos fuegiensis Stebbing, Das Tierreich, vol. xxi. p. 600.

1909. Lembos kerguelent Walker, Tr. Linn. Soc. London, vol. xii.

p- 337, pl. 43. fig. 6.

1909. (¢) LZ. kerguelena Chilton, Subantarctic Is. of N. Zealand,

p-. 646, text-figs. 12 a, 0.

In 1906 this species remained obscure, Dana having described

and figured it only in the femalesex. While naming it fwegiensis

as if it belonged to Tierra del Fuego, he assigned it to the “ Feejee”

Islands. Now that Mr. Vallentin has obtained a male and a

female specimen together from the Falkland Islands, I feel

pretty sure that the “‘ Feejee” Islands was not the original locality,

but assigned through some lapse of memory as the rendering of

JSuegiensis, yet the distribution must be extensive, since Walker

records the species from the Indian Ocean.

The male differs from Lembos kergueleni (Stebbing), taken

from a considerable depth at Kerguelen Island, by the hand of

the first gnathopod, which has a differently sculptured palm, and

also by the second joint of the second gnathopod, which is here

not a broadly expanded oval as in the other species. The expan-

sion, however, is also absent from the specimen which Chilton, in

1909, identified with Z. kerqueleni, but that identification seems

to me very doubtful, since the male here figured is apparently

adult, to judge by the antenne and gnathopods, and the size

slightly larger than that of the accompanying ovigerous female.

It scarcely needs observing that the expansion of the second joint

of the second gnathopod, though it occurs also in Hurystheus

exsertipes, is a very unusual feature. In the present species the

second joint is not expanded either in the gnathopods or in any

of the perzeopods.

The eyes are small and round. The first antenne have a long

peduncle and longer flagellum, first joint of peduncle rather

370 THE REY. T. R. R. STEBBING ON

shorter than the second, which is four times as long as the third;

flagellum of 30 joints in the male, 26 in the female, accessory of

6 and 5 joints respectively. Second antenne with long peduncle,

last joint a little longer than the penultimate, a little shorter

than the 13-jointed flagellum.

The mouth-organs and pereeopods do not seem to offer charac-

ters of value specifically. The first gnathopods of the male are

in near agreement with the figure and description given by

Mr. A. O. Walkerin 1909. My figure was drawn before I had

realized the necessity of comparing it with Mr. Walker’s. His

description is, ‘hand three times as long as wrist, hind margin

longer than palm, which is defined by a blunt, everted tooth,

setose on the side; behind this is a large sinus followed by a

prominent flat-topped tooth ; dactylus swollen near the middle.”

In my specimen the front margin is rather longer than that in

Mr. Walker’s drawing and the blunt tooth is not everted, but

such differences may well be individual. In regard to the first

gnathopods of the female, with their slightly excavated palm,

Mr. Walker’s figure agrees fairly well with Dana’s and with mine,

which also was drawn before comparison with Dana’s species had

occurred to me as desirable. Mr. Walker speaks of the fourth

and fifth pereeopods as having “the 2nd joints wider than in the

‘Challenger’ specimen” (of LZ. kergueleni). That would not

agree with the form here in question or with Dana’s figure.

In the uropods Dana notices the long spines apical to the

peduncles of the first and second pairs. In the first pair the

rami differ slightly from Dana’s figure in being a little longer

instead of a little shorter than the peduncle, but on Dana’s plate

they are drawn in situ, which is unfavourable to minute accuracy

of measurement.

The telson is considerably longer than broad, a fact that

would scarcely be suspected from a lateral view. Length of

male in much curved position 7 mm., at full stretch probably

10 mm. or more; female (with numerous eggs) nearly as long.

Locality. Falklands, from roots of Macrocystis at 2-4 fathoms,

Jan. 14, 1902.

Fam. PHOTID &.

1872-76. Photide Boeck, Skand, Arkt. Amphip. vol. i. p. 74,

vol. 11. p. 546.

1906. a Stebbing, Das Tierreich, vol. xxi. p. 602.

Gen. Haptocurrra Haswell.

1879. Haplocheira Haswell, P. Linn. Soc. N.S. Wales, vol. iv.

p. 273.

HAPLOCHEIRA BARBIMANUS (G. M. Thomson).

1879. Gammarus barbimanus Thomson, Tr. N. Zealand Inst.

vol. xi. p. 241, pl. 10. fig. 1.

CRUSTACEA FROM THE FALKLAND ISLANDS. 371

1879. Haplocheira typica Waswell, P. Linn. Soe. N.S. Wales,

vol. iv. p. 273, pl. 11. fig. 2.

1906. H. barbimana Stebbing, Das Tierreich, vol. xxi. p. 609.

1907. 5 Walker, National Antarct. Exp. vol. iii.,

Amphip. p. 35.

1912. iy Chilton, Tr. R. Soc. Edinb. vol. xivii. p. 510.

Mr. Vallentin obtained a specimen at Stanley in seaweed at

low water of a spring tide, and others from a sponge on a

schooner.

Fam. AMPITHOID &.

1899. Ampithoide Stebbing, Ann. Nat. Hist. ser. 7, vol. iv. p. 211.

Gen. AmprrHoE Leach.

1813-14. Ampithde Leach, Edinb. Eneyel. vol. vil. pp. 408, 432.

1906. Ampithoe Stebbing, Das Tierreich, vol. xxi. p. 631.

AMPITHOE BREVIPES (Dana).

1852. Amphithoe brevipes Dana, P. Amer. Acad. vol. ii. p. 216.

1853-5. __,, tia Dana, U.S. Expl. Exp. vol. xiii. p. 941,

pl. 64. figs. 5 a2, k-n, and 1, md, m\*.

1906. Ampithoe brevipes Stebbing, Das Tierreich, vol. xxi. p. 637.

A female specimen 20 mm. in length, carrying numerous small

eggs, agrees remarkably well with Dana’s figures and description,

except that neither the first nor the second uropods have the

rami so equal in length as his figure represents, nor is the

more slender (and longer) ramus so devoid of spines. The

glandular second joint in the first and second pereopods, so

important for nest-building, is, as Dana notes, conspicuously

swollen.

Locality. Found “with their ‘nests’ made on a frond of

Macrocystis pyrifera, 1 foot from the surface,” Nov. 24, 1898.

Fam. J ASSID &.

1906. Jasside Stebbing, Das Tierreich, vol. xxi. pp. 8, 647, 739.

Gen. JAssa Leach.

1814. Jassa Leach, Edinb. Encyel. vol. vii. p. 433.

1906. ,, Stebbing, Das Tierreich, vol. xxi. pp. 652, 739.

1913. ,, Chevreux, Deuxiéme Exp. Antarct. frangaise,

Amphip. p. 181.

JASSA FALCATUS (Montagu).

1808. Cancer (Gammarus) falcatus Montagu, Trans. Linn. Soc.

; London, vol. ix. p. 100, pl. 5. fig. 2.

1906. Jassa falcata Stebbing, Das Tierreich, vol. xxi. p. 656.

OTe Fo 55 55 sexton, J. Mar. Biol. Assoc. vol. ix. p. 212.

ON Deans ,, Chilton, Tr. R. Soc. Edinb. vol. xlviii. p. 511.

Under Montagu’s specific name Dr. Chilton groups a great

Proc, Zoou. Soc.—-1914, No. XXV. 25

BY THE REV, T. R. R. STEBBING ON

variety of synonyms, including Pfeffer’s Podocerus ingens, which

attains a length of 26 mm., Walker’s Hemijassa goniamera and

Jassa wandeli Chevreux. In 1913 Chevreux points out that,

though the adult male of his species proves to have second

gnathopods very similar to those of J. falcatus, it is nevertheless

distinguished among other things by a more elongate carpus of

the first gnathopods, the comparatively narrow second joint of

the fourth and fifth perzopods in which the hind margin is

almost straight, and by the less acute apex of the telson. In all

these respects it is distinct from the little specimen here recorded,

which has a length of only 3mm.; the antennz are well furnished

with long sete, and the second joint in the third, fourth, and

fifth pereopods has a very decidedly convex margin.

Locality. Stanley Harbour, among seaweed at low water of

spring tide.

Another specimen in the collection, which J should be inclined

to identify with Jassa ingens (Pfeffer), measures 8°5 mm. in

length. It has the second joint of the very elongate second

enathopod much curved, and the tooth of the large sixth joint

irregular in shape, as described and figured by Pfeffer.

Fam. COROPHIIDS.

1888. Corophiide Stebbing, Rep. Voy. ‘Challenger,’ vol. xxix.

p. 1154.

1906. iy Stebbing, Das Tierreich, vol. xxi. pp. 8, 662,739.

Gen. Cororuium Latreille.

1806. Corophiwm Latreille, Gen. Crust. Ins. vol. 1. p. 58.

1906. 4, Stebbing, Das 'Tierreich, vol. xxi. pp. 663, 685,

(3),

gs Chevreux, Bull. Soc. Zool. France, vol. xxxiil.

p. 69.

1908.

CoroPHIUM CYLINDRICUS (Say).

1818. Podocerus cylindricus Say, J. Ac. Philad. vol. i. p. 387.

1873. Corophium cylindricwum Smith & Vervill, Rep. U.S. Fish

Comm. vol. i. p. 370.

~ 1905. i A Holmes, Bull. U.S. Bureau Fish,

vol. xxiv. p. 521, text-figs.

1905. a - Paulmier, Bull. New York Mus.,

Bull. 91, Zool. 12, p. 167, fig. 37.

1906. a Bs Stebbing, Das Tierreich, vol. xxi.

pp. 692, 740.

The figures and description of the female supplied by Dr. S. J.

Holmes leave no doubt that Mr. Vallentin’s specimens belong to

this species. Holmes gives the length as 3-4 mm. Paulmier

gives it as 5 mm., probably with reference to a male specimen

which he figures in full. The Falkland Island specimens, collected

during low water of spring tides, measure only 3 mm. It is

CRUSTACEA FROM THE FALKLAND ISLANDS. _ 373

possible that Dana’s young Corophium (?) quadriceps, 2 mm. long,

from Rio Janeiro, may be a synonym, and probable that C. con-

tractum Stimpson, 1855, from Japan, later identified by G. M.

Thomson in New Zealand waters, may likewise be another name

for Say’s widely distributed species.

Fam. PoDOCERID4.

1906. Podoceride Stebbing, Das Tierreich, vol. xxi. pp. 8, 694, 741.

TOMO: Mg Stebbing, Mem. Aorstrere Mus. vol. iv. pp. 622,

650.

Gen. PopocEerus Leach.

1814. Podocerus, Leach, Edinb. Encyel. vol. vii. p. 433.

1906. » Stebbing, Das Tierreich, vol. xxi. pp. 700, 741.

PoDOCERUS BRASILIENSIS (Dana).

1853 & 1855. Platophium brasiliense Dana, U.S. Expl. Exp.

vol. xill. p. 838, pl. 55. figs. 9 a—I.

1906. Podocerus brasiliensis Stebbing, Das Tierreich, vol. xxi.

p. 704.

In this species Dana’s figure shows the fifth joint of the second

gnathopod in the male as quite distinct from the long joint which

follows. In the example from the Falklands, as in one from the

West Indies, this fifth joint is scarcely visible, in this respect

agreeing with Podocerus mangareve Chevreux, 1907, of which the

author says that the wrist is not clearly distinct from the hand.

It is possible that the species is not a true member of the fauna

of the Falkland Islands, since Mr. Vallentin records his specimens

as obtained from mud on a hulk sunk in Stanley Harbour.

Tribe CYAMIDEA.

1852. Caprellidea Dana, Amer. Journ. Sci. ser. 2, vol. xiv. p. 307.

1906. us Stebbing, Das Tierreich, vol. xxi. p- A,

1910. Cyamidea Stebbing, Ann. 8. Afr. Mus. vol. vi. p. 464.

Fam. CAPRELLID &,

1847. Caprellide White, List of Crust. Brit. Mus. p. 91.

1910. iy Stebbing, Ann. 8. Afr. Mus. vel. vi. p. 464

(with synonymy).

Gen. CapRELLA Lamarck.

1801. Caprella Lamarck, Syst. Anim. sans Vertebres, p. 165.

CAPRELLA PENANTIS Leach.

1813. Caprella penantis Leach, Edinb. Encyel. vol. vii. p. 404.

1816. C. acutifrons Latreille, Nouvean Dict. d’Hist. Nat. ude Vv.

p- 433.

1843. C. dilatata Kroyer, Naturh. Tidsskr. vol. iv. p. 585, pl. 8.

figs. 1-9.

25*

374 THE REV. T. R. R. STEBBING ON

1871. C. dilatata Cunningham, Tr. Linn. Soc. London, vol. xxvu.

. 478.

1890. €. neuen Mayer, F. & FI. Neapel, vol xvii. pp. 50, 51.

1910. C. penantis Stebbing, Ann. 8. Afr. Mus vol. vi. p. 465.

Mr. Vallentin obtained this species from dMacrocystis after a

gale on the 16th of Jan., 1910, and on other occasions from

hydroids and fronds of Jacrocystis at the mouth of Roy Cove.

Tribe PHRONIMIDEA.

1890. Hypertidea Sars, Crustacea of Norway, vol. i. p. 5.

1906. Stebbing, Das Tierreich, vol. xxi. p. 4.

1910. Phronimidea Stebbing, Ann. 8. Afr. Mus. vol. vi. p. 473.

Fam. HYPERIID®.

1889. Hypertide Bovallius, K. Svenska Vet.-Ak. Handl. vol. xx.

INOS (i, (ace

Gen. Hyreria Latreille.

1823: Hyperia Latreille, in Desmarest, Dict. Sci. Nat. vol. xxviil.

p. 347.

HyprriA GAUDICHAUDIT Milne-Edwavrds.

1840. Hyperia gaudichaudiw Milne-Edwards, Hist. Nat. Crustacés,

Vole mip. (id:

1888. x Hh Stebbing, Rep. Voy. ‘Challenger,’

vol. xxix. p. 1394, pl. 169.

1903. Hl. garvidichaudi Walker, J. Linn. Soe. London, vol. xxix.

p- 40.

IYO i Walker, Nat. Antarct. Exp. vol. ii,

Amphip. p. 7.

The specimens were “removed from large Beroe,’ March 11,

1910.

LEPTOSTRACA.

1888. Leptostraka Claus, Arbeit. Zool. Inst. Wien, vol. vii, pt. 1,

jd,

Claus here supplies a full discussion of his own and other

views on the systematic position of this subclass, insisting strongly

on the points of agreement with the Malacostraca.

Fam. NEBALIID&.

1850. Nebaliade Baud, Brit. Entomostraca, Ray Soe., p. 31.

1896. Nebaliide Sars, Fauna Norvegie, vol. i. p. 6.

1900. 53 Stebbing, Willey’s Zool. Results, Part 5, p. 659.

1904. ‘ Die Leptostraken’ Thiele, Ergeb. der deutschen Tiefsee-

Exp. vol. viii.

LOO ers. 53 Thiele, Deutsche Stidpol. Exp. vol. ix.,

Zool espe, Olle

In the interests of carcinological phylogeny attention may

CRUSTACEA FROM THE FALKLAND ISLANDS, 375

here be called to Mr. C. D. Walcott’s remarkable account of

Middle Cambrian Crustacea (Smithson. Misc. Coll. vol. lvii. no. 6,

1912), in which he rather confusingly refers to the present group

as Phyllocarida, Nebaliacea, and Leptostraca, without clearly

indicating that he uses them as synonyms.

Gen. NEBALIA Leach.

1814. Nebalia Leach, Zool. Miscellany, vol. i. p. 99.

1888. » Claus, Arbeit. Zool. Inst. Wien, vol. viii. pt. 1,

p. 122: °

1909. » Chilton, Subantarctic Is. of N. Zealand, p. 669.

Other references coincide with those given above. It may

be noticed that Leach assigned his genus to the subclass

Malacostraca.

NEBALIA B1PES (O. Fabricius).

1780. Cancer bipes O. Fabricius, Fauna Groeulandica, no. 223.

1888. Nebalia bipes, var. chilensis Claus, Arbeit. Zool. Inst. Wien,

vol. viii. pt. 1, pp. 127, 132.

1904. V. 6. chilensis Thiele, Ergeb. der deutschen Tiefsee-Exp.

vo]. vill. p. 13, pl. 4. f. 73.

1905. NW. bipes Thiele, Deutsch. Siidpol. Exp. vol. ix., Zool. i.

p- 67.

A dissected specimen, apparently a young male, shows a rostrum

with the proportions of length to breadth, 16:6, or slightly less

than 6. The eyes have a little lateral projection. The first

antenne agree with those which Dr. Thiele describes and figures

for his Nebalia longicornis magellanica. He allows that, on the ~

whole, the southern JV. longicornis Thomson differs little from

the northern 1, bipes. Itseems to me very doubtful whether the

specific distinction can be maintained. In one of Mr. Vallentin’s

specimens the second antenne nearly reach the end of the caudal

ram.

Locality. Whales Bay, May 17, 1910.

THYROSTRACA.

1893. Thyrostraca Stebbing, History of Crustacea, Internat. Sci.

Ser vole bexave pp yO, bl ail.

1902. 5 Stebbing, Encycl. Brit. ed. 10, suppl. vol. xxxiii.

(®)) joe BU

Fam. LEPADIDA.

1851. Lepadide Darwin, Monogr. Cirrip., Ray Soc., vol. i. 1851.

Gen. Leras Linn.

1758. Lepas Linn., Syst. Nat. ed. 10, p. 667.

376 THE REV. T. R. R. STEBBING ON

LEPAS AUSTRALIS Darwin.

1851. Lepas australis Darwin, Monogr. Cirrip., Ray Soc., vol. 1.

Ps S95 pie Ethos

This species was taken in King George’s Sound, on the shore

after a gale, Sept. 15, 1910.

Fam. BALANID ®.

1854. Balanide Darwin, Monogr. Cirrip., Ray Soe., vol. ii. p. 33.

Gen. Eimintus Leach.

1825. Hlminiws Leach, Zoological Journal, vol. ii.

ELMiInius KinGu Gray.

1831. Elminius kingii Gray, Zoological Miscellany, p. 13.

1854. i » Darwin, Monogr. Cirrip., Ray Soc., vol. ii.

p. 348, pl. 11. figs. 6 a6 e.

1911. #. kingi Ortmann, Princeton Univ. Exp. Patagonia, p. 637.

- Points by which I have verified this species are the scutum

without an adductor ridge, the labrum deeply notched, with five

little teeth on each side, the mandible with four or five teeth.

Darwin speaks of the first pair of cirri as having ‘‘ one ramus

nearly twice as long as the other.” In the specimens dissected,

the difference was not so considerable. The penis was stout

except near the apex in one specimen, the thin part considerably

prolonged in the other.

Mr. Vallentin’s specimens were taken at low water of a spring

tide affixed to Mytilus edulis in Stanley Harbour. He speaks of

them as scarce.

EXPLANATION OF THE PLATES.

Prats I.

Tanais ohlini Stebbing.

n.s. Line indicating natural size of female specimen figured below.

C. Dorsal view of head and first pereon segment, with first antenne and first

gnathopods ; frontal line of head conjéctural.

Pl., urp. Dorsal view of pleon, with lett uropod.

0c., a.8., a.t. Hye, first and second antenne.

m., mx.1, mxp. Mandible, first maxilla, maxillipeds.

gn.1, gn. 2, prp.1, prp.5. First and second gnathopods, first and fifth perwopods.

urp. Right uropod.

All the separate parts are magnified to a uniform scale, except the terminal joint

of the fitth perwopod, which is further given in higher magnification.

Prats II.

Exospheroma calcareus (Dana).

m.s. Lines indicating natural size of male specimen figured. below in dorsal aspect.

C.V. Ventral view of cephalon with side-plates of first peraon segment, to give an

idea of the epistome and first and second antenne in position.

plp.1. The first pleopod, along with the male organs on the last perewon segment.

plp. 2,3, 4,5. Second, third, fourth, and fifth pleopods, with apical parts of fourth

and fifth more highly magnified.

CRUSTACEA FROM THE FALKLAND ISLANDS. 377

Puate III.

Tryphosites chevreuxi Stebbing.

n.s. Line indicating natural size of male specimen figured below.

a.s., a.%. First antenna and part of the second.

gv.1, gn.2. First gnathopod with distal portion more highly magnified, and second

gnathopod.

prp. 2, 3,5. Second and third pereopods, the latter without the branchial vesicle ;

fifth pereopod, with second joint only partially figured for economy of

space.

urp.2, urp.3, T. Second and third uropods; telson in dorsal view, with higher

maguitication of the apex of the lelt division.

The separate parts are enlarged to a uniform scale, with additional enlargements

of the first gnathopod and the telson.

Prats IV.

Pariphimedia normani (Cunningham).

n.s. Line indicating natural size of female specimen figured below.

C. Partial side view of head.

l.s., l.s’. Upper and lower lips.

m., me. 1, mx.2. Mandible, first and second maxille.

gu.1, gn.2. First and second gnathopods.

All the separate parts, except the head, are magnified to a unitorm scale. The

marsupial plate of the second guathopod is omitted.

Puate V.

Pariphimedia normani (Cunningham).

mep. Maxillipeds.

prp.2, 3,5. Second and third pereeopods incomplete, and fifth pereeopod.

urp.1. First uropod, the peduncle incomplete.

urp. 2, urp. 3, T. Dorsal view of second and third uropods, with the telson.

For economy of space the perzeopods are given on a lower scale of magnification.

The other parts are uniform with those of the preceding Plate.

PuateE VI.

Monoculopsis vallentini Stebbing.

n.s. Line indicating natural size of the specimen figured below. The figure is partly

schematic, as prior to dissection details of the crowded overlapping limbs

could neither be clearly seen nor satistactorily represented.

a.d., a.i. First and second antenne.

1.s., mx.1, maz. 2. Upper lip, first and second maxille.

gn. 1, gn.2. First and second gnathopods.

Puate VII.

Monoculopsis vallentini Stebbing.

m., 1.i. Mandible and lower lip. :

prp. 2, 3,4,5. Second, third, fourth, and fifth perseopods. The branchial vesicle and

marsupial plate of the second perzeopod are omitted.

plp. A pleopod. ; :

urp. 1,2, 8,3. The right first uropod, the left second uropod, the pair of third uropods,

with the peduncle only of the left first and the right second. The telson

appears as if it were attached to the fifth segment of the pleon, but is really

attached high up on the sixth which it overlaps.

In this and the preceding Plate the mouth-organs are more highly magnified than

the other parts, but each set is on a uniform scale.

(sy)

ie)

ON CRUSTACEA FROM THE FALKLAND ISLANDS.

Pruate VIII. ;

Bovallia reyis Stebbing.

a.s. Line indicating natural size of specimen figured below in curved position, some

of the limbs omitted to prevent contusion.

as. First antenna.

1.i., m., m. Lower lip, the two mandibles.

gn.1, gn.2, prp.5. First and second gnathopods, and part of fifth perseopod.

urp.1, urp.2. First and second uropods.

T.,urp.3. Telson and third uropod.

The mouth-organs are more highly magnified than the other parts. The limbs,

uropods and telson are on a uniform scale, but the limbs and mouth-organs are not

from the specimen figured as a whole, though of one approximately of the same

S1Z€.

Prats IX,

Lembos fuegiensis (Dana).

n.s.@. Line indicating natural size of male specimen figured below in lateral view.

a.s.,a.i. First and part of second antenne.

1.i.9. Lower lip of female.

gn.1, gn.2, gn.12,gn.29. First and second gnathopods of the male and of the

female.

urp.1, wrp.2. First and second uropods.

urp.8, I’. Sixth pleon segment with telson and third uropods attached, m dorsal

view.

All the parts are drawn to a uniform scale, and all are from the male specimen,

except those with the sign ¢.

ON THE ANATOMY OF THE OPHIDIA. 379

25. Further Contributions to the Anatomy of the Ophidia.

By Josnpa C. THompson, Surgeon, U.S. Navy *.

Received October 22, 1913: Read April 21, 1914. |

INDEX.

Anatomy : Page

Methods employed ...........0.....0.......... 879

List of species examined ....................... 9380

Meruops EMPLOYED.

A routine procedure is needed for recording and correlating

observations nade on the anatomy of serpents. In comparative

anatomical studies of mammals, the position of an organ is given

in terms of its relation to other viscera. With animals of a

compact build this is amply sufficient ; with serpents on the other

hand, owing to their attenuated bodies, it is not adequate to state

that one organ lies posteriorly to another. This drawing out of

the body is accompanied by much variation as to the relative

lengths of different organs and their distances apart. For

example, the tip of the liver may be situated the distance of a

dozen vertebre behind the apex of the heart, with a thick-walled

right lung filling the space between them, or it may overlap the

apex of the heart by several millimetres, and the lung be a mere

air-sac dorsal to the liver. The length of certain organs varies to

a degree unknown in any other Order. For example, the right

lung may terminate a few millimetres posterior to the end of the

liver, or extend nearly to the cloaca.

If the gastrostege level at which the various structures occur

is recorded, one may obtain data satisfactory for comparing

different species or for determining the range of variation in

a series. ‘lake for example the data relating to ZThammnophis

ordinoides: the apex of the heart is recorded as being at the 26th,

the anterior tip of the liver at the 70th, and the posterior end at

the 86th gastrostege. It may be seen at a glance that the liver

is nine gastrosteges posterior to the heart and that it is of a

definite length, extending over thirty-five shields. For com-

paring series it will be found of advantage to reduce the records

to a percentage. In each case the number of gastrosteges in the

specimen is to be taken as the base.

Returning to Z. ordinoides: the number of gastrosteges in an

example of that species is 157, and the position of the heart or liver

in terms of its distance down the spinal column is as follows :—

Heart apex 16°6 per cent.; liver tip 22°3 per cent.; liver end

44-6 per cent. Behind the posterior pair of geneials there are

usually one or two pairs of small gular shields ; these are followed

by from one to three shields in the median line which increase

* Communicated by Dr. F. E. Beppsarp, M.A., F.R.S.,-F.Z.8. :

+ To avoid frequent repetition the term gastrostege will be abbreviated to “ g

380 SURG. J. C. THOMPSON ON THE

in width in pyramidal fashion. ‘The shield regarded as the first

gastrostege is the one that is nearly the standard width; it is

frequently distinguished by having a colour similar to the rest of

the ventrals and not white or cream like the throat.

List oF SPECIES EXAMINED.

Xenopeltis unicolor, p. 380.

Polyodontophis geninatus, p. 380.

Polyodontophis collaris, p. 381.

Thamnophis ordinordes, p. 381.

Tropidonotus vibakart, p. 384.

Tropidonotus sauteri, p. 385.

Pseudoxenodon sinensis, p. 385.

Zamenis constrictor, p. 386.

Zamenis rhodorhacis, p. 387.

Zamens florulentus, p. 389.

Coluber oxycephalus, p. 389.

Philothamnus semivariegatus, p. 394.

Cerberus rhynchops, p. 394.

Leptodira hotambeia, p. 397.

Psammophis sibilans, p. 398.

Ophiophagus bungarus, p. 398.

Doliophis bivirgatus, p. 400.

Causus rhombeatus, p. 401.

XENOPELTIS UNICOLOR Reinwavrdt.

Specimen.—California Acad. Sci. No. 16750. Singapore.

Adult female: total length 480, tail 58 mm.

The vertebral artery arises from the right aortic arch at the

AD5th g. and courses superficially to the 12th g. It gives off an

intercostal branch to each vertebra, and these enter in the

median line. The right aorta is nearly the same diameter as the

left. It gives off eight intercostal branches; each arises as a

single trunk which bifurcates midway to the parieties, and the

forks enter on either side of the median line. The dorsal aorta

begins at the 52nd g. The intercostal branches arise anteriorly

as a single trunk which upon reaching the parieties bifurcates,

one fork entering on each side. Before the gall-bladder these

bifurcate nearer to their origin, and behind this organ they are

symmetrically paired. The right kidney receives two renal

arteries; the first enters the second lobe, and the second the sixth

lobe. The left kidney receives one artery which enters at the

second lobe.

PoLYODONTOPHIS GEMINATUS Boie.

Specimen. — California Acad. Sci. No. 16751. Botanical

Gardens, Singapore. Adult male.

The trachea enters the lung at the level of the 36th g.; it

ANATOMY OF THE OPHIDIA. 381

terminates two gastrosteges further on, and is without an intra-

pulmonary prolongation. The tracheal membrane is bare to a

trifle above the base of the heart,at which point it acquires a thin

lining of shallow-celled respiratory tissue. The right lung is lined

with air-cells to the level of the 55th g. There is a free apex

reaching forward for the distance of two gastrosteges and in

relation with the right side of the trachea. The left lung is

entirely suppressed. The heart apea is at the 38th, and the tip

of the liver at the 50th g.

There are 144 gastrosteges. The scales are in 17 rows anteriorly

and the same posteriorly. — .

PoLYODONTOPHIS COLLARIS Gray.

Specimen.—California Acad. Sci. No. 26783. Mo Kan Shan,

Chekiang Prov., China. Adult male.

The tracheal membrane begins to be lined with respiratory

tissue a little before the base of the heart. The trachea enters

the lung at the level of the 38th g. The right lung is lined

with respiratory tissue to the 60th g. There is a free apex

3 mm. long in relation with the right side of the trachea.

The lefé lung is entirely suppressed. The heart apex is at the

level of the 36th and the tip of the liver at the 54th g.

This is an unusually long distance between these organs, as the

average of a Colubrine serpent is about ten gastrosteges, The

points of all the teeth on the maxillary, palatine, and pterygoid

bones are brown. The mandibular teeth are not in the normal

alignment ; those in the anterior third of the series are abruptly

faulted inwards, so that they are set in about ‘7 mm. from the

line of the remaining teeth.

There are 172 gastrosteges. ‘The scales are in 17 rows anteriorly

and the same posteriorly.

THAMNOPHIS ORDINOIDES Baird & Girard *.

The salient character is the range of variations that occurs in the

number of scale-rows. A series of over one hundred specimens

has been studied, and the following eight formule recorded :—

21-19-17, that is 21 rows forward, decreasing to 19, and further

along to 17; 19-21-19-17, that is 19 rows forward, increasing to

21 at about the level of the heart and continuing to the end of

the liver, then decreasing to 19, and further on to 17; 19-17;

19-17-15; 19-17-15-13 (13 is the lowest number of scale-rows

on the body that has been recorded in Zhammophis); 17-19-17—

15; 17-15; 15-17-15; these lower counts prevail in specimens

from the State of Oregon.

* According to the classification of the North American Garter-snakes as set

forth by A. G. Ruthven, Esq., in his memoir, “ Variations and Genetic Relationships

of the Garter-snakes,” 1908, Bulletin 61, U.S. National Museum, this species

includes Tropidonotus leptocephalus (B. & G.) and T. vagrans (B. & G.).

382 _ SURG. J. C. THOMPSON ON THE

Specimen. — U.S. National Museum No. 504006. Sausalito

Peninsula, California. Adult female.

The tracheal membrane begins at the glottis; anteriorly it lies

along the dorsal and right quadrant of the tube, and further on,

at about the 16th gastrostege, it gradually winds to the dorsal

quadrant. Anteriorly it is narrow and the ends of the carti-

laginous rings are in apposition; at the 6th g. it is 1 mm. wide

and begins to be lined with alveolar tissue; where the trachea

enters the lung the membrane is 3°D mm. wide and its alveolar

lining becomes confluent. with the respiratory tissue of the lung.

The air-cells on the membrane near the end are smaller than

those lining the anterior portion of the lung. The trachea

enters the lung at the 24th g. From without it appears to

terminate in an obtuse angle a few millimetres posterior to the

base of the rudimentary lung ; from within it is seen to continue |

along the ventral surface of the lung as a narrow, straight, and

shallow gutter of fibrous tissue which reaches nearly. to the

anangious portion. The right lung is lined with pulmonary

tissue to about the 40th g.; from this point on the air-cells

become larger, more irregular, and deeper; they may be said to

terminate, and the anangious portion to begin, at about the 45th ¢

From this poimt the lung continues as an air-sac to the level of

the 106th g.

There is a free apex: extending forward 8 mm. in relation with

the right side of the trachea; this apex opens into the lung by a

restricted aperture, 2°5 sam. in diameter, which is just aritenion

to the opening in the rudimentary lung. The rudimentary left

lung is 7 mm. long, lined with air-cells, and opens into the trachea

at a point 4°5 mm. anterior to where the trachea terminates in

the right lung.

The vertebral artery vises from the right aortic arch at the level

of the 20th gastrostege and is superficial in its course to the

10th g. It gives off five intercostal branches: I-20; IJ.-16;

T11.-15; 1V.-14; V.—-12; and one branch to the esophagus at

the 14th g. The right aorta gives off a single intercostal branch

close to its junction with the left; this branch arises at the

29th g., courses upwards and forwards and enters at the 27th g.

The left aorta is only a trifle larger than the right and is without

branches. The dorsal aorta begins at the 29th g.

There are sixty intercostals, and all but Nos. LAVAL: and LIX.

enter in the mid-dorsal line. The gastrostege level at which

they leave the aorta is as follows:—I.-31; I1.-35; III.—41;

LV.-44; V.-50; VI.-53; VIL-55; VIII-—57; 1X.-60; X.-62;

XI.-64; XIT.-66; XIIT.-68; XIV.-72; XV.-73; XVI-75;

XVIIL-77 ; XVIII.-79; XTX.-82); XX.—84; XXI.-86; XXIT—

88; XXIIT.-90; XXIV.-92; XXV.-93; XXVI.-95; XXVIT.—

96; XXVITI.-99;° XXIX-101; XXX.-103; XXXI.-104;

XXXIT.-106; XXXTIT-108; XXXIV.-110; XXXV.-111;

RRRV I A1I2 5 XXX VAT 3) 5, ROO Ti OXOS RES

116; XL-117; XLI-119; XLIL-121; XLISI.-122 ; XLIV.—

_ ANATOMY OF THE OPHIDIA: 383

193; XLV.-126; XLVE-128; XLVII-129; XLVIEL-131,;

NID LISS = ID NEGe IMLS e JIN sie INDE AUT

WAGs TN TS) S TUL = WIS ENT VG

153; LX—-156. .

The courses of the intercostals vary slightly in different parts of

the body. All but the last, LX.—156, enter the parieties anterior

to their points of origin. ‘Anteriorly the difference is only a

millimetre or so; at the posterior end of the liver it has in-

ereased to an average of three to four millimetres, equivalent to

the length of one vertebra. In the region of the kidneys each

artery runs forward the distance of about two vertebr before

entering the body-wall. These latter branches are crossed by the

renal arteries; the blood in the intercostal arteries flowing

forward and in the renal arteries flowing backward. This

arrangement is what might be expected to result from a

lengthening of the body in this region. As the kidneys move to

the rear the arteries entering them would be drawn out to the

rear. As the dorsal aorta is drawn to the rear there is a tendency

for the intercostal branches to maintain their former points of

entrance ; to accomplish this necessitates that they course for-

ward. A few are peculiar in the, following details :—I.-31 is the

longest, measuring 11 mm. ; IL.-35 bifurecates, sending branches

to the lung and cesophagus; III.-41, XX.-86, and XXI—-95

bifurcate and send one fork to the lung: LVIII.-152 enters

9-7 mm. to the left of the median line, and forks just before

reaching the parieties; LIX.—153 enters a similar distance to the

right ; LX.-156 is the only one that enters the parieties at the

same level as its origin, all the others trending forward to a

greater or lesser extent.

The gastric artery is at the level of the 81st g.; upon reaching

the stomach it bifurcates into anterior and posterior branches,

the latter a trifle larger. There are three mesenteric arteries :

1-119; 11.99; I1I1.-108. There are ten intestinal, rectal, and

cloacal arteries; 1-113 is 32 mm. long; II.-119; II1.-129;

VAIS OE MVE aS Ore Wale 4 5) Velie ol Wel 52 5 xe 153;

X.-155; the majority of these are long straight vessels.

There are four renal arteries on the right and seven on the left

side. The right kidney extends from the 116th to the 139th g.,

and its four arteries leave the aorta as follows :—I.-118 ; I1.-126 ;

TI1.-129; IV.-135. All of these enter the organ posterior to

their origin. No. I. is the largest; at the kidney it divides into

an anterior branch which forks to supply the kidney and the

ovary, and a posterior branch to the kidney alone. The left

kidney extends from the 125th to the 145th g., and is practically

the same size as its mate. The seven arteries leave the aorta

Aeifollows et 122—) 11 IAT Vesa VE 39 5

VI.-141; VII.-143. No. I. is the largest and forks to supply

the ovary. Nos. I. to VI. are quite oblique; Nos. V. to Vil.

enter the kidney almost directly.

384 SURG. J. C. THOMPSON ON THE

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :—

Gastroste ces mentees. versus 157 100 per cent.

Hearst apexursatce tesa. chiag 26 16°8

TA Viers MUN waeme ce oc aio hibe «tite ay 30 22°3

said ROI Gren ete oc uaiiehcssnes 70 44-6

UE eval IOie) ga Redanedlopeoonbeosr 86 54:8

IKercliveneiacle iii bl) pasate eee Sela 73°8

a BN CAG GUE tiie en aeRO 139 88°6

Pee We Lbs CLD seers lia acs 125 79°6

- gt OW et emen ce Sok 145 92:3

The number of scale-rows on the body, the sequence in which

they are added and suppressed, and the gastrostege level at which

these changes occur may be thus presented :—

Neck.

21 rows, IV row ends, right 8th, left 7th gastrostege, leaving :

Body.

19 rows, V row added, right 26th, left 24th gastrostege, making:

2 aoe A Ba LUC IS gm COA Drove ‘6 leaving :

Tg centile Vo Manet Usa Be Sihdae, ” Go SOila

17 +4, ~+~which are continued to the vent.

99 39

An inspection of this table will show that on the body the

scale-row count is raised from 19-21 rows at about the level

of the base of the heart, and that this maximum number of rows

continues to the posterior end of the liver, where the original

count of 19 rows is resumed. A further tapering of the body is

indicated by the reduction of the 19 rows to 17, and this occurs

just posterior to the gall-bladder. The relation existing between

the position of the viscera and the added and suppressed scale-

rows has been studied in fifty specimens of this species. The

example just recorded offers the normal condition for individuals

inhabiting the Sausalito Peninsula.

TROPIDONOTUS VIBAKARI Bole.

The salient character of this species is the peculiar type of

intromittent organ. This has warranted the removal of the

species from Z'ropidonotus and the establishment of a new genus

Hebius*.

Specimen.—California Acad. Sci. No. 15861. Yokohama,

Japan. Adult female: total length 644, tail 140 mm.

The trachea ceases abruptly at a point just posterior to the

apex of the heart. The tracheal membrane begins at the glottis

and extends along the right side of the tube; from the origin

to the 15th g. it is finely plicate; it then becomes lined with

respiratory tissue, the alveoli being at first very shallow and

indistinct ; at the level of the auriculo-ventricular septum the

* Thompson, P. Z, 8. 1913, p. 424.

ANATOMY OF THE OPHIDIA. 385

shallow cells change rather suddenly to the type of those in the

lung. The trachea ceases abruptly at a point just posterior to

the heart apex. The right lung is lined with pulmonary tissue

to about the 44th g. There is a free apex 1-5 mm. in lengt).

There is a rudimentary left lung, a mere sac without air-cells or

an opening into the trachea.

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :—

Gastroste ges) ey ecsewet cen ess 147 100 per cent.

inlieambhape xeaeeebe ses 2 ome cen 29 19:5

JRIGRETSS WYO)» Beoecins soneM tare Ae? 36 25:7,

Bhi t) MEYOGL Acie ores ree ee 64 50°7

GalllEbladder gees. 5-210: 76 55-7

Tk rainy, TEMG, 18)0) coogsosancsacc= 119 76°5

ssp datOT aN lita Sac ee he Hitt 128 87

= Vente bi wes teense 126 82°7

. shi ACTING SA a aati sin ad Laz 93

ihe -czecalllavall Vie rsecon cree cite 128 85:5

There are 19 scale-rows anteriorly ; at the level of the gall-

bladder the IV row is dropped, leaving 17 rows which are

continued to the vent.

TROPIDONOTUS SAUTERI Boulenger.

Specimen.—California Acad. Sci. No. 18988. Kanshirei,

South Formosa. Adult female.

The tracheal membrane begins to be lined with respiratory

tissue a little anterior to the base of the heart. The trachea

enters the lung at the level of the 24th gastrostege. The right

lung is lined with respiratory tissue to the 55th g. There is

a free apex about one and one-half gastrosteges long, in rela-

tion with the right side of the trachea. The rudimentary /ef¢

lung is at the 25th g.: it is a mere fibrous sac, 2 mm. long,

without pulmonary tissue. The heart apex is at the 24th, and

the tip of the liver at the 32nd g.

Thereare 17 rows of scales anteriorly and the same posteriorly.

Gastrosteges 128.

PSEUDOXENODON SINENSIS Boulenger.

Specimen.—Brit. Mus. Nat. Hist. Ex. No. 713-18. Yunnan

Fu, China. Adult male.

The tissues in this specimen are in too fragile a state to permit

of a satisfactory autopsy. The tracheal membrane is developed as a

broad thin sheet; the air-cells begin at the 28th g. The carti-

laginous rings are broader and thinner than in most species.

There is a cervical set of air-chambers above and to the right of

the trachea ; these are divided into about seven compartments by

transverse partitions. Each compartment opens into the trachea

by a large round foramen situated on the tracheal membrane;

these are opposite the following gastrosteges: 3, 7, 11, 15, 19,

386 SURG. J. GC. THOMPSON ON THE

23,25. The right lung has a large free apex. ‘There is no trace:

of the left lung.

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :—

GastLOsteses) Wesens.c cudnt ee a0 140 100 per cent.

IFlicai Grameen in aici el Cao oa. 30 21°4

UG Ive rem DU meetce eet hese. ote tcins 40 28°6

ny (EN 00 ba ak Bn a 66 42°8

Gall-bladder. 2.01.02. .s2sne:sn2c22 75 53:6

Kidweyarieht, tip 2...t2.c0c--2- 98 67:2

Hi see MOIMGEsc et) teats as 0 120 85:7

*, lefitstioy tenes seeeeee IAL 19°2

4; s/s CUNCS etc tera er 128 ies}

The number of scale-rows, the sequence in which they are

suppressed, and the gastrostege level at which they terminate

are as follows :—

19 rows, IV row ends, right 68th, left 71st gastrostege, leaving:

17 99 Vi 99 99 Pie] 81st, bP) 81st 9 27

15 which are continued to the vent.

ZAMENIS ConstTRiIcToR Linnzus.

Specimen.—_U.S. National Museum No. 504301. Sausalito

Peninsula, California. Aduit female.

The tracheal membrane begins at the glottis and extends along

the left quadrant of the tube. It is bare anteriorly but beconies

lined with respiratory tissue at the 15th g.; at first the air-cells

are large and shallow, further on assuming the type of those in

the lung. The trachea enters the lung at the 34th g.; a few

millimetres posterior to the rudimentary lung it tapers to a

narrow fibrous intrapulmonary band, less than 1 mm. wide,

which continues along the ventral wall of the lung to the

anangious part. The intercartilaginous fibrous portion of the

trachea is pigmented a dark grey, which is in sharp contrast to

the white of the cartilages and gives the trachea a regularly

banded appearance. There is a free apex 5 mm. long, closely

bound to the right side of the trachea, and with two small

openings into the lung. The right lung is lined with alveoli to

the 54th g., where they cease rather abruptly and the lung

becomes anangious. The left lung, 2°5 mm. long, is at the

36th g.; it is lined with air-cells and opens into the trachea.

The vertebral artery courses superficially over seven gastro-

steges; it arises from the right aortic arch at the 30th g. and

enters in the median line at the 23rd g._ It gives off two inter-

costal branches: I.-28, I1.-26, and two minute twigs to the

esophagus: I.-27, I1.-25. In another specimen (No. 504043,.

same locality and sex) having the same number of gastrosteges

and with the heart at the same level, this artery courses over

nine gastrosteges and gives off six intercostal branches: L-28 ;

ANATOMY OF THE OPHIDIA. 387

Te 6 Ven NE 22 Vil 2 and) one: to the

cesophagus: I.-23.

The eaternal landmarks of the principal viscera in terms of

gastrosteges are as follows :—

CARTTOSHEROS pabodnassednes5060000 168 100 per cent.

JBLGATA AY DEK a coocecsn0b spsoncgoo02ee 30 20°9

JLANTEIE HID” \pacosodondoacevoeongdane AT 28

55) hang S1OK leas Sam coenGs SacweepEE ose 77 45°8

Gallebladders repent etc. 99 59

TR GlINi 7 TMEANG, GN) aeceonoobeocce 135 80-4

ue Wee MING ee ay a ciaacacadels 155 92°2

ss Webb satuon ones e soce 142 *84°5

LA Ca Bigg 1010 bag SPR Seat me 160 95:2

The parietal plewra is pigmented along an irregular line on the

flanks. The parietal peritonewm is darker and the pigment is

more diffused until at the level of the pancreas it is quite black.

The visceral peritoneum covering the liver is slightly darkened.

The number of scale-rows on the body, the sequence in which

they become suppressed, and the gastrostege level at which they

terminate may be presented thus :—

Neck.

19 rows, IV row ends, right 5th, left 136th gastrostege, leaving :

Body.

17 rows, IV row ends, right 94th, left 95th gastrostege, leaving :

15 ,, which are continued to the vent.

ZAMENIS RHODORHACIS Jan.

The salient characters are:—The sequence of suppression of

the scale-rows which is first No. IV, then IX, and finally

VII ;—over one hundred species of Colubrine snakes have been

examined for this character, and this is the only one recorded

with this formula. The caudad position of the heart. The

attenuated liver. The of distance the gall-bladder from the

liver,—fifteen gastrosteges, while it 1s usually ten in a Colubrine.

Specimen._Brit. Mus. Nat. Hist. Ex. No. ’13-7a. Aden,

Arabia. Adult male.

The tracheal membrane lies along the right side of the tube

from the glottis to a little before the base of the heart where it

becomes dorsal; it is narrow throughout and lies in a lax state,

the interrupted ends of the cartilaginous semirings being in

apposition ; it is not attached to the extremities of the carti-

lages, but along a line about °5 mm. distant. The aiv-cells begin

at about the 55th g. The trachea ends abruptly at the 62nd ¢.,

there being no trace of an intrapulmonary bronchus. ‘The left

lung is completely suppressed. The right lung loses the lining

of air-cells at the 77th g. Thereis an apex 3 mm. long, attached

to the left side of the trachea and communicating with the lung

by a restricted opening.

Proc. Zoou. Soc.—1914, No. XX VI. 26

388 SURG. J. C. THOMPSON ON THE

The heart is situated 27°6 per cent. down the vertebral column ;

this is a little further to the rear than is usual in a Colubrine

snake. The liver is much attenuated both anteriorly and pos-

teriorly. The parietal plewra is very dark, the parietal peritoneum

is almost black, and the visceral coverings are unpigmented.

The vertebral artery rises from the right aortic arch at the

54th g., courses superficially to the 44th g., and enters in the

median line. There are four intercostal branches: I-53;

I1.—52; I11.-50; IV.-47. The right aorta has one large inter-

costal branch, 1.60, which enters to the left of the median line.

The dorsal aorta begins at the 62nd g. The intercostal arteries

enter the left side at the following gastrostege levels: 1-70;

172 wif > VV 3 Vee Vil=84 Vv M85 Ve oie

TX.-89 ; X.-92; XI.-95; the branches beyond are injured.

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :—

Gasbrosbe Ses Ui eann.c saan ee 221 100 per cent.

IRIGENED EY NEG od sunoossGsconoacepaese 61 27°6

Tei Ver MitO ye eae crmaru tan: 66 29°8

aE Mea Vee ea Be UTR ae ani 109 49-5

Gall-bladder ..... Dersatte eG 134 60-4

TRESS, TBIEAN TINO! a spcossrocasee 160 (ARS

Ke neha PCI ask Aaa eae 167 75:8

Ne Netti bi wane es eee oe 172 77°8

as MEERCTNG pce ena nae aye nee 179 81-0

Keine ye avelity sblp ees seer errr: 184 84:2

ss remainder injured.

The number of scale-rows, the sequence in which they are

suppressed, and the gastrostege level at which they terminate 1s

as follows :—

Neck.

21 rows, IV row ends, right 7th, left 6th gastrostege, leaving :

Body.

19 rows, IV row ends, right 129th, left 128th gastrostege, leaving :

Vite LEX peo eels 5 Weal a

ANS yan gi eA Al GLa er as 5 albiist ee a lost 4s

13 ,, which are continued to the vent.

The bodies of most serpents taper posteriorly and this is

usually associated with the loss of definite scale-rows. The rows

that are suppressed are prone to be either on the middle of the

flanks or one or two rows distant from the median. When the

rows on the flanks are dropped the levels at which they terminate

are usually bilaterally asymmetrical, there being a difference of

from one to four gastrosteges. When the rows adjoining the

median are suppressed, the rule is for them to terminate at the

same level. ‘he pioneer work in this field has been done by

A. G. Ruthven, Esq. Since the days of Cope, no American

worker has given so astute a suggestion to students of herpetology

as is contained in his memoir on the genus Zhamnophis.

)

ANATOMY OF THE OPHIDIA. 389

ZAMENIS FLORULENTUS Geoft.

Specimen.—Brit. Mus. Nat. Hist. Ex. No. ’13-5. Blue Nile,

Soudan. Adult male.

The tracheal membrane lies on the left side of the tube, and is

narrow, being 1°5 mm. wide at the base of the heart. The air-

cells begin at the 40th g. The trachea terminates at the 50th g.

The remnant of the bronchus continues as a fine intrapulmonary

streak of fibrous tissue as far as the anangious region. The right

lung is without a free apex ; the air-cells extend to the 66th g.

A vestige of the left lung is present at the 50th g.; it is without

air-cells or an opening into the trachea.

The vertebral artery leaves the right arch of the aorta at the

43rd g. and enters the parieties at the 35th. It has four inter-

costal branches which enter in the median line: I.-43; II1.—41;

I11.-39; IV.-37. The right aorta extends from the 43rd to the

5ist g. It has two intercostal branches: I.—45; I1.-49.

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :—

(CASIROSIOHES Sentecssaccbncneenace 197 100 per cent.

Jal@nien, GOSS Fouaasys WEA W nent 48 24-4

NGI VET UNM a acne ce lasccmes: 56 28°5

cto MUGEN gk ts 3 er eta ONS SAIN ae 96 48-7

Gall-bladder ............. Cotscee 124 62:9

ARSSeTS, SAElANG, WOb secucc ssc doodboo- 144 Ce

=H iel(s hac maya nek eine rca ear 148 75°2

5) Wiel esas Nail ah gaaae mee meen 158 80:2

3 Pos) 06 IR geen er Een Ac 162 82°3

TAGhNe iy, TENE NES Hl Oso sdneoaden asc 166 84:2

“3 PSR HCSi 010 RSE es 179 90°9

5 Neht sulpaeessee acre Way 90

- Basra Wie eea sane ciao 189 96

The number of scale-rows, the sequence in which they are

suppressed, and the gastrostege level at which they terminate, are

as follows :—

Neck.

23 rows, LV row ends, right 11th, left 10th gastrostege, leaving:

Body.

2lrows, IV row ends, right 120th, left 120th gastrostege, leaving:

IGG) easel ea] 2 Bl ae teen “ LePial, Wezel

Tene ee D Gia - Wiatiecl, . USahec!

15 ,, which are continued to the vent.

9? 9?

99 9)

CoLUBER OXYCEPHALUS Bole.

The salient characters of this serpent are as follows:—The

system of air-chambers opening into the trachea. The length of

the right lung and the considerable development of the left; both

lungs are larger than in any Colubrine as yet recorded. The

intercostal branches of the vertebral artery do not all enter in the

26*

390 SURG. J. C. THOMPSON ON THE

median line. The highly differentiated intromittent organ.

Finally the ugly and fierce disposition.

Specimen.—Brit. Mus. Nat. Hist. Ex. No. 713-17 a. Bala-

kappan, Dutch Borneo. Adult male.

The trachea terminates abruptly at the 65th g. There is no

trace of an intrapulmonary bronchus, the last vestige, the linear

arrangement of the air-cells even being absent. The intercarti-

laginous membrane is black, and in sharp contrast to the white of

the cartilages. Hach end of a cartilage where the ring is in-

terrupted by the membrane is free to the extent of -5 mm.; this

gives a comb-like appearance to the line of junction of the

cartilaginous and membranous portion. The tracheal membrane —

is not extensively developed, being uniformly about 2 to 3 mm.,

and at the base of the heart 4 mm. wide. Anteriorly it lies

along the right and posteriorly along the dorsal side of the

trachea. It is bare to the 49th g., where it acquires a lining of

air-cells that posteriorly become continuous with those of the

lung.

The body-cavity beneath the trachea from the level of the

paired gular shields to the base of the heart is converted into an

air-chamber. This structure gives the impression of crowding

and filling all the available space. The walls are of stout

connective tissue and are closely bound to the surrounding

structures. The air-chamber is divided into fifteen separate com-

partments by connective tissue bulkheads that are imperforate.

The position of the posterior wall of each compartment in terms

of gastrosteges is as follows:—I.-3; IT—8; IT1.-13; [V.—18;

V.-23; VI-26; VII-31; VIII-36; IX.-39; .X.-42;

X44 = OX 49 = X= 52 XT VR a4 = XeV 565 “Thesaiver

compartment is in relation laterally with the vena cava and the

carotid artery and posteriorly with the base of the heart; it even

sends a prolongation for a few millimetres along the ventral surface

of the pericardium. Compartments XII. to XV. cease to be

attached to the parieties and lie between the vessels of the neck

a the trachea. Hach compartment opens into the trachea by an

aperture ; in several cases there are two and even three openings.

These apertures are situated on the tracheal membrane, close to

the ventral row of cartilage-ends. They are oval, with the long

axis in the sagittal plane, from 2 to 3°5 mm. in diameter, and

extend over the space of several tracheal cartilages. Where they

exist, the cartilages terminate flush with the rim of the opening,

and do not have the free extremity that is present elsewhere.

The first structure of this nature was discovered by Dr. Beddard *

in Ophiophagus. In the Hamadryad it is dorsal to the trachea,

whereas in Gonyosoma it is ventral.

The right lung extends posteriorly to the 230th g., which is just

four shields anterior totheanal. It is one of the longest recorded

in a Colubrine and resembles Coluber corais in this respect T

* P.Z.S. London, 1903. vol ii. p. 319.

+ Beddard, P. Z.S. 1906, p. 520.

ANATOMY OF THE OPHIDIA. 391

There is an apex about 18 mm. long, reaching to the 61st g.; it

is above and to the right of the trachea, to which it is closely

adherent posteriorly, the terminal 6 mm. only being free. The

lumen of the apex communicates directly with the lung and is

not set off by a restriction. The respiratory tissue ceases rather

abruptly at about the 80th g. ; its disappearance is not accompanied

by the gradual change in size and shape of the air-cells so

frequently seen.

The left lung, 86 mm. long, extends from the 62nd to the

80th g., and is the largest recorded in a Colubrine snake. The

base is attached to the right side of the ventral surface of the

right lung; it is 8 mm. long and corresponds to the 64th and

65th g. Though adherent to the right lung there is no direct

communication between the two, the interpulmonary septum being

imperforate. Just posterior to the base the diameter of the left

lung is 7 mm. There is a free apex, 12 mm. long, extending

forward nearly to the auriculo-ventricular septum; it is in

relation with the left side of the pericardium, to which it is

closely adherent. The posterior end of the lung extends 6 mm.

beyond the anterior tip of the liver. The left lung is closely

_ adherent to the inferior vena cava, and hides this vessel when the

ecelum is opened from beneath. The left bronchus is merely a

little bulge on the ventral side of the trachea 4 mm. anterior to

its termination. There is no trace of a left intrapulmonary pro-

longation or fibrous band. The lung is lined throughout with

regular air-cells ; these are larger and more shallow than in the

right lung.

The pulmonary artery courses along the right lateral edge of

the right lung. Its principal branches are as follows: No. I, a

minute one to the ventral surface of the apex. No. IT, a large

one which is at the level of the middle of the apex and winds

around to its outer side and then turns to the dorsal surface of

the lung. No. III enters the lung at the base of the apex.

No. IV, very small, crosses to the right over the middle of the

ventral surface of the isthmus joining the two lungs. No. V,a

large branch, which runs along the posterior border of the isthmus

and divides, sending several twigs to the right lung and a small

branch to the left lung. The branches beyond are too irregular

to describe in detail; some enter the lung direct, some wind

around the right side to the dorsal surface, and one large one

crosses the ventral surface and is distributed to the left side.

The right aorta gives the impression of being a branch of the

vertebral artery *. It arises from the right side and is less than

ove half the calibre of that artery. It flows directly to the

rear, forming an acute angle with the vertebral. It crosses to

the left aorta and upon reaching that vessel at the 64th g. it

courses alongside to the 68th g. before joining to form the

dorsal aorta. There is a single intercostal branch which enters

to the left of the median line at the 61st g.

* Quite as in Zamenis flagelliformis, Beddard, P. Z.5. 1904, vol. i. p. 388.

392 SURG. J. C. THOMPSON ON THE

As it was not feasible to study the vertebral artery without

injuring the air-chambers, this record was taken from another

specimen (17 6, same locality), an adult female, with 243 gastro-

steges and the heart apex at the 63rd g. The artery arises at

the level of the 56th g. and courses superficially to the 23rd g.,

where it entersinthe median line. There are fourteen intercostal

arteries arising as follows: I.—56, this arises quite in the anterior

angle formed by the vertebral and the right aorta and enters

in the median line; IJ.—56, enters to the left; III.-52, bi-

furcates, and one fork enters on each side of the body ; TV.—50,

bifurcates, both forks enter in the median line; V.—47, this and

the rest of the branches are single and enter in the median line;

VI-43; VII.-42; VIIL-40; [X.-39; X.-37; XI.-35; XII.-

33; XIII—30; XIV.—27.

In almost all serpents the intercostal branches of the vertebral

artery enter the body-wall in the mid-dorsal line*. In this

species the second and third branches offer two distinct types

which are exceptions to this general rule.

The right aorta, °5 mm. in diameter, joins the left, 1-5 mm. in

diameter, to form the dorsal aorta at the 68th g. The intercostal

arteries anterior to the kidneys enter the parieties to the right or

to the left of the median line: I.-73 and II.-83 to left; III.-87 to

right, and at the entrance it sends a branch to the left side which

bifurcates, sending one branch into the parieties and the other to

the peritoneal muscle. In the lumbar region the intercostal

arteries enter as follows:—At the 199th, 203rd, 204th, and

206th g. to the left. At the 208th g.a pair, one from each side

of the aorta, and entering on either side of the vertebral column.

At the 215th g. a single artery enters to the left. At the 215th

and the 218th g. the vessels are paired. At the 226th g. the last

artery is single and enters to the left.

The right spermatic artery arises at the 167th g. and enters

just anterior to the middle of the gonad. ‘The left arises at the

181st g. and enters the middle of the organ. Just posterior to

each testis there is an artery arising from the aorta which enters

the cord. There are five renal arteries on the right side. They

arise as follows: I.-203, courses to the rear for about one gastro-

stege, while the remainder enter nearly directly; II.-205;

II1—208; IV.-215; V.—222. The left kidney receives six

arteries as follows: I.-207; II.-209; III.-212; IV.-217; V.-219;

VI-222.

The anterior vena cava at the 40th g. is looped to form an

S-shaped curve, parallel to the median plane, and with the re-

current limb measures 6 mm. This suggests the provision for

the lengthening of the vein when the neck is inflated.

In the female (No. 17 6) the gall-bladder is 14 mm. in diameter,

the pancreas 18 mim. long, and the distance between the two

organs 6°5 mm.

The hepatic duct divides into about half a dozen tubes of un-

* Beddard, P.Z.S. 1904, vol. i. p. 368.

ANATOMY OF THE OPHIDIA. 393

equal size. The cystic duct, 29 mm. long, arises from the anterior

half of the ventral surface of the gall-bladder. Midway between

the gall-bladder and the pancreas it divides into two branches.

The smaller branch is one-third the diameter of the larger and

divides into two branches near the anterior border of the pancreas.

On the inferior surface of the pancreas the three branches of the

cystic duct join the several branches of the hepatic duct to form

a small but complex rete. Similarly complicated anastomoses

have been recorded in the Python by Poelman and in the Hama-

dryad by Beddard.

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :—

Male. Female. Male. Female.

per cent. per cent.

Total number of gastrosteges 234 243 100 100

TEI@RWAB RYDEN soso eceunmoodddceseas 69 63 28 26

iver sanbertor Ul) eared 81 75 35 ol

55 (DOSUETAIOIE GING! oe cesses 115 108 AD 44

Gall-bladder, centre ......... 139 137 59 56

JERWNGIRERIS, COIMNGWE covccocasccnses 142 ay) 61 57

Testis, right, anterior tip ... 168 a 72

X », posterior end... 173 ne 74

,, left, anterior tip ...... 180 aa il

i », posteriorend ... 185 Si: 79 it

Kidney, right, anterior tip... . 202 208 86 85

4s ,, posterior end. 226 233 96 96

aes. left, anterior tip ... 204 211 87 87

ue 5, posterior end... 233 Doe SY) 96

Tt will be noted that in the female there are nine more

gastrosteges ; in other words, the spinal column contains nine

more vertebre. In each specimen the posterior end of the liver

and the anterior tip of the right kidney are very nearly at the

same relative level. In each specimen the posterior end of the

left kidney is exactly at the 233rd gastrostege. This indicates

that the additional vertebrz in the female are situated between

the end of the posterior kidney and the vent. The position of

the vagina may require that the body be lengthened in this

region.

» This serpent has received considerable attention from the

early savants. It was discovered by Reinwardt in Java. By

Wagler it was removed from the genus Coluber and made the

type of the genus Gonyosoma; the determining character for this

change was the lateral angles on the ventral shields ; these are

well shown in the plates given by Schlegel and Jan. Gunther, in

the ‘Reptiles of British India,’ writes: “It is described as ex-

ceedingly strong and fierce, defending itself feroctously when

attacked. It raises nearly the anterior third vertically from the

ground before it strikes.” Cope, in the ‘Crocodilians, Lizards, and

Snakes of North America,’ gives a figure of the hemipenis and

places the genus apart from all the Colubrine, owing to the fact

394 SURG. J. C. THOMPSON ON THE

that the intromittent organ is highly differentiated. To this

may now be added the character to be found in the system of

air-chambers opening into the trachea. These structural con-

ditions will justify the removing of this serpent from the genus

Coluber and reinstating the genus Gonyosoma of Wagler.

PHILOTHAMNUS SEMIVARIEGATUS Smith,

Specimen.—Brit. Mus. Nat. Hist. Ex. No. ’13-6. Blue Nile,

Soudan. Adult female.

The trachea is black and terminates at the 51st g. The mem-

brane is well developed and lies along the right side of the tube:

the air-cells begin at the 45th g. The right bronchus persists as

an intrapulmonary streak to the 57th g. The right lung is lined

with air-cells to the 68th g., where they end very abruptly. The

left lung is a small air-sac opening into the trachea at the 5lst g.

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :—

FASUROSUCCES: Sipe puree ae reece nee 204 100 per cent.

ear bay ens.ce-tans ec eee eas 51 25

VET RUN) woe secs e eee cee 65 31:8

sh Wy OidG Neem acre oe a state 98 48

Gallbladder ete .cc-n-c eee oe: 114 55°8

Kaidineysnie bite tips sarees se: 184 90

ay 3 end!) Raeaee es 198 97

a2 Nehte taoae ner oseeince ce 187 91-5

ss Os ENGR cer ota eae 200 98 :

The scale-rows, the sequence in which they are suppressed, and

the gastrostege level at which they terminate are as follows :—

Neck.

17 rows, IV row ends, right 4th, left 4th gastrostege, leaving :

Body.

15 rows, VI row ends, right 112th, left 110th gastrostege, leaving:

Spee LV Gn ct cee pre UMN yy itl - oh

11 ,, which are continued to the vent.

CERBERUS RHYNCHOPS Schneider.

The salient characters of this species are: All the teeth,

without exception, are grooved. The tracheal membrane is

lined with a peculiar type of deep air-cells. The shifting of the

heart to the rear. The reduction of the renal arteries to one on

one side and two on the other. The particularly tough con-

nective tissue, which in this respect resembles that found in an

Hydrophid and differs from the texture in a Colubrine.

Specimen:—California Acad. Sci. No. 15301. Cavite Viejo,

Luzon, Philippine Islands. Cranium.

There are 16 mawillary teeth which are grooved on the external

quadrant; separated from these by a small space are two slightly

enlarged teeth, in a special compartment of mucous membrane,

ANATOMY OF THE OPHIDIA, 395

and grooved on the anterior quadrant. There are 9 large palatine

teeth which are grooved on the external quadrant. There are 21

pterygoid teeth, about one half the size of the palatine teeth and

grooved on the internal quadrant. There are 23 mandibular

teeth which are grooved on the external quadrant.

Specimen.—California Acad. Sci. No. 15309. Caloocan, Luzon,

Philippine Islands. Adult male.

The tracheal membrane begins at the glottis and lies along the

right side of the tube; at the 9th g. it is 1 mm. wide and at the

17th g. it is 2 mm. wide and begins to be lined with air-cells;

at the 44th g. it has increased until it forms three-fourths of the

circumference of the trachea; towards the base of the heart it

decreases somewhat in size and merges with the lung. The

respiratory tissue lining the tracheal membrane is of a specialized

type, the individual air-cells being from three to four times deeper

than wide. When the cartilaginous portion is laid open the

lumen appears to be but 2 mm. in diameter; to the left may be

seen the openings of the air-cells, which are in longitudinal rows.

The cartilages lie at first on the right and beyond the 4th g. on

the ventral side of the tube. The trachea ends abruptly at the

54th g., and there is not a trace of an intrapulmonary bronchus.

The right lung terminates at the 94th g., which is the same level

as the end of the liver. The posterior extremity is free and

rounded as in the Boine type, and not bound to the parieties ;

upon opening the ccelum it may be seen just posterior to the end

of the right lobe and to the right side of the left lobe of the

liver. The lung is lined with air-cells throughout, but towards

the end they become shallow and irregular. There is a free apex,

2mm. long, on the right of the trachea, and extending to the

52nd g. The left lung is completely suppressed.

The heart is located rather to the rear, being in terms of

gastrosteges 33 per cent. down the vertebral column, in Colu-

brine snakes the prevailing position is about 25 per cent. The

liver is divided by an incisure along the ventral surface into

right and left lobes; the right lobe begins at the 68th and ends

at the 92nd g.

The pulmonary artery divides into two equal branches opposite

the 5lst g. The anterior is distributed to the trachea, the

posterior to the lung. The branch to the lung divides just before

the tip of the apex of the lung into dorsal and ventral branches.

The vertebral artery arises from the right aortic arch at the

48th g.and runs superficially to the 14th g.; just before dis-

appearing it sends a minute twig forward to the next inter-

vertebral space. It gives off 23 intercostal branches which enter

in the median line; the gastrostege level at which they enter the

parieties is as follows:—I. arises just at the origin of the ver-

tebral and courses alongside of it to the level of the 45th g.;

E435 ee 4A VAs Veal Vili 30) Vil 38 3 iV ED

Bie ID Seaais) a Ca aVLs D832 OUTS a 0)s OGLE Me) 3s INE 27)

XOVES26r, AX 2 5 XG Hai, EVE 2 XO;

396 SURG. J. C. THOMPSON ON THE

XX.-20; XXI-18; XXII-16; XXIIT-14. The right aorta

gives off two intercostal branches which enter in the median

line: I-47; I1.-54. The dorsal aorta begins at the 58th g. and

as nearly as can be determined gives off 44 intercostal branches

which enter in the median line; the first is at the 57th g. and

the last at the 148th g.

There are four gastric arteries, the largest being at the 87th

and the 91st g. The superior mesenteric artery is at the 102nd

and the inferior mesenteric artery is at the 112th g. Each testis

receives a single spermatic artery; the right artery leaves the

aorta at the 112th g. and enters the organ just below the middle;

the left arises at the 119th g. and enters near the posterior end

of the gonad. The right renal artery is single; it leaves the

aorta at the 121st g.and enters near the anterior tip of the organ.

The left kidney has two arteries: I. leaving the aorta at the

128th g. and entering near the tip; IJ. at the 137th g. and

entering near the end of the organ.

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :—

Gasbrosteses: iro 5s.stesnears eee: 151 100 per cent.

Bleartvapem.:cpsca less urwet=e- es 3 35°]

NEIVIOK GUN a2 2c epe te Rene Nahas 62 42

Degg yl =50\(0 Litera ta ep ar SOR hes 94 62:2

Gallbladder: 2 e). 22th suena 102 67°5

eS tis svi lata eenes aeenee ethan 1) 13°7

wy Re) (V0 MANN ae ok aN as 118 18:2

sd Pay MONIC taonen ee ane ei 123 81:5

Kerelnteiye eit, rte en eee eee 122 80°8

i SMG SraaneNe iorict esc 135 89-4

‘a ORG wu mares. erences 129 86°5

- Day CT Gi rps aereece ee 141 93°5

The number of scale-rows, the sequence in which they become

suppressed, and the gastrostege at which this change occurs may

be thus presented :—

Neck. There are from 28 to 25 rows, which to the level of the

15th g. are asymmetrical and too irregular to record.

Body.

23 rows, VI row ends, right 84th, left 98th gastrostege, leaving :

DN IRE . ey LOOtha eNO aih oo ”

19 ” Til 9 ” 132nd, ) Polish 39 ”

17 ,, which are continued to the vent.

If this scale formula be compared with the one given under

Thamnophis ordinoides, it will be seen that two fundamentally

different types exist.

Jn the Colubrine snake the rows that were added or suppressed

were in adjoining series.

In the Opisthoglyphine serpent they were not in adjoining

ANATOMY OF THE OPHIDIA. 397

series, 2. e. one, No. IX, was next to the median row, and the

other, No. IV, was on the flank.

The scale formula of over one hundred species has been studied

and the serpents possessing the non-adjoining type have usually

been in the Dipsadomorphine. The following were the excep-

tions :—Loodon lineatus Dum. & Bibr., Zamenis rhodorhacis, Jan,

4. florulentus Geoff., and Coluber oxycephalus Boie.

This species is common in the vicinity of Manila, where it

inhabits the brackish water swamps that border the Bay. Over

a score have been seen in shallow places resting or moving slowly

on the bottom. They are able to remain beneath the surface for

several hours at a stretch. This may explain the need of the

extensive respiratory area in the trachea and lung.

LEPTODIRA HOTAMB@IA Laurenti.

Specimen.—Brit. Mus. Nat. Hist. Ex. No. 713-10. Fort Hall,

British Hast Africa. Adult male: total length 950, tail 85 mm.

The trachea terminates at the 34th gastrostege; it ends

abruptly, and the only trace of an intrapulmonary continuation

is in a regular arrangement of the air-cells in an imaginary line

in its prolongation. The tracheal membrane begins near the

glottis on the left side; further on it winds to the dorsal side of

the tube, and at the level of the 28th g. it becomes lined with

shallow air-cells. The right lung is reticulate to the 47th g.

The left lung, 2°3 mm. long, is at the level of the apex of the

heart; it is closely adherent to the left side of the pericardium,

and opens into the trachea at the 33rd g.

The vertebral artery arises from the right arch at the 29th

gastrostege, and is superficial to the 18th; it gives off five

branches, all of which enter in the median line. The gastro-

stege level of these branches is as follows :—I.-27; I1.-26;

IIT.-23; 1V.-22; V.-20.

The right aortic arch joins the left at the 38th gastrostege ;

at the 31st it gives off a single intercostal branch, which bifur-

cates close to the entrance in the median line, one fork supplying

each side. The intercostals from the anterior part of the dorsal

aorta enter in the median line.

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :-—

Gastrostesesh ease te ce 163 100 per cent.

team bua exce yaa ite tnee eae rele 34 20:9

vers, bio Mee saree feecee ars a 4] 25°2

pict) (8) ONC CBr eae MPRA ORT RAR 81 A9°7

Gall-bladdemeywyes, aes 1590 553

TEGiC hares, TAEINE SM co ceonensgencoe 129 80

aA PAE TiC) eh sk 2 155 95

Himlehty! tuoi Weck eine 136 83°5

MCP, sae. ee sme. 156 95:7

oP) 39

The number of scale-rows on the body, the sequence in which

398 SURG. J. C THOMPSON ON THE

they are increased or suppressed, and the gastrostege level at

which these changes occur may be thus presented :—

17 vows, [X row added, right 15th, left 17th gastrostege, making :

OT EX eters amide, al Oth _ leaving :

ip oy a. LN ee » 105th, ,, 104th

15 ,, which are continued to the vent.

9? 2)

PSAMMOPHIS SIBILANS Linneeus.

Spectmen.— Brit. Mus. Nat. Hist. Ex. No. ’13-1. Nile Delta,

Egypt. Adult male: total length 969, tail 315 mm.

The tracheal membrane is narrow, being only 1:5 mm. wide at

the base of the heart; it is confined to the left side of the tube

and the air-cells commence at the level of the auricles of the

heart. The bronchus is continued as a fine intrapulmonary

fibrous band to the level of the 55th g. The right lung is lined

with pulmonary tissue to the 58th g. and is without a free apex.

The left dung, 3 mm. long, opens into the trachea one gastrostege

below the apex of the heart.

The vertebral artery arises from the right aortic arch at the

36th g.; it gives off six intercostal branches which enter in

the median line:—I.-33; 11.-32; IJI.-29; IV.-28; V.—25;

VI.-23; a branch to the esophagus at 20 g.; at the 18th g. it

bifurcates, sending a small intercostal branch into the parieties,

and a larger one onto the dorsal surface of the cesophagus.

The right aorta gives off an intercostal at the 36th g. The

common aorta begins at the 43rd g.; the first few intercostal

branches enter on the left side.

The external landmarks of the principal viscera in terms of

gastrosteges are as follows :—

CRISUKOSIEGIVES \ooecadnescsen’ 2k 3 166 100 per cent.

Me ebNPO Xe aa naaes sce eeee eae A] 24:7

Mai RENes AUN se. Sesser eae os 48 28°9

Sy Sia eB ee aaa AON ea 82 49-4

Gallebladder o.2 j42e erode se 100 60°3

KGolmeyampiohit. hips. oes rece 142 85:7

is ae ae TCU 2 hae eae 163 98

go iptledibs: GW eeccetetieee ciuehes ie 146 88

5 Rae RETA LY eocters Wan aes 164 98°8

There are 19 scale-rows on the neck. On the body there are

17 anteriorly, decreasing to 15 and finally to 13 rows posteriorly.

OPHIOPHAGUS BUNGARUS Schlegel.

The salient characters of the Hamadryad may be thus

enumerated :—The large postparietal plates. The enlarged

scales in the vertebral row. (This was first noted in a stuffed

specimen in the Leyden Museum in which the skin had been

stretched, until, each scale being free, the exact size and form

was readily seen.) The extreme degree to which the grooving

is developed on the teeth ; every tooth except the fangs has one,

and some have two and even three distinct grooves. The series

ANATOMY OF THE OPHIDTA. 399

of air-chambers in the neck that open into the trachea. The size

it attains and its aggressive disposition.

Specimen.—California Acad. Sci. No. 16777. Singapore.

Cranium.

The mawillary bone extends beyond the palatine a distance of

‘8 mm. There are two fangs: the inner, 10 mm. long, are

cemented in the erect position; the outer, of the same length,

are loose and horizontal. There are three small teeth, 2°5 mm.

long, situated on the posterior two-fifths of the bone; each has

a groove along the antero-external quadrant that extends the

entire length and is deeper and wider towards the base. The

ectopterygoid bone is widened anteriorly and has a concave border

forming an antero-external and an antero-internal process. The

palatine bone has two processes. The maxillary process is short

and directed inwards; its base extends from the middle of the

third to the middle of the fifth socket. The vomerine process is

narrow and is arched upwards, inwards, and towards the end

slightly downwards; its base extends from the socket of the fifth

to the sixth tooth. There are eight stout teeth, 2°8 mm. long,

and each has three distinct grooves. The deepest groove is on

the antero-internal quadrant and extends from the base to

the tip. The next is not quite so deep; it is on the inner

quadrant and extends from the base nearly to the tip. The last

and least pronounced is on the external quadrant ; it is deeper

on the distal half of the tooth. The pterygoid bone is broadly

expanded posteriorly and but slightly curved. The articulation

with the ectopterygoid extends from the space between the fourth

and fifth tooth to a point 2 mm. bebind the last tooth. There

are ten stout teeth, 1-5 mm. long, situated on the anterior two-

fifths of the bone. Each tooth has two grooves; the deeper is

on the internal quadrant and the other on the external quadrant.

The dentary bone bears fourteen teeth, the anterior of which are

a trifle the larger. All are deeply grooved along the exterior

quadrant.

With the modification of the anterior maxillary teeth as fangs,

and the small teeth on the maxillary and dentary bones being

grooved, the King Cobra exhibits a further development of the

poison apparatus, as the teeth on the palatine and even on the

pterygoid bones are also grooved.

Measurements in millimetres.

Base, tip of premaxille to rim of

NOMAHTOH6) Toa eTANOO, 325 aoe oe aeoueodane 46

WIRSUIENAy poossenosospaogpedasos beausoue 13:9

Bichopbery Gord cm. o-een ase ese: 17°6

JES ENGILN®=eapee mob oyee Beas Aes onaaaqoor 17

IP BSIAVFEONG! Noacobtioptsoasobnbobeocoooaccss 32:5

IM Leyoye MONE" coceasogaHencsoopescuBocnoss0% 61-2

IDOI TAY co ome bdospenonaeceoeosetaetanas 25°2

@uraicinaieliens. seca ccene ne eee aes ee 2023

SORANTNCSAI nccasnadeonsseccqunooooesgear ASH

SRE Sw otscten eeaseiawie min nasiiaieiety oie c= 10

400 SURG. J. C. THOMPSON ON THE

Specimen.—California Acad. Sci. No. 15340. Butuan, Min-

danao, Philippine Islands. Head and neck of an adult.

A native was bitten by this beast at noon and died at seven in

the evening. In conformity with the local folk-lore the serpent

was decapitated, and the brain scooped out and applied to the

wound as an antidote.

The anatomy of this form has been made the subject of a most

interesting paper by Dr. F. KE. Beddard*, who discovered the

existence of the remarkable system of air-chambers connected

with the trachea. This specimen contributes a couple of minor

details to the description. The cartilaginous portion of the

trachea is U-shaped, with the interrupted ends to the right. The

tracheal membrane begins close to the glottis and runs along the

right side as far as the 16th gastrostege ; at this point it is tense,

parallel to the sagittal plane, and 6°5 mm. wide. The transverse

diameter of the trachea is also 6°5 mm. at this point. The air-

chambers are to the right of the trachea. The first compartment

is 55 mm. long and reaches from the level of the 2nd gastrostege

to the 11th g.; it has one foramen, 1°7 mm. in diameter, at the

6th g., and another, 2°7 mm. in diameter; these are situated on

the tracheal membrane and communicate with the lumen of the

windpipe. The second compartment, 12 mm. long, extends

from the 11th g. to the 13th g.; it has one foramen, 1°3 mm. in

diameter, near the posterior wall. At the middle of this chamber

there is an oval thin spot on the tracheal membrane. These thin

spots have been observed and referred to by Dr. Beddard as

‘imperforate foramina.” The third compartment, 9:5 mm. long,

extends from the 13th to the 15th g.; near the anterior wall is

a foramen 2°3 mm. in diameter, and near the posterior wall an

imperforate foramen.

The only other serpents known to possess a similar system of

air-chambers are Psewdoxenodon chinensis and Coluber oxycephalus.

The latter has in addition one of the longest right lungs recorded

in a Colubrine. To complete the data for the Hamadryad it is

necessary to ascertain the distance the anangious portion of the

right lung enters the celum. In the ordinary museum specimen

this is not easy to determine. A useful procedure is to suspend

_the specimen and to inject into the trachea a little 95 per cent.

alcohol, tinged with eosin, and allow it to trickle down into the

lung. With the least bit of luck the fluid will reach quite to

the end and stain the lung, so that it stands out in fair contrast.

The main point in the technique is to employ an extremely weak

staining solution.

DOLIOPHIS BIVIRGATUS Bole.

Specimen.—California Acad. Sci. No. 33059. Sarawak, Borneo.

Adult male: total length 1319, tail 128 mm.

The tracheal membrane in this species is of large dimension.

* P.Z.S. 1903, vol. ii. p. 319.

ANATOMY OF THE OPHIDIA. 40 1

It begins at the glottis and rapidly widens to comprise 50 per cent.

of the circumference of the tube; at the 42nd gastrostege it has

increased in width to 66 per cent., and just before the heart it is

fully 80 per cent. ‘The cartilaginous rings are markedly thin and

weak ; towards the end of the trachea they lie along the dorsal

and left segment of the tube. The trachea for its entire length

is destitute of air-cells. It gives every indication of fiiling all

the available space along its course. There is no line of demar-

cation to show where it ends and the lung begins. The alveolar

tissue of the left lung is thin; it begins very gradually at about

the level of the auriculo-ventricular septum, and continues

to the 116th g.; from this point to the 128th g. there are

a few large irregular air-cells before the anangious portion

begins. There is no free apex and no rudimentary left lung.

The heart apex is at the 91st g., and the tip of the liver at the

101st. Gastrosteges 272. There are 13 rows of scales from the

neck to the level of the 265th g.; at this point the median row

is suppressed, leaving 12 rows.

Causus RHOMBEATUS Lichtenstein.

Specomen.— Brit. Mus. Nat. Hist. Ex. No.’13-lla@, Fort Hall,

British Hast Africa. Adult male.

The trachea increases in diameter and occupies a large part of

the cervical region ; its walls are almost entirely formed by the

membrane. The tracheal membrane begins on the left side at

the level of the glottis; it rapidly widens, and after a few

millimetres becomes lined with pulmonary tissue which in no

respect differs from that found in a normal lung.

The cartilaginous portion at the origin almost surrounds the

tube, being incomplete only on the left ; at the 12th gastrostege

it becomes ventral and forms a small U-shaped gutter, 1-5 mm.

in diameter; as it approaches the heart it gradually flattens out

and is continued onto the anangious portion as a band. This

intrapulmonary bronchus ends at the 65th g.; it is not a mere

strip of fibrous tissue but a flat band of similar structure

to the tracheal wall, with the alternate sections of cartilage and

connecting membrane. The right lung has a very scanty lining

of pulmonary tissue ; thickly-set air-cells cease on the ventral

surface of the tracheal membrane at the 32nd g., dorsally they

are continued a little further; from this on to the heart apex

there are large oblong cells extending along each side of the

tracheal band ; on the lateral walls of the lung these alter to an

irregular shallow honeycomb structure; posterior to the heart

apex the lung is a simple air-sac, with the remnant of the

bronchus along its ventral wall. The left lung is completely

suppressed.

The heart is situated principally to the right of the median

line; it is rotated on its long axis until the right auricle is

nearly dorsal to the left.

The pulmonary artery divides 3 mm. from its origin into

402, ON THE ANATOMY OF THE OPHIDIA.

pulmonary and tracheal branches. The pulmonary branch is much

the smaller ; it runs along the left lung, at first to the left, then

down the centre of the bronchial band. The tracheal branch

runs anteriorly along the tracheal membrane.

The first portion of the descending limb of the left aortic arch

lies to the right of the left ventricle; this would be equivalent to

lying on the ventral surface of the heart were that organ not

rotated. The left aorta crosses to the left side ventrad to the

space between the heart apex and the liver tip; it then winds

around the cesophagus at the 46th g. and continues closely

adherent to and beneath the lung just to the left of the bronchial

band. The few intercostal arteries noted enter in the mid-dorsal

line. There is a single spermatic artery on each side. There are

nine right and ten left renal arteries: these are short and enter

directly. The iver is situated almost entirely to the right of the

median line; like the heart it also is rotated on its long axis;

the right lobe is dorsal to the left and posteriorly is 3-5 mm.

longer. The hepatic duct joins the cystic duct at the 85th g. and

the common duct enters the ventral surface of the pancreas. The

parietal peritonewm is greyish-brown; the visceral coverings are

but slightly tinged with the exception of that on the testis which

is very dark.

The external landmarks ot the principal viscera in terms of

gastrosteges are as follows :—

GaStroshemres. oes. cn jcceraeennts 145 100 per cent.

Heart apex...... sinc enlace Rept seated 42 29

Vier AUD mess tec ecea eee: see 4] 28:3

ova ol 89 a1 Raaes sl SESH a TRA HE eS 72 49-7

Gail) add Cru snout emergent 85 D713

Me shisseri& libs supe were cee 95 65°6

ye Ae OUNCE hist cimean tava 100 69

PRS elUs UL Open. carc nce hee 105 72°5

i amg Cl tera a, te eye 109 15-2

IKerdimeye nto bi pp eaerceee ese 108 (4:5

a Hefei! WSN Nee aS: SI, 138 95-2

3 KS erin Oe eaeeR See ese 116 80

oS SH C1216 Uae Amat SS le) 95:9

The scales are in 19 rows anteriorly, decreasing to 17 and

finally to 15 rows posteriorly. Gastrosteges 145,

ON THE AFRICAN HUNTING DOG. 403

26. The Coloration of the African Hunting Dog (Lycaon

pictus). By Major J. Stevenson-Hamivton, C.M.Z.S.

[Received March 17, 1914; Read April 21, 1914. ]

INDEX.

Variation and Atiology.

The classification into several subspecies of the Hunting Dogs

of South Africa has been apparently based upon the supposed

fact that animals inhabiting certain regions of the sub-continent

display colour-patterns clearly distinguishing them from their

relatives in other such regions, and so justify their division

into different sub-races.

My own acquaintance with the Hunting Dogs of other parts

of South Africa is purely cursory; but of those inhabiting the

Transvaal, especially the north-eastern portions, I have a fairly

intimate knowledge. J should remark here that, while, as I shall

endeavour to show, some of the distinctions of pattern and colour

brought forward as evidence of distinctive race are merely

individual peculiarities, there seems no doubt that the animals

native to the deserts of the south-west of the Union of South

Africa tend to be very much lighter in general colour than those

inhabiting the forests and savannas of the north and east.

I do not know to what type or types the Hunting Dogs of the

Transvaal Province would be assigned, but the natural features

dividing the western from the eastern portions of the Province are

very much more formidable than those which separate, on the one

hand, the eastern Transvaal from Rhodesia on the north and from

Zululand on the south, and, on the other, the western Transvaal

from Bechuanaland and Cape Colony respectively. Seeing, then,

that the Hunting Dogs of Zululand, which certainly rub shoulders

with the animals from the eastern Transvaal on the common ground

of Swaziland, have acquired the dignity of a separate subspecies, it

might reasonably be expected that the animals from the eastern

and western Transvaal and bush countries, separated as they are

from one another by several hundred miles of plateau country,

which is civilized and practically devoid of all wild game for many

years past, would be still more mutually distinct.

It is the case, however, that in the eastern Transvaal there are

found, in the same locality and even within the same pack, colour-

patterns which would fit, not only any of the determined sub-

-species, but, if produced singly for classification on the same

grounds, no doubt seemingly would justify the establishment, of

a good many more.

Proc. Zoou. Soc.—1914, No. XX VII. 27

404 MAJOR J. STEVENSON-HAMILTON ON

Again, it would be quite impossible to distinguish a group of

animals obtained, say, from Rustenburg in the west, froma similar

group selected haphazard from Lydenburg in the east. Almost

every individual shows certain special peculiarities, but the

general type of animals in the two widely separated districts in

no wise differs. In each we find the three colours of the coat

variously predominating in different individuals: throat ruffs,

backs of ears, black streaks on neck, show little uniformity. The

markings of the tails vary in extreme cases from all white to all

black, and while white tips are the rule, black ones are far from

uncommon.

The eastern Transvaal borders on the Mozambique Province

of Portuguese East Africa, from which it is separated only by

the low ridges of the Lebombo Hills, crossed freely by all wild

animals. It is known, in fact, that the various packs of Hunting

Dogs which ravage the Transvaal Game Reserves on the eastern

frontier, travel long distances into Mozambique, and no doubt

breed largely within that territory. It might, therefore, be con-

fidently anticipated that these animals would adhere closely in

type to the described subspecies Lycaon pictus typicus.

In the large number of skins which I have seen at various

times, some of which are still in my possession, it is, however,

quite impossible to reconcile all the colour types and markings

with one described subspecies. I am speaking now of the area

of the eastern Transvaal extending from the Limpopo River in

the north to the Crocodile River in the south, a range of not quite

four degrees of latitude.

T would specially mention three skins obtained in 1913. Two

of these represent a male and a female shot in November from the

same pack, at a spot near the Portuguese border of the Transvaal,

approximately 24° 46'S lat. I selected them from several others

secured on the same occasion, as presenting the most divergent

types. In one the yellow was considerably in excess of the black,

and the tail was mostly white with a black patch in its basal half.

In the other the two colours were reversed in quantity upon the

body, and the whole of the distal half of the tail was black and

the rest yellow. The latter skin, however, had less black upon

the ears than the former. The others in the pack all displayed

various colour-patterns of an intermediate nature, some with

white patches in the coats.

The skin of the third specimen was that of a large male shot

in about the same longitude as the others, but some thirty miles

farther north, in August, 1913. It belonged to a different pack.

Its coloration represented a third type, and between it and

the two extremes one finds all sorts of variations. It was

THE AFRICAN HUNTING DOG. 405

the representative of a rather common intermediate type of

colour-pattern, in which there is a fair amount of white shown

on the body and limbs, and the tail is about half black and

half white.

The northern part of the Sabi Reserve is covered with dense

thorn bush, and in the neighourhood of the large Olifants River

is rather stony. Then comes a stretch of savanna country some

70 miles in width, after which, in the neighbourhood of the

Sabi River, the country is the same as that first described. In

both the first and last the game is of similar character, mainly

Impala and smaller antelopes, with large numbers of Reedbuck

in certain localities. Im the savanna country are found mainly

Waterbuck and the plain-dwelling antelopes. Thus the Hunting

Dogs in these areas have to adapt themselves to slightly different

conditions of hunting, and while there is no doubt that in the

south and north they find that their prey comes comparatively

easily, in the centre they have to work hard to secure it.

So far as my observations have gone it seems pretty certain

that the savanna packs stick in the main to their own country,

though they occasionally raid north and south into the bush.

The same may be said inversely of the bush packs, north and

south. One might, perhaps, recognise some general similarity of

type between the bush packs of the north and south respectively,

in which, on the whole, the individuals are larger than are those

of the savanna country, and agree in being less extreme in

colour-patterns of their coats. But these animals of the north

and south probably never came into direct contact with one

another, whereas the intermediate packs touch each of them.

Instead, then, of having a gradual transition of type from north

to south, we find that the extremes have certain affinities in

common, mainly as regards size, and this may perhaps be

accounted for by the relative amount of food available for the

young animals. This is doubtless the foundation for the native

belief that there are two species of Hunting Dogs within the

same area, one larger than the other.

As regards colour-types, however, all agree in their individual

dissimilarities.

The following are a few measurements, taken at various times;

of full grown animals from the Portuguese boundary of the

eastern Transvaal.

27*

406 ON THE AFRICAN HUNTING DOG.

Head Tail

Sex. and (gs Be Neck. toe Girth. Height.

body. en eo

hairs)

Male ...... 43 133 16 5 25

a adetece 41 14 oe Be oe 274

rhe meade: 45 144 ‘ 26 284

ane aad 42 14 Ao 254

oe Sse os 41 14 13 23 28

a 40 13

33) aes 43 14 16 6 26

herr ee 40h 142 162 5h 212

ah Bae 45 14 17% 6 27% 30*

fone 41 14. ep 25

aay eee 39 134 15 24, 265

Female ... 38h 133 154 54 9A,

‘ 37 12 23 264

29 39% 145 1

is S62aa ele Se Hes iy 26

* The weight of this specimen, uncleaned and two hours after death, was 60 lbs.;

it was in spare condition (Huzting Dogs are mostly fat in my experience).

Steel tape measurements In inches.

Heights measured from highest point of withers to back of

palmar pad of fore foot.

Measurements round centre of fore leg midway between elbow

body behind elbows. [and wrist.

centre of neck.

? 99

29 99

ON A NEW AVIAN CESTODE. 407

27. On a New Cestode from an Albatross,

Diomedea irrorata. By H. A. Bayutis, B.A.*

[Received April 21, 1914; Read May 19, 1914.]

(Text-figures 1-4.)

IypeEx.

CESTODA : Page

Account of Tetrabothrius strangulatus, sp. n., in Diomedea

APTORALA TOI, Pe \beeserer tent se eee ee eae oie Se sden co seecens 407

Among some material presented to the British Museum by the

Hon. N. C. Rothschild, one tube contained specimens of a small

cestode collected from Diomedea irrorata Salvin by Dr. H. O.

Forbes. Locality : Lobos de Tierra Island, Peru.

Examination of these specimens showed them to belong to the

genus Tetrabothrius Rudolphi, to which in fact all the Cestodes

found in this group of birds have been assigned. The species

appears to be a hitherto undescribed one, for which I propose the

name Tetrabothrius strangulatus, on account of the sudden con-

striction observed behind the head, which gives the animal the

appearance of having had a thread tied round its neck.

TETRABOTHRIUS STRANGULATUS, sp. n. (Text-figures 1—4.)

External Features.

This is a very small form compared with most of the other

species of the genus. The material is unfortunately in a rather

fragmentary condition; the longest piece measured 57 mm.

This fragment contained 222 proglottides, in the first 5 or 6 of

which there were as yet no traces of genital organs developed.

The only fragment which includes a head shows about 30 seg-

ments, in which there are no genital organs, and the length of

this piece is about 3°5 mm., including the head. At the lowest

computation, therefore, a complete specimen probably measures

60 mm. in length and contains about 250 proglottides. The

maximum width attained is about 0°77 mm.

As already stated, only one head could be found among the

specimens. It is of a somewhat oblong shape, flattened dorso-

ventrally—i, e., in the same direction asthe strobila. At the apex

it has a flattened area, from which arises a slight conical pro-

jection. The four suckers are large, placed at the anterior end

of the scolex, and occupying less than half of its length. They

have a considerable ear-shaped fleshy expansion antero-laterally,

and their apertures are irregularly triangular in outline.

The total length of the head is 0°54 mm., while at its widest

* Communicated by the SzrcrETARY and published by permission of the Trustees

of the British Museum.

408 MR. H. A. BAYLIS ON A

part (through the lappets of the suckers) it has a transverse

wmeasurement of 0°36 mm. Its width behind the suckers is

0-28 mm. The suckers are 0°22 mm. long, and 0:15 mm. broad

in the middle of their length.

There is no rostellum, and no hooks.

Text-figure 1.

Head of Tetrabothrius strangulatus, X ca.70.

Ex. Aperture of excretory canal (paired).

All figures were drawn with Abbé’s drawing apparatus.

The apex of the head bears two pairs of curious apertures in

front of the suckers. These have the form of small, clear, circular

areas, crossed by a minute transverse slit. ‘They are probably

apertures of the excretory canals, fine branches of which can be

seen passing up to the openings in the head. A similar struc-

ture has been noted by von Linstow in Zetrabothrius heteroclitus

(7. auriculatus*|. Behind the head there is a sudden sharp

constriction, forming an isthmus only 0-08 mm. in width. After

this the width of the neck increases rapidly again to 0:17 mm.

Segmentation begins at about 0-25 mm. behind the constriction

the earliest proglottides being about 0°05 mm. long by 0:14 mm,

* Voyage of H.M.S. Challenger, Report on Entozoa (1884), pp. 14-18.

NEW AVIAN CESTODE. 409

wide. They increase rapidly in both dimensions on_ passing

backwards, but are always broader than their length.

Internal Anatomy.

Musculature.—The longitudinal bundles of muscle-fibres are

arranged in two layers, which are most strongly developed dorsally

and ventrally, The outer layer consists of slightly smaller bundles

than the inner layer, and is scantily developed towards the middle

line of the segments, being more in evidence towards the sides.

But neither layer is strongly developed quite up to the lateral

margins.

The inner layer consists of about 25 to 30 bundles.

Transverse and dorso-ventral fibres are only feebly developed.

Text-figure 2.

Eel

Tetrabothrius strangulatus.

Semi-diagrammatic view of the anatomy of a young, sexually mature, segment,

as seen from above by transparency.

Ov., Ovary; R.S., the coil destined to become the receptaculum seminis ;

Te., testes; Ut., uterus; Vag., vagina; Vit., vitelline gland; V.D., vas deferens.

Nervous System.—The usual pair of longitudinal lateral nerves

is present ; they are situated slightly beyond the most lateral of

the inner longitudinal muscles.

Exeretory System.—As usual, two pairs of wide longitudinal

lateral vessels are present. The dorsal pair is situated almost

exactly vertically above the ventral pair, and are of nearly

the same diameter. The ventral canals are connected in the

posterior part of the segments by transverse vessels.

410 MR. H. A. BAYLIS ON A

In the scolex fine canals can be seen running up to open by

the peculiar slit-shaped apertures on the apical projection.

Genital Organs.—The testes are proportionately large, com-

pared with those of other species of the genus, and are also

remarkably few in number. ‘They are nearly spherical, and have

a diameter of about 0°055 mm. ‘They are situated in the dorsal

part of the central field of the proglottis, and arranged on a

horseshoe plan round the other organs, which occupy the centre

and more ventral parts. The two ends of the horseshoe are

directed towards the right side of the animal, on which the

genital pores are always situated, and the male and female genital

ducts pass between them.

Text-figure 3.

py Te Re yl,

Tetrabothrius. strangulatus.

A. Transverse section through a sexually mature segment.

D.v., Dorsal excretory vessel; G.A., genital atrium; ., outer, M’., inner, longi-

tudinal muscles; IV., lateral nerve: Ov.,ovary; Ve., testes; V.D., vas deferens ;

Vit., vitelline gland; V.V., ventral excretory vessel.

B. Horizontal section through a gravid segment.

B.S. Receptaculum seminis; 7.V., transverse excretory vessel; Ut., uterus.

NEW AVIAN CESTODE. 411

The number of testes varies between 7 and 9, but the most

usual number is 8.

The ovary is a somewhat loose and irregularly lobed organ in

which the ova do not appear to be very closely packed. It

occupies the greater part of the centre of the proglottides in the

younger portion of the strobila.

Below the ovary, and extending a little in front of it, lies the

large, rounded vitelline gland, while above the ovary, in young

progiottides, the uterus appears as a very narrow crescentic tube

with conspicuous cell-nuclei, the convexity of the crescent being

towards the head of the worm.

In such a young segment, the genital ducts and apertures are

difficult to separate, the common genital atrium being as yet

imperfectly developed.

The vagina first appears as a rather wide canal running inwards

from the thickening on the right side, which is to be the genital

atrium ; at first it runs a nearly straight course, with a slight

forward inclination. It then turns backwards at an obtuse angle,

and forms a remarkable loop, returning over itself and running

upwards and forwards for a short distance, finally taking a sharp

bend downwards and descending perpendicularly to the ovary.

At the point where the backward loop of the vagina occurs,

there is at first no perceptible enlargement of the duct, which

appears to be of the same width throughout. But in older seg-

ments there is in this position an expansion to form a recep-

taculum seminis. In sections of the older proglottides this is

always seen to occupy the space between the backwardly-directed

‘“‘ horns” of the uterus.

The vas deferens is, in young segments, with difficulty dis-

tinguished from the vagina, lying as it does above it, and

following an approximately parallel course. Later it lengthens,

widens, and becomes very elaborately coiled, at the same time

becoming gorged with sperm. It comes to occupy a con-

siderable part of the right side of the segment, its coils being

concentrated mainly towards the anterior border.

Distally it passes into a large spherical cirrus-sac, which measures

0:055 mm. in diameter. Within this the duct continues to coil

about, and finally passes out on the opposite side, as the cirrus,

or male cloacal canal, This projects into the upper portion of a

large genital atrium, with thick muscular walls. At the point

where it leaves the cirrus-sac, the lumen of the cirrus shows a

considerable dilatation, which is always found to be full of sperm.

This may perhaps be regarded as serving the purpose of a seminal

vesicle, no other organ of that kind, apparently, being present.

Immediately below the male cloacal canal lies the opening of

the vagina, which passes inwards from the genital atrium, below

the cirrus-sac, curves upwards behind this to pass between the

dorsal and ventral excretory vessels, and thence passes, as

described, to the ovary.

As the proglottides grow older, the uterus is seen to enlarge at

412, MR. H. A. BAYLIS ON A

Text-figure 4.

Tetrabothrius strangulatus.

A. Portion of a horizontal section through a sexually mature segment.

B. Portion of a transverse section passing through a genital atrium and cirrus-sac.

C., Cirrus (‘‘male cloacal canal”); C’., dilated portion of cirrus; C.S., cirrus-

sac; D.V., dorsal excretory vessel; G.A., genital atrium; L.M., longitudinal

muscles; Ov., Ovary; 7.M., transverse muscle-fibres; Vag., vagina; V.F.,

ventral excretory vessel; JM., excretory canal.

NEW AVIAN CESTODE. 413

the expense of the ovary. At first it keeps its well-defined

crescentic shape, but becomes crammed with ova, and finally

expands so as to occupy nearly the whole of the segment. Mean-

while the rest of the genital organs become broken down and

disappear, though for a long time the outlines of the now vacant

testes can be seen, and the receptaculum seminis and vas deferens

are still distinguishable by reason of the spermatozoa contained

in them.

General Remarks.

Tetrabothrius strangulatus is distinguished from other members

of the genus by some interesting peculiarities, quite apart from

its size, which is unusually small,

One of its most distinctive features is the very sharp demarca-

tion of the head from the neck, giving the worm a “strangled ”

appearance. In other forms the head, behind the suckers,

usually passes almost imperceptibly into the neck, and there is

not, as in this species, any considerable portion of the head

between the posterior border of the suckers and the beginning

of the neck.

The small number of testes is also characteristic. As stated

previously, they do not exceed 9 in number, and the usual com-

plement is 8. In the majority of species of this genus there

are at least 22 testes (according to Fuhrmann). There may be

almost any number up to 60, and the number in a given species

generally varies slightly, but only in one species hitherto described

ure there as few as 8 (7. monticellii Fuhrm., from Fulmarus

glacialis. ‘Testes 8-12).

In the general arrangement of its internal organs 7. strangu-

latus approaches closely to 7’. heteroclitus Diesing. But this is a

considerably larger form, and its testes, though arranged some-

what similarly in a rosette or horseshoe pattern, are much more

numerous, and smaller in proportion. Fuhrmann* gives the

number as 28, but in some specimens in the British Museum

I have counted 43 in several successive segments, and some-

times an even larger number (probably about 50) +.

In conclusion it may be mentioned that hitherto no species of

Tetrabothrius—nor, so far as I am aware, any other Cestode—has

been recorded from Diomedea irrorata.

* Proc. Roy. Soc. Edinburgh, xxii. 1899, p. 649.

+ ‘The specimens of J. heteroclitus referred to are the types of “ Tenia dio-

medee”’ v. inst. and “ Tenia sulciceps”’ Baird, respectively, both of which

Dr. Fuhrmann considers identical with Tetrabothrius heteroclitus Dies. After

examining specimens of both, in spite of the discrepancy in the number of testes—

43 in“ T. diomedee”’; about 50 in “ T. sulciceps””—I have no doubt that this view

is correct.

ON A REMARKABLE CASE OF AFFINITY BETWEEN ANIMALS. 415

28. On a Remarkable Case of Affinity between Animals

inhabiting Guiana, W. Africa, and the Malay Archi-

pelago. By OxprieLp THomas, F.R.S., F.Z.S.*

[Received April 28, 1914: Read May 5, 1914.]

INDEX. Paze

Introduction ........... season occ sanesdaanscdaab cog Hull

E Gtesini Sg, n., or Guicre: Mm baadocaadtancn asl

In the lower Vertebrates a considerable number of cases are

known where there is an undeniable and direct attinity between

forms inhabiting the opposite sides of the Atlantic, but among

mammals such cases are excessively few, so that the discovery

of an additional one deserves a special record. That this case,

like the others, may be explainable without recourse to a land-

bridge—which there is little reason to believe persisted into

mammalian times—does not make it any the less advisable to

publish the case, so that it may be properly considered by

students of the subject.

Examples of a striking relationship between certain mammals

of West Africa and of the Malay region are of course numerous,

however difficult to explain quite satisfactorily, but it is note-

worthy that the present instance of transatlantic affinity is also

one of the best marked of the West Africa-Malay cases, and one

that has been often recorded in that connection.

This is the case of the Pigmy Squirrels, of which there are

some half dozen species in the Malay Archipelago, while a single

form—Sciurus minutus Du Chaillu, the basis of my genus

Myosciurus—inhabits Western Africa. Of the close relationship

of Myosciurus to the Hastern Vannosciwrus there can be no

doubt whatever.

In 1789 Buffon described = a little squirrel from Cayenne as

“Le petit Guerlinguet,” a technical name, Sciurus pusillus, being

attached to it by Desmarest later on. Then in 1867 Gray

described a small squirrel bought from the dealer Parzudaki and

said to have been collected by Castelnau in Brazil, as Macroxus

kuhlii, and this was, and I believe rightly, synonymized with

S. pusillus by Alston and other authors. Probably the specimen

was not collected by Castelnau at all, but was accidentally included

with Castelnau specimens by Parzudaki.

The skulls, both of this specimen and of another which was

fortunately so broken that no proper judgment on the characters

* Published by permission of the Trustees of the British Museum.

+ [The complete account of the new genus described in this communication

appears here, but since the name and a preliminary diagnosis were published in the

* Abstract,” ‘No. 133, 1914, the genus is distinguished by the name being under-

lined.—Ep. ]

+ Hist. Nat. Supp. vil. p. 263, pl. 66.

416 MR. OLDFIELD THOMAS ON A REMARKABLE

of the animal could be obtained from them, and merely on the

basis of its small size and its possession of five cheek-teeth it has

been placed in Microsciurus, a group of true squirrels found in

Central and North-western South America.

Now at last the Museum has received from Mrs. McConnell,

widow of the late Mr. F. V. McConnell, so long and frequent a

contributor to the Museum collections, three specimens of the

“Guianan Pigmy Squirrel, one of them with a practically perfect

skull.

An examination of this skull shows that instead of being in

any way related to Microsciwrus or other forms of American

Sciurine, the Guiana Squirrel is a member of the Nannosciur ne,

in which it forms a special genus closely related to Vannosciurus.

This genus may be diagnosed as follows :—

SCIURILLUS.

Abstract P.Z.S. 1914, p. 36 (May 12th).

General structure of skull and number of teeth as in Vanno-

sciurus, agreeing with that genus and differing from Myosciwrus

in all the characters recorded by me* as distinguishing these

genera from each other. An ectopterygoid present, broad, but

not so long as in Vannosciurus.

Postorbital processes over posterior root of zygoma. Inter-

orbital space as broad as the brain-case. Zygomata very broad

and strong. Anteorbital foramen small, far in front of the

teeth, as in Vannosciurus, its opening continued upwards as a

peculiar curved groove along the front edge of the anteorbital

fossa.

Cheek-teeth 4? as in Nannosciurus. Molars low, as in other

Nannosciurine, their set normal, as in Nannosciurus, the last

molar not facing outwards as in Myosciurus. Their upstanding

cusps, both above and below, very little developed. Their surface

more smoothly basin-shaped, with less evident transverse ridges.

Type. Sciurus pusitlus Desm.t

As a genus, Sciurillus is very closely related to Nannosciurus,

the reduction in the prominent transverse ridges of its molars,

the peculiar structure of its anteorbital foramina, and its high

but abruptly truncated ectopterygoids being its chief distin-

guishing characters. From d/yosciwrus, though both are un-

doubtedly of the same group, it is more widely separated.

In presenting this highly interesting case to students of

geographical distribution, I may point out that the whole of

North America is full of Squirrels of the other subfamily, the

Sciurine, and that these have penetrated into South America as

* Ann. Mag. N. H. (8) iii. p. 474 (1909).

+ Should any doubt be thrown on the determination of Sciwrus pusilius, the

genus should be considered as founded on the species represented by the type of

S. kuhlit.

CASE OF AFFINITY BETWEEN ANIMALS, 417

far as the tropic of Capricorn, and Sciurine cover the whole of

Europe, North Africa, and the continent of Asia, the Nanno-

sciurine being in the Old World rigidly restricted to a small

part of West Africa and to the Malay Archipelago. The addition

of Guiana to the known distribution of the group is therefore of

extraordinary interest.

Here, if ever, there would seem to be a case supporting the

persistence of the now generally admitted * transatlantic con-

nection into mammalian times, but bearing in mind how other

cases—such as those of the Tapirs and Opossums—have become

weakened by the discovery of fossil members of the groups in

N. America, Europe, and N. Asia, it would be wise not to lay too

much stress upon it, isolated and absolutely tropical as are the

three genera of Nannosciurine.

Moreover, the fact that in every character which separates the

other two, the Guianan Sciurillus agrees with the Malay Nanno-

sciurus and not with the African MZyosciurus, is also against this

ease having any connection with the ancient ‘“‘Gondwana-land,”

which at a time almost or quite pre-mammalian is supposed to

have extended from Eastern 8. America across Africa into the

Malay region.

* Cf. Andrews, ‘Tertiary Vertebrata of the Fayum,’ Introduction, p. xxvi (1906).

)

fin? ik aed

De aaa

ah Adena

IGN4S, ILI WWIS), IP.

Bale & Danielsson, Lt?

CITTOTASNIA LAGORCHESTIS.

Po 4 S UI4, MIA WIHS, Pl.

Ae ew.

oS ee

OZ Ii wigs ISS

PR ee NES 5 ae oe.

arn

ANNA

iyi

69590 OP og" a

2 @ ee

= SCS

ae ==

CEEMTUDPOS py

6 R.S.

Bale & Danielsson, Ltt

Cl WOMADINUA ILANCOIRCIE Sars,

IC,

1 Ay ty USNs, IVE WS), IPI, TM.

10 . Bale & Danielsson, L*®

Clr VOWARINWA Ib/ANGOINCisha)S Wiis).

Jo hg to USA IIE WATS), IP, NY,

SN &

Lose

——

—ra

»)

Ves.Sem.

Bale & Danielsson, Lt¢

CLT VOWATINIA iL/NGORCHIDSTIS:

Pi 4. Sy, USI4s, ILIDWWIS, El. WW.

sae See ier iy

eee

ae SEN ee

=< ide 35)

IS _ Bale & Danielsson, 1*¢

Ia, 1S. CMP WOWwAZE INIA” L/ACORCHIE STIS.

16. CITTOLANIA VILLOSA.

PA. S, IN, IWS, el. WI,

rm Ferny

rT me

peach Ce BY sail

a it nl oe

ee = N

ra fas i ERS Cole

0 op £0 Poe0s as b . . Re

}?-] 260 ® oor Oat ANS 046 Gy

Us cy 8 oe a 3 YH 02°) Wire yh \ epee

Tae 3

= b+

UI Cie AL?” oe

a4 opis

vines een on ge

Bale & Danielsson, ptd |

I7, 18, Clap WOWADINIUN LAGORCHESTIS.

Is, BO. CITTOTANIA VILLOSA.

Witt,

LOA LEWES Ee

PB, ZB. 8:

Bale & Danielsson, L*¢

NIA VILLOSA

Cir vO

Is 4. S, UIs, IDWS, 1, Witt:

KD.

Bale & Danielsson, 1*¢

Pa Sy VSG, ILI NWS 5 IP. IE.

Bale & Danielsson, 1*¢

IP, Za S, Ile, IGnWis, il. 3X,

Vés.Sem.

‘Bale & Danielsson, Ltd

Ce EOLA D INU, WALILIOWSVAL, *

ON TWO NEW MAMMALIAN CESTODES. 419

29. On Two New Species of Tapeworms from the Stomach

and Small Intestine of a Wallaby, Lagorchestes conspi-

cillatus, from Hermite Island, Monte Bello Islands.

By R. C. Lewis, M.A.(Cape), (1851 Exhibition

Scholar) *.

[Received March 17,1914: Read May 5, 1914.]

(Plates I.-X. + and Text-figures 1-3.)

INDEX.

Systematic : Page

Cittotenia lagorchestis, Sp. W. .....-.2.....s..eeveesenseereeees 420

OOGOEEO, BUNGE, D> WS 90 caes00 gondod coaese Sun hod ae6 opOR CHD 427

J am indebted to Mr. P. D. Montague, Caius College, Cam-

bridge, for this material obtained by him on the recent expedition

to the Monte Bello Islands. The parasites come from the intes-

tine and stomach of a species of Wallaby, Lagorchestes conspi-

cillatus, found only on two or three of the Monte Bello Islands.

The specimens were compared with those in the collection at

the Berlin Museum, thanks to the courtesy of Dr. A. Collin,

who kindly assisted in comparing these species with those in

the Museum, including Rudolphi’s collection. A comparison was

also made with the collection in the British Museum (Natural

History), where Cittotenia pectinata Goeze (21 & 25) and Citto-

enia denticulata Rudolphi (21) were examined, and no spe-

cimens bearing any resemblance to the species under discussion

were seen in the collection at the Zoological Gardens or at

the London School of Tropical Medicine.

Both species are characterized by the presence of two sets of

genital glands and two lateral genital pores to each segment. The

genital canals cross the longitudinal canals and nerves dorsally.

Interproglottidal glands are absent, and there are no calcareous

bodies. They belong to the genus Cittotenia Riehm. The history

of the genus is of some interest, and is discussed by Stiles and

Hassall (26) and by Lyman (21). Riehm (23 & 24) established the

genus in 1881, but later rejected it, placing his type Cittotenta

latissima Riehm in the genus Dipylidiwm Leuck. In 1891 Blanch-

ard included double pored leporine cestodes in the genus Monzezia,

taking Tenia expansa Rudolphi as his type. Later, 1893, Railliet

included these forms in the genus Ctenotenia with Tenia mar-

mote Frolich as type. It was not until 1896 that the genus

Oittotenia was re-established by Stiles and Hassall with the

following characteristics :—Anoplocephaline cestodes with seg-

ments broader than long and longer than thick, end segments in

some cases becoming longer and narrower. ‘T'wo full sets of genital

glands and two lateral genital pores to each segment; generally

* Communicated by Prof. E. W. Macprips, F.R.S., V.P.Z.S.

+ For explanation of the Plates see pp. 431433.

Proc. Zoou. Soc.—1914, No. XX VIII. 28

420 MR. R. C. LEWIS ON

one, sometimes two, simple transverse tubular uteri in each

segment, uterus generally with proximal and distal diverticula.

Vagina ventral to cirrus-pouch on both sides of segment. Dorsal

canal usually between ventral canal and nerve. Genital canals

cross longitudinal canals and nerves dorsally. Interproglottidal

glands absent. Caleareous bodies not yet recorded. Eggs with

well-developed pyriform bodies, the horns of which are long and

filamentous, crossing each other. Riehm based his diagnosis on

the noteworthy spread of the excretory system, which is not

ladder-like, but consists of three to five chief side stems and nume-

rous net-forming branches; on the double reproductive organs,

which open on the projecting lateral borders of the segments ;

and on the head appearing pushed-in in front and hookless.

Blanchard’s diagnosis of the genus Moniezia differs from that

of Cittotenia in that on the right side the vagina is ventral

to the cirrus-pouch and on the left side dorsal. Interpro-

glottidal glands present, and the horns of the pyriform body do

not cross, and end in a disk. The genus Thysanosoma, founded by

Diesing, 1835, contains double pored cestodes very similar to

those included in the genus Cittotenia, but differs in that the

genital canals cross the ventral canals and nerve dorsally and the

horns of the pyriform body are absent. There is one transverse

undulating uterus, and forms with one set of genital glands and

one genital pore are also included. Stiles states that from the

analysis of characters Cittotenia forms a genus intermediate

between Moniezia and Thysanosoma. The first of the species

now to be described was at first thought to belong to the genus

Thysanosoma Dies., resembling most closely Thysanosoma jum-

briata Dies., but it was found that the genital canals cross the

excretory canal and nerve dorsally, and that the species shows

considerable resemblance to Cittotenia zschokket von Janicki (10)

and agrees with this species in the absence of the pyriform body.

Von Janicki therefore holds that the generic diagnosis of the

genus Cittotenia should be altered from pyritorm body present to

pyriform body present or absent. Von Linstow in his description

of Cittotenia quadrata (20) describes and figures the vagina

running out and opening dorsal to the cirrus-pouch, and does

not state that this is contrary to the normal occurrence in the

genus. It is evident from the large number of synonyms for

each species, and from the departure from the generic charac-

teristics in some of the original descriptions, that the genus

Cittotenia requires revision. The two species from the Wallaby

show the generic characteristics as described for the genus Citto-

tenia by Stiles and Hassall (26) and are therefore included in that

genus.

CITTOTENIA LAGORCHESTIS, sp.n. (Pls. I.-IV.; V. figs. 14,15;

VI. figs. 17, 18.)

The specimens vary in length from 40-60 mm. and the greatest

breadth is 4 mm. There are usually about 150 proglottides.

TWO NEW MAMMALIAN CESTODES. 421

The strobile are uniform in breadth save at the anterior and

posterior extremities, where they become narrower. The scolex

is ‘6 mm. long by ‘75 mm. broad. The proportion of breadth to

length in young segments is as | to 3, in older as 1 to 6.

The scolex is rounded with no rostellum and no hooks. It is set

on the top of the strobila lke a dome-shaped knob. There are

four oval suckers. Each of the four is °35 mim. long by °25 mm.

broad. There is no neck, and the proglottides commence imme-

diately behind the head (figs. 2&3); each of them possesses a well-

marked projection at its hinder border which overlaps half of

the succeeding proglottis. The edges of these projections are

fimbriated, showing an even serration which gives the animal an

ornamented appearance resembling the condition found in Thysa-

nosoma fimbriata Dies. (22). The genital pores are found in the

middle of the projecting lateral margins of the segment. The cirri

are often seen projecting from the pores (figs. 12 i: 14). The mus-

culature does not present any marked peculiarities. It is lke

the structure described for (ittotenia zschokkei von Janicki (10),

where the musculature forms numerous fine-threaded bundles.

The subcuticular layer consists of well-developed longitudinal

and transverse muscular layers. There is also a system of

sagittal fibres which run dorso-ventrally, forming a dense network

of fibres between the transverse plates. They branch greatly

and these branches form a network around the testes and various

female glands. There is a ventral and a dorsal longitudinal

excretory canal on each side. The ventral canal is large, thin-

walled, and seldom circular in cross-section. The dorsal canal is

smaller and circular. It has a thick cuticular lining around

which is a layer of muscle-fibres. All the canals unite in the

head, forming an anterior loop between the suckers. The ventral

canals are connected in the posterior portion of each proglottis

by a transverse canal. There are no secondary longitudinal

canals connecting the transverse canal as described for C. pectinata

Goeze. The excretory system so far resembles that in C. varia-

bilis Stiles (21). On the terminal segment the canals open by a

simple pore situated towards the anterior end of the proglottis

(fig. 15). The large ventral canal discharges by simple trans-

verse tubes into this pore. There is no reservoir as described

for CO. pectinata Goeze (21), nor is there a projection of the

parenchyma into the ventral canal at the posterior end of each

segment, as described by Fuhrmann in C. kuvaria Shipley (7).

The first rudiments of the genital organs appear in the pro-

glottides immediately behind the head (fig. 3). There are two

sets of genital glands in each segment (fig. 7), and two

uteri in the early segments up to the 80th segment, where they

meet in the median field, and it appears ag if there were only

one uterus (fig. 1). The uteri possess proximal and distal diver-

ticula. Very young segments in which the reproductive organs

are in their first rudiments, show the vagina running out to

open into the genital atrium posterior to the cirrus (figs. 5 & 6).

28*

422 MR. R. C. LEWIS ON

The vagina is a thick-walled tube with a narrow lumen. It is

connected with a very large receptaculum seminis. With in-

creasing age, before the ovary is functional, the vagina atrophies

from the pore inwards, and finally, when female and male canals

are fully developed, the vagina remains only as a small tubular

projection from the receptaculum (figs. 9 & 11). This resembles

the condition found in C. zschokkei von Janicki (10), and von

Janicki concludes that the young proglottides, whose testes are

not yet developed, function as females, and that the older pro-

glottides with abortive vagine function as males, although the

female organs are otherwise fully developed. The eggs must all

be fertilized by the sperm stored up in the receptaculum seminis,

which receives sperm through the vagina from the same worm or

from another worm by copulation in the young state, and this is

proved by the fact that the receptaculum of proglottides in which

testes are not yet developed are filled with spermatozoa.

Male Reproductive Organs.

The first rudiments of the testes are seen about the 12th seg-

ment. They reach maturity about the 60th segment, and are seen

as small rounded sacs lying in the dorsal field of each segment

between the transverse excretory canal of the preceding segment

and the cirrus-sac (figs. 7 & 11). They do not extend in towards the

median field as in C. kuvaria Shipley (7), but are confined to the

dorsal region between the ventral excretory canal and the inner

termination of the cirrus-sac. From these testicular sacs, about

50 on each side, little ducts are given off which connect with the

vas deferens. The vas deferens is a slender duct which runs in

towards the median field and then loops back above the recep-

taculum seminis to run obliquely down to the cirrus-pouch. The

vas deferens does not extend further towards the median field

than the inner extremity of the receptaculum seminis. Before

entering the cirrus-pouch there is a circular swelling, the vesicula

seminalis, with a diameter of *1 mm. anda thick wall. The cirrus-

pouch is elongated and of uniform size and thickness in mature

segments, 1 mm. long by -2 mm. broad (figs. 9 & 11). It has mus-

cular walls consisting of an outer longitudinal layer and an inner

thicker circular layer. The cirrus also has circular and longi-

tudinal fibres. The space between the muscular wall of cirrus-

sac and cirrus is filled with spongy tissue. The cirrus is thick-

walled with a narrow lumen. It is often seen protruding toa

great extent, nearly as far as the length of the cirrus-pouch. It

is covered externally with closely set bristles. The outer end of

the cirrus-sac discharges into a space, the genital atrium. Its

outer end is guarded by a circular muscular sheath, the sphincter ;

and beyond this sheath, in the genital atrium, is a circular pro-

jection of the floor of the latter.

Testes are not observed after the 100th segment, though the

cirrus-sac persists to within twelve segments of the posterior

TWO NEW MAMMALIAN CESTODES. 493

extremity. In this respect this species differs from C. pectinata,

where the testes are described as persisting to the last proglottis.

It is noteworthy that in this species at the tapering posterior

extremity no genital glands are observed in the last 12-15

segments (figs. 13 & 15).

Female Reproductive Organs.

These appear earlier than those of the male. The first rudiments

of the genital glands are found in the segments immediately

behind the head (fig. 3). ‘The vagina opens to the exterior up to

_ the 25th segment, but after that aborts from without inwards. The

receptaculum is prominent from the first and persists throughout

the strobila, except in the last 12-15 segments where, as already

mentioned, no genital glands are observed. The ovary is rudi-

mentary up to about the 40th segment, where it takes on its mature

form, and at about the 50th segment the first traces of the uteri

are seen. After the 100th segment all of the female organs

except the uterus begin to atrophy, and in the posterior segments

only the uterus and receptaculum seminis persist (figs. 12 & 14).

The vagina, only complete in the early segments, runs out ventral

to the cirrus-pouch and opens into the genital atrium posterior to

cirrus (fig. 6). At its distal end the lumen is very small and the

cuticular lining very thick. In the region beneath the cirrus-pouch

it is surrounded by deeply staining cells as in C. variabilis Stiles

(21), and in this region has numerous bristles on its walls. It is

a straight tube, and runs in to open into the large receptaculum

seminis. The receptaculum seminis is elongate, °2 mm. long by

‘1 mm. broad. It is surrounded by a thin layer of circular muscle-

fibres, and outside the fibres is a single layer of cells which stain

deeply. The ovary consists of anumber of ‘“ Indian club” shaped

pouches. Each pouch opens into a common reservoir by a narrow

neck (fig. 10). The oviduct runs from the reservoir of the ovary

towards the yolk-gland. Beyond the opening of the vitelline

duct into the oviduct, the latter passes through the shell-gland

and enters the uterus at a spot situated just below the ovary.

The ovary is situated dorsal to the receptaculum. Its outer

tubules overlap the receptaculum, but the inner tubules are

slightly nearer the median field. Ventral to the ovary and near

the posterior margin of the segment is the small vitelline gland.

Alongside it, towards the median field, is the comparatively large

shell-gland (fig. 11). The duct from the receptaculum seminis

opens into the reservoir of the ovary at the side nearer the

margin.

The uterus is formed dorsal to the ovary. The female genitalia

are crowded close together. The uteri, at first separate, gradually

extend towards the median field, and their blind ends become

closely applied, so that in older segments it is difficult to distin-

guish two uteri. The uterus does not cross the longitudinal

canals and nerves, but in older segments pushes them nearer the

AQA4A MR. R. C. LEWIS ON

margin. It is possessed of numerous proximal and distal diver-

ticula. As already mentioned, the egg possesses no pyriform

body.

The nervous system has not been worked out, as the specimens

were fixed in alcohol, but from the sections it appears that the

condition is like that in C. quadrata von Linstow (20), with a

single trunk on each side to the outside of and dorsal to the

two excretory canals, and a ganglionic mass is observed beneath

the loop formed by the excretory canals in the head.

Text-figure 1.

C. zschokket von Janicki. Young already fertilized proglottis in optical section,

from von Janicki (10), p. 130. 45.

A.w.Dr., rudiments of female genital ducts ; C.b., cirrus-pouch ; #, excretory vessel ;

Ha., vudiments of testes ; WV, longitudinal nerve; #.s., receptaculum seminis ;

Ut.a., rudiments of uterus; V.d., vas deferens ; Vg., vagina.

Affinities and Contrasts with other Species.

In describing this species it has already been pointed out that

of all the species of Cittotenia this one resembles most closely

C. xschokker von Janicki (10). The chief points of resemblance are

the possession by both species of a projecting sheath, which is

Text-figure 2.

H. (= Vs, [2 N Ch

Tae 3G, GOR @)

OD Re GaP aba Bee e ct ra lol =

} Li i:

C. zschokkei von Janicki (10). p. 130. Ripe proglottis in optical section. X46.

Dst., vitelline gland; H., testes; Ks#., ovary ; Ut., uterus ; V.s., vesicula seminalis.

Other lettering as in text-fig. 1.

fimbriated, at the posterior border of each segment overlapping

half the next segment: also the development and position of

the reproductive organs; the absence of the pyriform body in

TWO NEW MAMMALIAN CESTODES, 425

the egg; the large receptaculum seminis, and the vagina aborting

in later segments. There are, however, important points of

difference which mark C. lagorchestis as a distinct species. In the

first place, there are only two longitudinal excretory canals and

one transverse excretory canal in each segment in (. lagorchestis,

whereas in C. zschokkei there are three longitudinal excretory

canals and two transverse canals to each segment. The most

important contrasts are found in the reproductive organs, and

may be tabulated as follows :—

C. lagorchestis. C. zschokkei.

(1) Vas deferens a slender tube

with an oval swelling, the vesi-

cula seminalis, near entrance into

cirrus-sac. Vas deferens recurves

to receive ducts from testes.

Vas deferens much swollen through-

out its length to form an elongate

vesicula seminalis. Vas deferens

passes straight to testes.

(2) Cirrus-pouch extends across the

longitudinal canals and nerve

towards the median field for half

its length.

(3) Testes situated dorsal to pro-

jection of cirrus-pouch in inner

parenchyma between ventral

longitudinal excretory canal and

inner termination of cirrus-

pouch.

(4) Ovary dorsal to receptaculum

seminis, and consisting of a circle

of “Indian club” shaped tubules

Cirrus-pouch does not extend in-

wards over excretory canals and

nerve,

Testes more median, situated dor-

sal to ovary, receptaculum

seminis, and vitelline gland, and

not extending outwards to longi-

tudinal excretory canal.

| Ovary on same plane as receptacu-

lum seminis, nearer median field

consisting of an irregular mass

Opening into a common reservoir not showing tubular arrangement.

by slender necks.

(5) Two uteri, whose origin is dis-

tinct as separate tubules, but

which with increasing age

approach each other and fuse in

the median field.

Uterus originates as a single tube.

The only resemblance with C. kuvaria Shipley (7) is that the

cirrus-pouch is much elongated and stretches over excretory

vessels and nerve into the inner parenchyma. Also there is a

large shell-giand in both species, and in ripe proglottides the

uterus fills the whole of the inner parenchyma, not extending

over the excretory canals and nerves but pushing these nearer

the margin. The contrasts are well marked. In C. kuvaria the

excretory system consists of extraordinarily wide channels, two

large ventral and one smaller dorsal vessel. and the hind end of

each proglottis shows a tongue-like projection of parenchyma

into ventral vessel which lies on a projection of the wall of the

latter and forms a barrier which prevents a backward flow of

excretory fluid. No such structure is observed in C. lagorchestis.

426 MR. R. C. LEWIS ON

Although the male organs are doubled the testes are not clearly

divided into two groups. They are united in the median field

of the proglottis. In OC. lagorchestis there are two distinct

groups lying in dorsal anterior field of segment, next to ventral

excretory canal, and not spreading further towards median field

than the inner border of the cirrus-pouch or the outer border of

the receptaculum seminis. In C. kuvaria the ovary is ventral and

the vitelline gland dorsal, while in C. lagorchestis the ovary 1s

dorsal and the vitelline gland ventral. There is a single uterus,

and the receptaculum is covered with a star-shaped epithelium in

C. kuvaria. In C. lagorchests the uterus is double, and there 1s

no star-shaped epithelium covering the receptaculum seminis.

Text-figure 3.

Sa ie

Foon gon gonodson Oe

Transverse section through a lateral portion of Cittotenia quadrata von Linstow,

from (20) fig. 3, p. 680. c, cirrus; %, nerve; g, excretory vessels; d, vitelline

gland; s, shell-gland; , vagina; 7, receptaculum seminis; &, ovary;

h, testes.

C. quadrata von Linstow (20) shows very little resemblance

with CO. lagorchestis. There are two longitudinal excretory vessels,

a larger ventral and a smaller dorsal, and a longitudinal nerve

dorsal to these, as in C. lagorchestis. The chief points of -con-

trast are that the vagina runs out and opens dorsal to the

the ovary; the shell-gland is ventral to the vitelline gland, and

the testes are dorsal to the ovary and extend in towards the

median field. The eggs are four-sided and possess a pyriform

body, and there is a single uterus,

C. pectinata Goeze (21 and 25) was examined at the British

Museum (Natural History). The head is not sharply marked off

from the strobila as in C. lagorchestis, nor are the suckers, which

are set on the rounded anterior extremity, so large and prominent.

The lateral margins of each segment are rounded and not

TWO NEW MAMMALIAN CESTODES. 427

projecting as in C. lagorchestis. There is also a slight rostellar-

like projection at the anterior end of the scolex. he posterior

projecting border of each proglottis is smooth and only slightly

overlaps the following segment, and the cirrus is never found

protruding from the pore. There are secondary longitudinal

canals uniting the transverse excretory canals. In the posterior

proglottis the excretory canals open into a reservoir which opens

at the excretory pore through many canals. The testes do not

disappear, but are found in the posterior proglottis grouped round

the excretory pore. There is an enlargement just inside the

cirrus-pouch filled with sperm, the vesicula seminalis, and there

are two seminal receptacles, a smaller external and a larger

internal sac. There is a single uterus which extends over

excretory canals and nerves into the lateral margins, The horns

of the pyriform body are long and filamentous. The above points

are In marked contrast with C. lagorchestis.

Cittotenia denticulata Rudolphi (25) was also examined at the

British Museum (Natural History), and differs from C. lagorchestis

chiefly in that the testes are in two groups in the lateral portion

of the median field. The cirrus-pouch is short and does not reach

the longitudinal nerve. There is a single uterus, and in ripe

proglottides it extends over the longitudinal canals and nerves

into the lateral margins.

These contrasts suffice to show that C. lagorchestis is a distinct

species. It has been compared in the same way with all the

described species mentioned in the Index Catalogue (27) and in

the Zoological Record, and the disparity has been sufficient to

indicate that it is a new species.

CIrroTHNIA VILLOSA, sp.n. (Pls. V. fig. 16; VI. figs. 19, 20;

Vil—X.)

These specimens have a leaf-like strobila and the shape re-

sembles a large liver-fluke (fig. 16), being widest near centre and

gradually tapering towards both extremities. The posterior

proglottides become narrower but not longer, and thus differ

from CO. pectinata Goeze (21), in which species they also become

longer. The length varies from 40-80 mm. and the greatest

width is 6 mm. The number of proglottides in strobila

vary from 120-200. The proportion of breadth to length

in youngest proglottides is 1:3, in older 1:8. The scolex

stands out from the anterior end of the strobila like a truncate

cone (fig. 21). It is 8 mm. long by 6 mm. broad. The suckers

are large and conspicuous and set on the widest part of the head.

They are 3 mm. broad by 4 mm. long. There is no rostellum

or hooks. In transverse section the head appears quadrate with

rounded corners. The suckers occupy the four corners and nearly

meet in the median line (fig. 19). In C. lagorchestis the head

appears circular in transverse section (fig. 4). There is no neck,

and the proglottides commence immediately behind the head.

In this species the posterior borders of the proglottides have

a backwardly projecting sheath with more markedly fimbriated

428 MR. R. C. LEWIS ON

posterior border than in C. lagorchestis (figs. 16, 22). The fim-

briation is in the form of a regular serration in the earlier

proglottides (fig. 21), only overlapping half the succeeding pro-

glottis, but in older proglottides these fimbriations are much

elongated to form a mass of elongate hair-like projections which

cover two or three of the succeeding proglottides. These pro-

jections become more elongate towards the posterior end and

project 4 or 5 mm. beyond the posterior segment (fig. 16). In

no described Cittotenia is this sheath from the hind border of

each proglottis broken up at its free edge in such a marked

degree into hair-like lappets which project so far backwards over

the succeeding proglottides.

Considerable difficulty was experienced in working out the

internal structure of this species, because several of the stvobile

were mounted whole, and no trace of reproductive organs was

observed in any of the proglottides (fig. 16). This was

confirmed by series of sections throughout the strobila, which

showed absence of reproductive organs (fig. 25). This was the

more remarkable, in that the specimens without reproductive

organs were not smaller in size, nor did they have fewer segments

than those found with reproductive organs.

In the forms found with reproductive organs no trace of these

was to be seen in the first 50 segments, after which an abrupt

transition was met with and proglottides were found with

reproductive organs. The reproductive organs reach maturity

very rapidly, and very few segments contain them in a rudimentary

state.

Again, after the 120th segment no reproductive organs were

found. The segments with reproductive organs therefore

were situated in the median portion of the strobila. This is in

contrast with all other species of Cittotenia, where the first

rudiments of the reproductive organs appear in the segments

immediately behind the head.

The musculature is similar to that described for C. variabilis

Stiles. The sub-cuticular layer, consisting of longitudinal and

transverse layers, is especially well developed. The excretory

system appears also to agree with the condition found in

C. variabilis, but no reservoir was detected on the posterior

proglottis, and the ventral canals discharge by a simple pore on

the last proglottis, as in C. lagorchestis. The ventral canal is

especially large and occupies a large part of the lateral field. One

main longitudinal nerve trunk is present on each side dorsal to the

two excretory vessels, but the details of the nervous system could

not be determined in these specimens killed and fixed in alcohol.

Male Reproductive Organs.

There is only one testis-sac on each side, which is large,

‘4 mm. in diameter, and filled with a mass of developing sperm

clusters (figs. 30 & 31). It is situated in the dorsal anterior

field of each segment, a little nearer the median field than

TWO NEW MAMMALIAN CESTODES. 499

the inner border of the receptaculum seminis. ‘The vas deferens

is coiled, and loops first towards the median field and then dorsally

back towards the margin. A large part of the vas deferens is

swollen to form an elongate vesicula seminalis as in C. zschokket

von Janicki (10). The vas deferens passes obliquely downwards

to the cirrus-sac. The cirrus-pouch is even more elongate than

in (. lagorchestis, 1:4 mm. long but not quite so broad. It is

possessed of the saine musculature on its wall as in C. lagorchestis,

and the cirrus is also thick-walled with a muscular wall, a narrow.

lumen, and is covered with numerous bristles. Also, between the

wall of the cirrus-sac and the cirrus is a mass of spongy tissue.

In a few cases the cirri are seen protruding from the male genital

pore (fig. 26), but in most cases cirrus and cirrus-sac are retracted

from the margin and the muscular sphincter is retracted into the

sub-cuticular parenchyma, and a wavy duct with wrinkled walls

passes to the exterior to open at the anterior border of the lateral

margin (figs. 23 & 24).

Female Reproductive Organs.

There is a large receptaculum seminis as in C. lagorchestis,

and lying dorsal and to the innerside of this are a number of

tubules, arranged fan-wise, which collect into a circular sac,

alongside of which, nearer the median field, is a small bean-

shaped vitelline gland. The duct from the receptaculum seminis

enters the circular sac of the ovary. The oviduct passes out

dorsally, and passes through the shell-gland near its origin from

the ovary and enters the uterus dorsal to the ovarian tubules.

The vagina is a thick-walled duct much as in C. lagorchestis,

but is not abortive. It does not open into a genital atrium

but opens separately in the middle of the projecting lateral

margin of the segment (figs. 23 & 24). The uterus arises as a

single transverse duct and has proximal and distal diverticula. It

passes above the female genitalia on each side to run out over

the longitudinal canals into the lateral margins as in C. pectinata

Goeze (fig. 27). The egg has a pyriform apparatus, the horns of

which are filamentous and cross one another (fig. 29).

The characters which show that this species is in distinct

contrast to other species are :—

(1) The extraordinary development of the fimbriations of the

projecting sheath at the posterior border of each proglottis (fig. 16).

(2) The presence of two large testicular sacs instead of, as

in other species, numerous small testicular sacs (figs. 30 & 31).

(3) The absence of reproductive organs entirely in some

strobile (fig. 16).

4) The absence of reproductive organs in the earlier pro-

glottides (fig. 25).

(5) The male reproductive opening situated at the anterior

edge of the projecting lateral margin and the female reproductive

opening in centre of margin (figs. 23 & 24).

430 MR, R. C. LEWIS ON

The species cannot be said to approach any of the described

species closely, though possessing the general characteristics of

the genus.

Of the works of reference the following list and also most of

those given in the bibliography of (14) Johnston’s ‘“ Entozoa

of Monotremata and Australian Marsupials” were consulted.

For comparison of the above described species the following

were found to be of most service:—7, 10, 11, 14, 15, 20, 21, 22,

23, 24, 25, 28, 29, 30, 31.

BIBLIOGRAPHY.

(1) Bepparp.—Proc. Zool. Soc. Papers, 1911-1913.

(2) Boas.—Zoologische Jahrbiicher Syst. xvii. 1902-3 (Triplo-

tenia mirabilis).

(3) Braun.—Cestodes, in Bronn’s Klass. und Ordn, des Thier-

Reichs, [V. Vermes, 1 Band, 1894-1900.

(4) Bremser.—Icones Helminthum: Systema Rudolphii Ento-

zoologicum illustrata, 1824.

(5) Cons.—Agricultural Gazette of New South Wales, vol. xvi.,

1905. (The Tapeworms of Australia.)

(6) Dissine.—Systema Helminthum, vol. i., 1850.

(7) FunrmMann.—Cittotenia kuvaria. Zool. Jahrb. Syst. xxii,

1905. (‘ Uber die asiatische Vogel-cestoden.”)

(8) ——. Bulletin of the Royal Belgian Academy, 1913.

(9) von JanicK1.—Zoologischer Anzeiger, Band xxvii., 1903-4.

(‘* Weitere Angaben iiber Zriplotenia mirabilis.”)

(10) Zoologischer Anzeiger, 1906, Band xxix. ‘ Beutler-

cestoden der Niederlindische Neu Guinea Expedition,

Zugleich einiges Neue aus dem Geschlechtsleben der

Cestoden.”

(11) Zeitschrift fiir wissenschaftliche Zoologie, Band 81,

1905. (“Studien an Saugetiercestoden.”)

(12) Krerrr.— On Australian Entozoa.” Trans. Entomol. Soc.

N.S. Wales, 11. 1873.

(13) Jounsron.—“ Notes on Australian Entozoa, No. 1, 1909.”

Records Austral. Museum, vol. vu.

(14) “The Entozoa of Monotremata and Australian

Marsupials, No. 1.” Proc. Linn. Soc. N. 8. Wales,

WO, socanvo, SOM)

(15) “The Entozoa of Australian Marsupials, No. 2.”

Proc. Linn. Soc. N. S. Wales, vol. xxxvi., 1911.

(16) ——. “On a new genus of Cestoda from a Marsupial.”

Austral. Assoc. for Advancement of Science, Jan. 1911.

(17) Australian Institute for Tropical Medicine: Report

for the Year 1911. (Cestodes & Acanthocephali.)

(18) Lerpy.—Proc. Acad. Nat. Sci. Philadelphia, vol. xxvii.,

1875. (Tenia bipapillosa from an Australian Wombat.)

(19) von Lixnsrow.—Compendium der Helminthologie, 1878.

(Ein Verzeichniss der bekannten Helminthen.)

TWO NEW MAMMALIAN CESTODES. 431

(20) von Liystow. Centralb. f. Bakt., Orig.-Bd. xxxvii., 1904.

(Neue Helminthen.)

(21) Lyman.— Studies on the genus Cittotenia.” Trans. Amer.

Micros. Soc. 1901-2, xxiii—xxiv.

(22) Ratiurer.—Traité de Zoologie Médicale et Agricole. 1895.

(23) Rreum.—Studien an Cestoden. Dissertation, Halle, 1881.

(24) Zeitschrift fiir die gesamten Naturwissenschaften,

Bd. liv., 1881.

(25) Srruus.—“ A Revision of adult Cestodes of Hares and

Rabbits.” Proc. U.S. Nat. Mus. vol. xxix., 1897.

(26) Srmes & Hassaun.—vw.S. Dep. of Agric., Bulletin No. 4,

1893. (“A Revision of the Adult Cestodes of Cattle,

Sheep and allied Animals.”)

Index Catalogue of Medical and Veterinary

Zoology : Cestoda & Cestodaria. Pub. Health. & Mar.

Hosp. Service of U.S., Hyg. Lab., Bull. 85, 1911.

(28) ZscHoKKe.—‘ Die Cestoden der Marsupialia und Monotre-

mata.” Semon’s Zoolog. Forschungsreisen in Australia

und dem Malayischen Archipel, 1898.

(27)

(29) Zool. Anz. 1898, Bd. xxi. (‘* Weitere Untersuch-

ungen an Cestoden aplacentaler Siiugethiere.”)

(30) Centralb. f. Bakt., Orig.-Bd. xxxvi., 1904. (‘‘ Die

Darmcestoden der Amerikanische Beuteltiere.”

(31) Centralb. f. Bakt. Abt. 1, Orig.-Bd. xliv., 1907.

(‘ Moniezia diaphana, n. sp. Hin weiterer Beitrag zur

Kenntnis der Cestoden aplacentaler Siiugethiere.”)

EXPLANATION OF THE PLATHS.

Lettering.

@!=Cirrus: Vit.Gl.=Vitelline gland.

C.P.=Cirrus-pouch. Res.= Ovarian reservoir.

Ex.C.=Excretory canal. L.Ex.= Longitudinal excretory canal.

G.A.=Genital atrium. Ex.P.=Excretory pore.

F.G.P.=Female genital pore. M.L.= Muscular layers.

M.G.P.=Male genital pore. V.D.=Vas deferens.

Ut.=Uterus. Ves.Sem.=Vesicula seminalis.

R.S.=Receptaculum seminis. L.L.= Lateral lappet.

Vag.= Vagina. Sk.=Suckers.

Sp.M.=Sphincter muscle. V.Ex.= Ventral excretory canal.

Test.=Testes. D.Ex.= Dorsal excretory canal.

S.T.=Spongy tissue. N. or L.N.=Longitudinal nerve.

L.M.= Longitudinal muscle-layer. Ov.=Ovary.

Ov.'T.=Ovarian tubules. E.Sh.=Fimbriated sheath.

T.Ex.=Transverse excretory canal. C.M.=Circular muscle-layer.

Sh.G].=Shell-gland. Ov. R.=Rudiments of female glands.

Prate I.

Cittotenia lagorchestis.

Fig. 1. Portion of strobila showing lateral projections of segment margins. The

two uteri distinct in the earlier sezments and meeting in median field

in older proglottides. Scolex with suckers, and no rostellum.

432 MR. R. C. LEWIS ON

Fig. 2. Coronal section through head and first 25 segments, passing through suckers

in head region.

3. Coronal section through middle of head between suckers showing origin of

genital glands immediately behind head; first sezments commencing

immediately behind head. Large receptaculum seminis. Excretory

ducts running up into head. Cirrus-pouch in some segments.

4. Transverse section of head across suckers.

Puate II.

Cittotenia lagorchestis.

Vig. 5. Transverse section of very young proglottis showing vagina as 2 thick-walled

duct below the cirrus-pouch.

6. Transverse section of an older proglottis showing vagina running out to

open into genital atrium.

7. Coronal section of a number of ripe proglottides.

Prats III.

Cittotenia lagorchestis.

Fig. 8. Transverse section showing transverse excretory canal connecting the larger

ventral longitudinal canals.

¢. Transverse section of proglottis, slightly older than in fig. 6, showing vagina

which has lost its connection with the genital atrium and is becoming

abortive.

10. Coronal section showing ovarian tubules arranged in a circle and opening

by narrow ducts into reservoir.

PratE IV.

Cittotenia tagorchestis.

Fig. 11. Enlargement of fig. 7, showing ovarian tubules, development of uterus, vitel-

line gland, shell-gland, testes, genital atrium, sphincter muscle, cirrus-

pouch, cirrus, spongy tissue, receptaculum seminis, transverse and longi-

tudinal excretory vessels, and longitudinal muscles.

12. Coronal section of amaturer portion of strobila showing uteri nearly meeting

in median field. Disappearance of genital glands except cirrus-pouch

and receptaculum seminis.

13. Coronal section of tail end showing reproductive organs to within twelve

segments of posterior end.

Puate V.

[Cittotenia lagorchestis.

Fig. 14. Coronal section same as fig. 12, showing protruded cirri.

15. Coronal section of posterior end showing excretory pore.

Cittotenia villosa.

16. Strobila showing fimbriations and liver-fluke shape.

Prats VI.

Cittotenia lagorchestis.

Fig. 17. Longitudinal section in region of cirrus-sac showing testes and receptaculum

seminis.

18. Transverse section of mature proglottis showing uteri nearly meeting in

median field.

Cittotenia villosa.

19. Transverse section of head through suckers showing’ quadratic cross-section

as contrasted with tig. 4.

20. Coronal section of head passing through suckers. Absence of neck and

absence of reproductive organs ; longitudinal excretory canals. Segmenta-

tion immediately behind head.

Fig. 24.

25.

26.

Fig. 27.

28.

29.

Fig. 30.

31.

TWO NEW MAMMALIAN CESTODES. 433

Jer Aga WIN.

Cittotenia villosa.

Enlargement of head showing excretory canals running up into head

suckers, and ornamented fimbriation.

Whole mount of a portion of strobila enlarged showing hair-hke

projections.

Coronal section showing male and female genital pores, uterus, cirrus, and

vagina.

Prate VIII.

Cittotenia villosa.

Coronal section showing uterus, vagina, cirrus-pouch, and cirrus.

Transverse section through young proglottis. | Well-developed sub-

cuticular musculature, absence of reproductive organs.

Transverse section showing protruded cirrus, sphincter muscle, cirrus-

pouch, portion of vas deferens and of vesicula seminalis.

Puate IX.

Cittotenia villosa.

Transverse section showing uterus in lateral margin.

Transverse section showing the two large receptacula seminis.

Egg with horns of pyriform body crossing.

PLATE X.

Cittotenia villosa.

Enlarged transverse section showing testis sac, part of coiled vas deferens

and of yvesicula seminis, cirrus-pouch, and cirrus.

Same series as fig. 30, showing the connection of vas deferens with cirrus-

pouch.

PZ. SS) 1914.0 DONOGHUE am

8 i

&) x

& ky

& & 2 4

\ B ei

Fi Fy

% F -

Bale & Danielsson, Lt amp

(G Jet. O Donoghue del.

MAINO S SsStS III OW SSALILWOIME CuNINTICUILYA.

PZ. S: 1914. 0 DONOGHUE soianne

ES MAL PLS SAA TEE AS SOOTNS SRA AD iE EON

aes

L.A,

VENOUS )SYSTE Me OF 1S @ Vin enUiviene Angelonia

ON THE CIRCULATORY SYSTEM OF ELASMOBRANCHS. 435

30. Notes on the Circulatory System of Elasmobranchs.

I. The Venous System of the Dogfish (Seyllium

canicula). By Cuas. H. O’Donocuus, D.Sce., F.Z.S.,

Senior Assistant in the Zoological Department, Uni-

versity College, London.

[Received April 2, 1914: Read May 19, 1914.1]

(Plates I., II. * & Text-figures 1-4.)

INDEX. Page

Ve JUMAROCIHG BOM ois dcdcesccscsc ess pecadadpseosenocen eat)

ee Developmenti eerste eee LOO

III. The Pre-Cardiac Vessels ..................... 488

IV. The Post-Cardinal Vessels .. ..... ......... 443

Vere lateral vieinsteees ess ee errr ee eee a>

VI. The Sub-Intestinal Vessels .................. 447

A. The Hepatic Portal Vein ............ 448

B. The Renal Portal Veins ............... 452

Walle her CoronanyaVierise eer cette eee) 402

WIII. The Ductus Cuvieri ........................... 452

IX. List of References.............................. 453

I, InrRODUCTION.

The foundation of our modern knowledge of the circulatory

ystem of Hlasmobranchs was laid by T. J. Parker in his

work on the venous system of the skate (Raia nasuta, 9), and

extended by his masterly description of the blood-vessels of

Mustelus antarcticus, in 1886 (10). This latter provides a full

account of the researches prior to that date, and also a full

bibliography.

Since that date only two papers have dealt with the subject.

Rand and Ulrich have discussed the posterior connections of the

lateral vein in the skate in 1905 (14), and Diamare the splanchnic

arteries and veins in Scylliwm catulus and Torpedo marmorata (2).

Since Balfour’s account (1) the general development of the

venous system in Elasmobranchs has been referred to by a number

of authors, including Hoffmann (5), Rabl (11 & 12), and Hoch-

stetter (3 & 4), and the development of the head veins by Grosser

(2 a), so that it is only necessary here to refer quite briefly to the

stage in a late embryo so as to indicate the order in which the

veins in the adult are described.

Although many dogfish have been dissected, there is no com-

plete account of the venous system. The descriptions given in

the text-books for the most part appear to be based upon

T. J. Parker’s work already referred to, and are not altogether

* For explanation of the Plates see pp. 454, 4565.

Proc. Zoou. Soc.—1914, No. X XIX. 29

436 MR. C. H. 0 DONOGHUE ON THE

satisfactory. Although the general disposition of the main vessels

in Mustelus antarcticus is somewhat similar to that in Seylliwm

canicula, the two species differ considerably in detail in the veins

and, as might be expected from the modification of form undergone

by the skates, the veins of these are markedly different from those

in the dogfish. In view of this and of the fact that Sceyllium

is frequently used as a type in the laboratory, the venous

system of the latter appears worthy of description.

An account of the arteries, on the other hand, is not so

necessary, as they do not vary nearly so markedly from those

described in other Elasmobranchs as do the veins and, moreover,

they are much more satisfactorily dealt with in text-books.

The animals were investigated by means of injection and series

of transverse sections through embryos of 37 and 56 mm. length,

and serial sections through a frozen adult were also examined

in order to check certain points. The injection-fluids used

were the gelatine mass, recommended by Tandler (15) for the

finer details, and the starch mass advised by Kingsley (6) for

the larger vessels. In a previous communication on the blood-

vessels of the grass-snake (7) I recommended using a mixture of

the solid matter that settles to the bottom of the starch mixture

with about twice its volume of gelatine, and I have now found

that an almost identical mixture was suggested for the blood-

vessels of the skate some time previously by Rand (18).

Perhaps the most striking feature of the venous system of

Scyllium is the dilatation of the vessels to form sinuses. These

sinuses render the injection of the system as a whole impractical,

and the indefinite nature of their walls makes it extremely

difficult to trace their exact course and extent with certainty

or to locate the points of entry of the smaller tributaries.

For the sake of ready reference the names of the vessels

adopted by Parker have been used as far as_ possible, and

wherever alteration has been made Parker’s terminology is also

given.

II. DEVELOPMENT’.

The pre-cardiac part of the venous system at a fairly late

stage of development is represented by the anterior cardinal

vein. This is morphologically composed of two sections: from

the ear back to the ductus Cuvieri it is the persistent anterior

cardinal, but the front part of this vessel has been replaced by

the lateral cephalic vein. It is convenient, however, to speak

of the whole trunk in the adult as the anterior cardinal sinus.

In the adult an inferior jugular vein is developed on each side

in addition to the foregoing sinus.

In the post-cardiac portion two distinct systems are to be

recognised : first, the posterior cardinal veins, and second, the

sub-intestinal vein. The posterior end of the first forms a loop

around the growing mesonephroi, and later the median portions

of the two loops fuse to constitute the inter-renal section of the

CIRCULATORY SYSTEM OF ELASMOBRANGHS, 437

definitive posterior cardinal sinuses. The second divides into

two parts—an anterior which gives rise to the hepatic portal

vein, and a posterior which acquires a connection with the outer

side of the kidney loop and forms the renal-portal vein.

Text-figure I

Su.

NHS.

Rea

sate

Diagram of the vessels in a fairly late embryo of Seyllium canicula.

V.C.L. Vena capitis lateralis. S.I. Sub-intestinal vein. V.I. Inter-renal vein.

For explanation of other lettering see p. 454.

Adopted and modified from Rabl (11),

According to Rabl (11), quoted also by Hochstetter (4), the

sub-clavian vein is described and figured as opening into the

posterior cardinal sinus, whereas in Seylliwm, in Mustelus, and

in the skates, this vein opens into the ductus Cuvieri, and

in the two former it opens into the ductus between the

point of entry of the anterior cardinal sinus and the sinus

29%

438 MR. C. H. 0’ DONOGHUE ON THE

venosus. In the adult Scylliwm and Mustelus the vessel opening

into the posterior cardinal vein in the position of that described

by Rabl as the * sub-clavian ” is really the sub-scapular.

In an embryo of 56 mm. it is quite easy to trace the relations

of these two vessels, which are found to be the same as in

the adult. The sub-clavian vein, formed by the union of

the brachial and lateval abdominal veins, opens into the sinus

venosus on its ventral side nearer the middle line than the

opening of the anterior cardinal sinus. The sub-scapular vein,

into which the lateral cutaneous vein opens, joins the posterior

cardinal vein in about the position indicated by the ‘‘ sub-

clavian ” vein of Rabl.

A sub-clavian vein is figured by Grosser (2 a, p. 184, fig. 4) in

the same position in an embryo of only 26 mm., although its

relation to the brachial and lateral abdominal veins is not shown,

nor is it identified as sub-clavian.

Rabl himself says (11, p. 233): “In den proximalen Theil

des Cardinalvenensinus miindet die Vena subclavia ein. Aus-

serdem tritt in die untere Wand des Ductus Cuvieri eine

Vene, die nach hinten bis zu den Bauchflossen zu verfolgen ist,

und deren Lage unter der Musculatur der vorderen Bauchwand

es nicht zweifelhafterscheinen lisst, dass man es hier mit der

Parietal-oder Seitenvene zu thun hat. Auf diese Vene habe ich

im Szshema keine Riicksicht genommen.”

It will be seen, then, that Rabl recognises two veins, but I

venture to think that ne has misinterpr eted one of them and in

ealling it the sub-clavian vein has suggested a connection with

the pectoral fin which does not exist. A muddle, probably re-

sulting from this, is to be found in nearly all the text-book

descriptions of Scyllium, in which it is stated that the sub-

clavian vein enters the lateral border of the front end of the

posterior cardinal sinus and brings back blood from the pectoral

fin. In all Elasmobranchs described, and also in Seylliwm, the

blood is returned from the pectoral fame by the brachial veins.

These open into the lateral abdominal veins, and the common

trunks so formed, which may be justly fommed sub-clavian veins,

open into the deans Cuvier.

For descriptive purposes it is easy to divide the venous system

according to its derivation from the embryonic condition just

described into pre-cardiac vessels, post-cardinal vessels, and sub-

intestinal vessels, to which must be added the lateral veins, the

coronary veins, and the cutaneous veins.

III. Tse Pre-CarpiAc VESSELS.

1. The Orbital Sinus.

1. i. The Nasal Sinus.

1. 1. The Orbito-Nasal Vein.

J.in. The Anterior Cerebral Vein.

l.iv. The Inter-Orbital Vein.

CIRCULATORY SYSTEM OF ELASMOBRANCHS, 439

2. The Post-Orbital Sinus.

3. The Anterior Cardinal Sinus.

3. i. The Hyoidean Sinus.

3. u. The Posterior Cerebral Vein.

3,11. The Myelonal Veins.

3.1v. The Nutrient Branchial Veins.

A, The Inferior Jugular Sinus.

v g

1. The Orbital Sinus (Parker) [ Pl. I., Or.] is a large irregular

sinus occupying the whole of the cavity of the orbit not occupied

by the eyeball and its muscles. It is easily recognised as a blood

sinus, for it almost always contains a considerable amount of

clotted blood in the freshly-killed animal and even in preserved

specimens.

It receives, in addition to the orbito-nasal vein, the anterior

cerebral vein, which enters it on its lateral wall a little way

behind the point of origin of the inferior oblique muscle. The

two orbital sinuses communicate with one another by means of

an inter-orbital vein.

1.i. The Nasal Sinus [Pls. 1. & II., N.S.j is a well-marked

crescent-shaped sinus situated somewhat ventrally on the

inner and hinder margins of the olfactory sac. ‘The two sinuses,

although approximating very closely in the middle line, are

completely separated by the cartilaginous inter-nasal septum.

They colleet the blood from the snout and olfactory sacs by a

number of more or less indefinite trunks, some of which come

from the anterior end of the roof of the mouth. They are figured

in an embryo of 26 mm. by Grosser (2 a, fig. 4).

Lui. The Orbito- Nasal Vein [ Pls. I. & Te, O.N.] is a small but

distinct vein passing through a canal in the cartilage separating

the olfactory sac from the orbit and entering the latter at its

lower, inner, anterior corner by a well-marked orbito-nasal

foramen. It conveys the blood from the nasal to the orbital

sinus.

This vein appears to correspond to the anterior facial vein of

Parker, who states that in Mustelus antarcticus he was unable to

get a satisfactory injection of it. In Scylliwm it is fairly easy

to inject with gelatine mass from the orbital sinus by means of

a pancreatic canula which will jamb into the orbito-nasal

foramen.

l. iii. The Anterior Cerebral Vein (Parker) |Text-fig. 2, A.C.]

is formed by the union of three factors at the posterior dorso-

lateral border of the diencephalon, whence it passes directly

outwards through a foramen in the cranial wall and empties

itself into the orbital sinus. The anterior factor is itself com-

posed of two smaller tributaries, one of which starts on the

ventral surface of the olfactory lobe and passes forwards and

then upwards to the dorsal surface. Here it runs backwards to

440 MR. C, H. O DONOGHUE ON THE

the main trunk, just before joining which it receives the other

tributary coming from the prosencephalon.

The ventral factor collects blood from the lower surface of the

prosencephalon and diencephalon as far back as the optic chiasma

and passes dorsally into the main vein. The third factor isa

posterior one that passes behind the edge of the prosencephalon

to anastomose with its fellow. In the middle line this vessel

receives a vein from the optic lobes and a large number of

smaller ones from the anterior choroid plexus roofing the third

ventricle.

Text-figure 2.

AC.

Ach,

Pch

PC.

LM.

Sketch of the dorsal side of the brain of Scylliwm canicula, showing the

arrangement of the anterior and posterior cerebral veins.

For explanation of lettering see p. 454.

l.iv. The Inter-Orbital Vein [Pls. I & II., 1.0.] is a small but

well-marked vein running from one orbit to the other in a canal

in the basi-cranial cartilage, and it enters the orbit towards its

posterior end just in front of and slightly below the large

CIRCULATORY SYSTEM OF ELASMOBRANCHS. 44]

foramen through which the sixth and main branches of the fifth

and seventh cranial nerves leave the cranium. It is situated

directly under a well-marked ridge in the floor of the cranial

cavity immediately behind the pituitary body.

2. The Post-Orbital Sinus [ Pls. I. & II., P.O.] 1s a moderate-

sized vessel placing the orbital sinus in communication with the

anterior cardinal sinus. It passes backwards from the posterior

external part of the orbit around the auditory capsule dorsally to

the spiracle. Its path around the capsule is indicated by a well-

‘marked groove, the post-orbital groove, which is to be found

between the ridge formed by the horizontal semi-circular canal

and the smooth surface for the articulation of the hyomandibular

cartilage.

3. The Anterior Cardinal Sinus* (Jugular Vein, Parker)

[Pls. I. & IJ., A.C.] is a very large and irregular sinus running

along the internal dorsal ends of the gill-clefts. It starts close

behind the spiracle at the anterior end of the first gill-cleft and

runs toa point a short way behind the fifth gill-cleft. At its front

end it receives the post-orbital sinus. The two anterior cardinal

sinuses are situated at about the same depth below the dorsal

surface of the animal as the vertebral column, with which they

are approximately parallel. The pharyngeo-branchial cartilages

project into the floor of the sinuses, forming well-marked ridges

which possess membranous flaps. The sinuses are readily exposed

from the dorsal side of the fish when making a dissection of the

cranial nerves, and, indeed, the branchial branches of the tenth

cranial nerves pass freely through their cavities, while the visceral

branches of the same are partially embedded in their mesial

walls. Just behind the fifth gill-cleft the anterior cardinal sinus

narrows down considerably, and passing ventrally opens into the

posterior cardinal sinus by an opening provided with a valve

(vide footnote, p. 444).

The anterior cardinal sinus receives small tributaries from the

dorsal ends of the gill-bars and from the surrounding musculature,

but their position is extremely difficult to ascertain owing to the

indefiniteness and irregularity of the walls of the sinus. At its

front end the anterior cardinal sinus receives the posterior

cerebral vein and the hyoidean sinus.

3.1. The Hyoidean Sinus (Parker) [Pls. I. & II, H.] is a

moderate-sized vessel situated in front of the first gill-cleft and

running parallel with it from the dorsal to the ventral side of the

fish. It lies in a shallow groove on the external side of the hyo-

mandibular cartilage, and at its dorsal end enters the anterior

cardinal sinus near the point where the latter receives the post-

ovbital sinus. The ventral end joins the inferior jugular sinus —

at the level of the posterior end of the thyroid gland,

The hyoidean sinus doubtless receives the blood from the

tissues surrounding it, but it is extremely difficult to make out

_* This name is used because, as Parker points ont, the vein is not in any way

homologous with the jugular vein of Iigher animals.

442 MR. C. H. O DONOGHUE ON THE

any definite tributaries corresponding to the nutrient branchial

veins and the posterior facial vein described by Parker in

Mustelus.

3. 11. The Posterior Cerebral Vein * (Parker) ['Text-fig. 2, P.C.]

commences at the anterior end of the cerebellum and passes back-

wards along its side. ‘Towards the posterior end of the cerebellum,

after receiving a well-marked tributary coming from its mid-dorsal

region, it runs out laterally over the sac-like lateral dilatation

of the posterior choroid plexus. Here it drains a remarkable

venous network, a kind of rete mirabile, composed of quite

well-marked veins. Its path now lies along the lateral border of

the median portion of the posterior choroid plexus, from which

it receives numerous tributaries, and just before the calamus

seriptorius it passes out through the cranium closely apposed to

the dorsal surface of the tenth cranial nerve. It parts company

with this nerve outside the cranium and apparently passes

through the muscles to the front end of the anterior cardinal

sinus, but it is difficult to follow. The fine veins from the

membranous labyrinth appear to open into the posterior cerebral

vein at its inner end. Parker describes and figures the veins

not as passing through the cranial wall but as running backwards

to unite and form the myelonal vein.

Grosser has figured this vein in an embryo of Sceyllium 26 mm,

long (2 a, fig. 4), but does not call attention to it in the text. In

dealing with Zriton and Salamandrina, however, he mentions

this vein as leaving the skull with the vagus nerve, and points out

that it is of general occurrence in many of the higher groups, a

fact also noted by Gaupp. It was readily made out in the sections

of embryos of 37 and 56 mm. length that were examined.

3. 11. The Myelonai Veins (Parker) [Text-fig. 2, M.]. Two

well-marked mvelonal veins are present, a dorsal and a ventral.

The dorsal myelonal vein runs the length of the spinal cord,

and at the anterior end forks just behind the calamus scriptorius.

The two limbs of the fork run into the posterior cerebral vein

just as it is leaving the cranial cavity. Parker describes this

vessel as forming a rhomboidal plexus in each vertebral segment,

but although the anterior end of the vein may be more or less

double for a short way behind the point where it divides, there

is no sign of this arrangement in Scylliwm. Segmental veins

are given off, and these often tend to form a lateral vessel on

each side of the cord by anastomosing in a longitudinal direction ;

it appears to be very irregular, however.

The ventral myelonal vein is formed by the union of two

* This vein and the anterior cerebral are very difficult to inject, as it is almost

impossible to insert even a hypodermic needle into them. However, I found that they

could be made distinct in the following way. After the anterior cardinal vessels

of a freshly killed fish have been quite filled with injection mass and plugged, the

cranium is dissected away so as to expose as much of the brain as possible. A

fixing fluid, corrosive formol, is slowly injected into the dorsal aorta until the

cranial arteries begin to get colourless and the veins distended with blood. If the

anterior end of the fish in this condition is placed in 5 per cent. formalin overnight,

it will be found that the blood in the veins has coagulated and become dark in colour.

CIRCULATORY SYSTEM OF ELASMOBRANCHS. 443

branches. Each commences by the side of the lobi inferiores just

behind the optic chiasma and passes backwards, draining the

saccus vasculosus, to unite behind the pituitary body to form a

median vessel. This runs the length of the spinal cord, giving

off small segmental branches.

3.iv. The Nutrient Branchial Veins (Parker) are a series of

four indistinct vessels on each side which bring back the

blood from the four holobranchs. Their anatomical relations

do not appear to be so constant and cannot be made out so

definitely as in Mustelus.

4. The Inferior Jugular Sinus (Parker) [Pls. I. & IL, I.J.] is

a moderate-sized, indefinite vessel situated below the floor of the

mouth. It commences as a small vein shortly behind the

symphysis of the lower jaw and passes backwards to the level of

the hinder end of the thyroid gland, where it receives the hyoidean

sinus and also anastomoses with its fellow. This anterier seg-

ment may perhaps correspond with the mandibular vein described

by Parker. The anastomosis between the two inferior jugulars

takes the form of an extremely irregular trunk passing along the

base of the thyroid gland (around which blood-clots are fr equently

to be found) and irowlniene the innominate arteries. From this

point the jugular sinus widens out considerably and runs along

the internal sides of the ventral ends of the gill-clefts, bathing the

proximal parts of the afferent branchial arteries, back to the wall

of the pericardium. Here it narrows and passes along the

wall to open into the proximal part of the ductus Cuvieri by an

opening common to it and the sub-clavian vein.

LV. THe Post-CAarDINAL VESSELS.

The Posterior Cardinal Sinus.

1. The Renal Veins.

2. The Genital Sinus.

A. The Ovarian Sinus.

B. The Spermatic Vein.

2. i. The Intestino-Mesenteric Vein.

. The Anterior Parietal Veins.

The Anterior Oviducal Sinus.

The Sub-scapular Sinus.

SR se

The Spinal and Hsophageal Veins.

The Posterior Cardinal Sinus (Parker) [Pls. I. & U., P.Ca.]

originates between the kidneys, where the two posterior comaline)

sinuses are united to form a median vessel. At the anterior end

of the kidneys the right and left sinuses are usually separated by

a partition passing frem the dorsal to the ventral wall in the

444 MR. C. H. O'DONOGHUE ON THE

middle line. Each cardinal sinus runs forwards as a fairly

narrow vessel to a point just in front of the anterior mesenteric

artery and then commences to widen out. At the anterior end

the two sinuses occupy the whole of the region dorsal to the

cesophagus right up to the pericardio-peritoneal septum and

outwards to the sides of the body. The condition of the septum

between the sinuses at the front end varies greatly in different

specimens. It may be practically absent, represented only by a

few strands, it may be well developed with perforations, or most

frequently it is in a condition between these two extremes. ‘The

posterior cardinal receives the renal veins, the genital sinus, the

anterior parietal veins, the anterior oviducal sinus in the female,

the sub-scapular sinus, the veins from the spinal cord, and at its

front end the anterior cardinal sinus and the ductus Cuvieri open

into it *.

The Renal Veins (Parker) are represented by a series of efferent

vessels leaving the kidneys. They do not appear as separate

vessels outside that organ, as the wall of the posterior cardinal

sinus is in contact with the mesial border of the kidney.

2. The Genital Sinuses differ in the two sexes and will be

dealt with separately.

A. The Ovarian Sinus is a large trunk composed of numerous

irregular factors and runs in the mesovarium dorsal to the ovary

from its posterior end. A very similar but smaller vein is

formed at the anterior end of the ovary, aud the two unite in the

first third of that body and pass dorsally into the posterior

cardinal sinus in the region where the anterior mesenteric and .

henogastric arteries are given off from the dorsal aorta. Just

before entering the posterior cardinal sinus it swells out some-

what and receives the intestino-mesenteric vein.

B. The Spermatic Vein (Parker) [Pl. I., R.S.] is subject toa

considerable amount of variation, and often each testis possesses

two separate veins. The posterior drains the hinder two-thirds

of the testis and, passing dorsally through the mesorchium, joins

its fellow of the opposite side to form a common genital sinus

which opens into the posterior cardinal sinus. The intestino-

mesenteric vein often joins the right spermatic vein just before

it unites with the left. The anterior spermatic vein drains the

front portion of the testis, in front of which it joins with its

fellow and opens into the posterior cardinal sinus. Sometimes,

perhaps more generally, this anterior spermatic vein is simply

a factor of the posterior one, which is always the main vein.

* It is stated that developmentally the anterior and posterior cardinal veins open

separately on the anterior and posterior sides of the ductus Cuvieri respectively.

That this description. given by Hoffmann (5), Hochstetter (8), Balfour (1), and

others, is the correct one can easily be verified by reference to a series of sections

of an embryo of S. canicula. In the adult, however, the actual anatomical

relations are different, possibly because of the dilatation of the veins to form

sinuses, and we find that the anterior cardinal sinus opens into the posterior cardinal ,

sinus by an aperture guarded by a valve, and the ductus Cuvieri projects as a short

tube into the posterior cardinal sinus, opening therein by an oval aperture.

CIRCULATORY SYSTEM OF ELASMOBRANCHS. 445

1. The Intestino-Mesenteric Vein [ Pl. I., 1.M.] is a small but

nevertheless well-marked vessel that collects the blood from the

right side of the intestine in the region of the Spiral valve and

runs through the mesentery, from cant it receives branches, to

open into the genital sinus. In the male it flows into the branch

from the right testis shortly before this joins with its fellow in

the middle line.

3. The Anterior Parietal Veins (ant. spinal veins, Parker) come

from the myotomes of the body between the pericardio-per itoneal

septum and the anterior end of the kidney and flow into the

posterior cardinal sinus.

4. The Anterior Oviducal Sinus (ant. ov. vein, Parker) is a large

vessel situated around the oviduct in the region -of the oviducal

gland in the full-grown female. It is a very large sinus, quite

separate from the posterior cardinal sinus into which it opens at

its anterior end. In the immature female this sinus cannot be

detected, but in the adult, and more especially when the oviducal

gland appears to be active, it is obvious enough.

d. The Sub-Scapular Sinus { Pls. I. & IT., $.8.] is a small sinus

situated on the dorso-lateral aspect of the dogfish immediately

behind the fifth gill-cleft and just ventral to the dorsal end of

the scapular cartilage. It joins the posterior cardinal sinus on its

dorso-lateral edge towards the anterior end by one or two small

openings guarded bya valve. Into it opens the lateral cutaneous

vein. As is pointed out above this vessel is generally but in-

correctly termed the sub-clavian vein. .

6. The Spinal and Csophageal Veins. The veins from the

spinal cord in the body region flow into the posterior cardinal

sinus. A few small veins from the extreme front end of the

esophagus may also enter this sinus, but the main part of the

blood from the cesophagus is collected by a factor of the hepatic

portal system.

VY. THe LATERAL VEINS.

The Sub-Clavian Vein.

1. The Lateral Abdominal Vein.

2. The Iliac Vein.

1.1. The Femoral Vein.

2.11. The Cloacal Vein.

3. The Rectal Vein.

4. The Brachial Sinus.

The Sub-Clavian Vein |Pls. I. & I1., 8.C.]. The condition of

this vein in the embryo has already been noted. In the adult it

is a short trunk passing from the union of the lateral abdominal

vein and brachial sinus dorsally along the edge of the coracoid

cartilage, and it flows into the ductus Cuvieri through a common

opening with the inferior jugular sinus. — Its position justifies it

446 MR. C. H. O DONOGHUE ON THE

being termed the sub-clavian vein and, in addition, it is homo-

logous with the similarly named vein in Kana *.

1. The Lateral Abdominal Vein (Lateral Vein, Parker) [Pls. I. &

TL., L.A.] is a moderate-sized vessel that runs immediately beneath

the peritoneum along the ventro-lateral wall of the body-cavity.

It originates as a continuation of the iliac vein on the dorsal

side of the pelvic cartilage, across which it anastomoses with its

fellow of the other side. Thence it passes forwards in the body-

wall to the pericardio-peritoneal septum, in the wall of which it

turns very sharply dorsalwards and slightly mesially along the

posterior edge of the coracoid cartilage. A short distance along

this it unites with the brachial sinus to form the sub-clavian vein.

2. The Iliac Vein (Parker) [P1. I., Ll.] is a short vessel formed

by the union of the femoral and cloacal veins on the inner side

of the basipterygium towards its anterior end. It runs into the

lateral abdominal vein on the dorsal surface of the pelvic bar.

2.1. The Femoral Vein (Parker) [PI. L., F.] drains the major

part of the pelvic fin and is situated laterally and slightly dorsal

to the basipterygium. It passes across the anterior end of this

cartilage, which is slightly notched to receive it, to unite with

the cloacal vein.

2. ii. The Cloacal Vein (Parker) [Pl. I., Cl.] lies on the inner

side of the basipterygium, and is formed by the union of factors

from the posterior and lateral walls of the cloaca and also from

the inner side of the pelvie fin.

it will be noted that the arrangement of the factors of this

vein agree more. nearly with those described by Parker in

Mustelus than with those figured by the same author for Rata

nasuta, and those in: R. erinacea and R. levis according to Rand

and Ulrich (14).,. ‘The additional factors in the skates are

doubtless to be correlated with the greater relative size of the

pelvic fin. A similar difference is found in the brachial veins ;

only one such is present on each side in Scylliwm, while two are

found in FR. erinacea and K. levis and three in Ff. nasuta.

Further similarity with Mustelus is shown by the presence

of a pelvic anastomosis between the lateral abdominal veins in

Scyllium but not in the skates.

3. The Rectal Vein [P1. I., Re.| isa small short vessel joining

the anastomosis between the lateral abdominal veins in the middle

line. Its branches form a fairly rich network of vessels spread

over about the last one and a half inches of the rectum and the

body-wall ventral to this.

This vessel is not represented in Mustelus, where the posterior

end of the rectum is drained by a fairly large proximal tributary

of the cloacal vein on each side. It more nearly resembles the

* Tn the tadpole the sub-clavian vein is formed by the union of the musculo-

cutaneous, brachial, and the epigastric veins, the latter being homologous with the

lateral abdominal veins. In the adult the two epigastrics are reduced to a median

vein, the anterior abdominal, which acquires a secondary connection with the

hepatic portal vein, though in certain abnormal specimens the primitive connection

is retained (8).

CIRCULATORY SYSTEM OF ELASMOBRANCHS. 447

condition in R. erinacea and R. levis, where paired vessels from

the end of the rectum run straight into the lateral abdominal

veins.

4. The Brachial Sinus (Br. vein, Parker) (Pls. I. & IT., B.]

collects blood from the pectoral fin and passes along near its

posterior edge. It leaves the fin and penetrates the body-muscles

to open into the lateral abdominal vein on the posterior edge

of the coracoid cartilage.

The Cutaneous Veins.

1. The Lateral Cutaneous Vein.

9. The Posterior Ventral Cutaneous Vein.

1. The Lateral Cutaneous Vein (Parker) [ Pls. 1. & Int, 10)

originates far back in the tail, and forms a well-marked vessel

running to the region of the pectoral fin, in the connective tissue

immediately underlying the lateral-line canal. Here it passes

inwards and opens into the sub-scapular sinus. Parker describes

anastomoses between it and the caudal vein, and such anastomoses

ean be found in sections of embryos of 56 mm., but I have been

unable to inject and display them by ordinary dissection.

2. The Posterior Ventral Cutaneous Vein (Parker) can be seen in

transverse sections both of embryos and of adult fish. It runs

forward embedded in the connective tissue in the mid-ventral

line from the tail, forms a loop around the anal fin, and forks in

the region of the cloaca. A similar vein is to be found under the

eutis in the median line of the abdomen, and this doubtless

corresponds to the anterior ventral cutaneous vein of Parker, but

the exact relations of its anterior and posterior ends could not be

ascertained as it is too small for injection.

Serial sections through embryos of 37 and 56 mm. length have

been studied, and the general arrangement of these cutaneous

vessels is apparently similar to that in Mustelus as described by

Parker. They are not dealt with in detail here as they have only

been followed in the above sections, and they cannot be studied in

the adult by ordinary methods of injection.

he ventral cutaneous veins are too small to inject successfully,

even with a hypodermic syringe, and the dorsal cutaneous vein is

barely visible to the naked eye.

VI. THe SuB-INTESTINAL VESSELS.

A. The Hepatic Portal Vein.

1. The Posterior I ntestinal Vein.

9. The Posterior Lieno-gastric Vein.

9. i. The Posterior Splenic Vein.

9. ii. The Median Gastric Vein.

8. The Pancreatic Veins.

448 MR. CG. H. O DONOGHUE ON THE

4. The Gastro-intestinal Vein.

4. 1. The Intra-intestinal Vein.

4, ii. The Anterior Intestinal Vein.

4. i. The Anterior Lieno-gastric Vein.

. The Dorsal Anterior Gastric Vein.

5. i. The Dorsal Gastric Vein.

5. li. The Dorsal Gisophageal Vein.

6. The Ventral Anterior Gastric Vein.

6. 1. The Ventral Gastric Vein.

6. ii. The Ventral Gisophageal Vein.

7. The Hepatic Veins and Sinuses.

B. The Renal Portal Veins.

1. The Caudal Vein.

2. The Renal Portal Vein.

2. i. The Posterior Oviduecal Veins.

2.11. The Posterior Parietal Veins.

THe Hepartc Porran System.

The hepatic portal system consists of a number of large well-

marked veins, mostly lying in the gut mesenteries, which convey

blood from the whole of the alimentary canal (from cesophagus

to rectal gland inclusive) to the liver. In the higher verte-

brates all the blood collected from the gut is taken to the

liver, but in Scyllium there is an exception to this general rule

in the presence of an intestino-mesenteric vein. There are

marked differences between the component veins of this system

in Mustelus antarcticus, according to Parker, and in Seylliwm,

and in consequence the nomenclature here adopted is descriptive

and does not necessarily imply homology. The system may be

conveniently and easily injected from the main trunk near the

liver.

A. The Hepatic Portal Vein | Pl. 11., H.P.] is formed in the pan-

creas, a short distance from its posterior end, by the confluence

of the posterior intestinal and posterior lieno-gastric veins. It

runs partially embedded in the right dorsal edge of the pancreas

to the anterior end of that body, and receives during this part of

its course a number of small tributaries, the pancreatic veins.

At the anterior end of the pancreas it is joined by two large

veins; one, the dorsal anterior gastric, enters it on the right,

and the other, the gastro-intestinal, enters it somewhat ventrally

on the left. From this point it runs for a short distance, about

1:5 cm., in the gastro-hepatic omentum before it receives its last

large tributary, the ventral anterior gastric vein. It is now an

extremely large vein with a diameter, when fully distended, of

CIRCULATORY SYSTEM OF ELASMOBRANCIS. 449

about 5 mm.,and it quickly divides into two main branches,

a right and a left, one feeding each lobe of the liver.

Text-figure 3.

VAG.

_vO.

¥ jess DO!

DAG

ink

Gl.

Pl.

Diagram of the factors of the Hepatic Portal System in Scyllium cantcula,

viewed from the ventral side.

For explanation of lettering see p. 464.

1. The Posterior Intestinal Vein [ Pl. I1., P.I.] commences as a

small vein on the ventral side of the rectal gland, and then,

turning sharply upon itself, passes along the dorsi-lateral wall of

the intestine to a point just posterior to the caudal end of the

pancreas. At this place, marked also as the point at which the

anterior mesenteric artery reaches the gut-wall, the posterior in-

testinal vein receives a fairly large factor from the anterior end

of the intestine, and then it runs freely to join the posterior

lieno-gastrie vein in the pancreas. During its course along the

intestine the vein receives a number (usually eight) of well-

marked paired tributaries, whose position on the outside of the

intestine marks the line of insertion of the folds of the spiral

valve within.

450 MR. C. H. O'DONOGHUE ON THE

2. The Posterior Lieno-gastric Vein | Pl. I1., P.L.G.] is formed

dorsally to the gut by the union of the posterior splenic and

posterior gastric veins, and it runs from this junction to the

posterior end of the pancreas and along the right dorsal edge of

this gland, until it unites with the posterior intestinal vein to

give rise to the hepatic portal vein.

2.1. The Posterior Splenic Vein | P\.I1., P.S.] is fairly large and

situated dorsally to the posterior end of the cardiac division of

the stomach. It collects blood from the ‘posterior portion of the

spleen, and it receives in addition one or two small branches from

the stomach.

2,11. The Median Gastric Vein [P\. II., M.G.] is also fairly large,

and it is formed by the union of several branches from the

posterior dorsal region of the cardiac division of the stomach.

It; leaves the stomach-wall at the point where the lieno-gastric

artery joins it, and it runs by the side of this artery until it

unites with the posterior splenic vein.

3. The Pancreatic Veins are a number of small tributaries that

flow into the hepatic-portal vein in its course along the edge of

the pancreas.

4, The Gastro-Intestinal Vein [Pl. I1., G.I.] is a short vein,

hidden away in the fold between the pylorus and intestine, which

commences about the middle of the anterior edge of the pancreas

and runs along this edge into the hepatic-portal vein. It is

formed by the union of the intra-intestinal, the anterior lieno-

gastric, and anterior intestinal veins, and in addition receives one

or two branches from the pyloric thickening.

4. i. The Intra-Intestinal Vein [Pl. II., I.1.] is a large vein

bringing back the blood from the spiral valve. ‘The valve is

extremely well supplied with large capillaries, and numerous

branches in each fold feed the intra-intestinal vein which runs in

the central core of the spiral valve. It perforates the wall of the

intestine at the anterior end of the spiral valve close against the

pyloric valve, and it emerges from the intestinal wall in the sharp

bend between the intestine and the pyloric thickening, where it

quickly unites with the anterior intestinal and anterior lieno-

gastric veins.

A. ii. The Anterior Intestinal Vein | Pl}. I1., A.I.] is of moderate

size, and originates in the line of insertion of the first fold of the

spiral valve in the intestine towards the dorsal side of the latter.

It follows the valve round and comes through the intestinal wall

on its ventral side close to the anterior lobe of the pancreas.

Here it receives one or two branches from the wall of the ventral

part of the anterior end of the intestine and runs straight to the

pancreas, where it unites with the anterior lieno-gastric vein.

4.11. The Anterior Lieno-gastric Vein | Pl. II., A.L.G.] is situ-

ated between the anterior lobe of the spleen and the pyloric

division of the stomach, and receives tributaries from both these

bodies. It arises near the level of the division between the two

parts of the stomach and runs to the beginning of the bend

CIRCULATORY SYSTEM OF ELASMOBRANCHS. 451

between the intestine and the stomach, where it branches off

to the anterior lobe of the pancreas.

5. The Dorsal Anterior Gastric Vein | Pl. I1., D.A.G.] is formed

by the union of the dorsal gastric and dorsal csophageal veins,

and runs in the mesentery from the dorsal anterior side of the

stomach into the hepatic-portal vein just as the latter is leaving

the front dorsal end of the pancreas.

Diamare (2) describes and figures a fairly large anastomosis

between the dorsal anterior gastric vein (vena gastrica dorsalis

anterior) and the median gastric vein (vena gastrica media) in

Seyllium catulus. I have examined eight specially injected speci-

mens of S. canicula and numerous fresh ones, but have been

unable to find this anastomosis. A large branch runs forwards

to join the dorsal anterior gastric vein, and a smaller one

originating near it runs backwards into the median gastric vein

(vide fig. 1, Pl. I1., P), but the two do not anastomose by any

well-marked vessel such as Diamare figures, although they are

indirectly in communication by means of capillaries.

5.1. Lhe Dorsal Gastric Vein [Pl. I1., D.G.j is a well-marked

vein composed of two main tributaries, which collect the blood

from the right and left sides of the anterior two-thirds of the

cardiac division of the stomach.

5.11. Lhe Dorsal Gsophageal Vein [ Pl. I1., D.O.] brings back

blood from the dorsal side of the cesophagus.

6. The Ventral Anterior Gastric Vein [ Pl. II., V.A.G.] is a large

trunk receiving one branch from the stomach, the ventral gastric

vein, and one branch from the cesophagus, the ventral cesophageal

vein. It leaves the stomach-wall on its ventral side, anterior to

the point of departure of the dorsal anterior gastric vein, and it

runs in the mesentery to flow into the hepatic-portal vein shortly

before the latter divides into its right and left branches.

6.1. The Ventral Gastric Vein (Pl. II., V.G.] is a conspicuous

vessel formed by the confluence of a number of tributaries from

the ventral side of the anterior two-thirds of the cardiac division

of the stomach.

6.11. The Ventral Wsophageal Vein [ Pl. I1., V.O.] collects the

blood from the ventral side of the cesophagus.

The esophagus has a very rich plexus of large capillaries

similar to that described by Parker (10) in M/ustelus, save that in

Mustelus the blood from this plexus is taken to the posterior

cardinal sinuses, while in Seylliwm it goes to the hepatic-portal

vein in the manner described above.

7. The Hepatic Veins and Sinuses (Pls. I. & II., H.S.]. The

hepatic vein is a large thin-walled venous trunk situated at the

anterior end of the corresponding lobe of the liver. The two

hepatic veins unite just outside the liver to form a large sac,

the hepatic sinus, which, when dilated, entirely fills up the space

between the ventral body-wall, the cesophagus, the anterior ends

of the lobes of the liver, and the pericardio-peritoneal septum.

Proc. Zoou. Soc.—1914, No. XXX. 30

452, MR. C. H. O DONOGHUE ON THE

This sinus is partially divided into two chambers by an incom-

plete vertical septum formed of interlacing trabecule. It passes

through the pericardio-peritoneal septum and opens into the

sinus venosus, however, by two small circular apertures, one

on either side of the middle line, by means of which all the

blood brought to the liver by the hepatic-portal vein and the

hepatic arteries is returned to the heart.

B. THe Renat Porta VEINs.

1. The Caudal Vein (Parker) [Pl. I., C.] originates far back in

the tail and runs forwards in the hemal canal ventrally to the

caudal artery to a point just posterior and dorsal to the anus.

Here it leaves the vertebral column and divides into two large

equisized branches, the renal-portal veins. It receives numerous

small branches from the myotomes of the tail.

2. The Renal Portal Vein (Parker) [ Pl. I., R.P.] starts from

the bifureation of the caudal vein and passes forward along the

dorsal and dorso-lateral edge of the kidney, to which it sends

numerous afferent renal branches. It gradually diminishes in

calibre, and dies away towards the anterior extremity of the

eaudal mesonephros at about the level of the front end of the

vesicula seminalis in male. ‘The two renal-portal veins are

completely separated in the middle line, and do not communicate

directly with the posterior cardinal sinus.

2.1. The Posterior Oviducal Veins (Parker) are small veins

from the dorso-lateral wall of the posterior portion of the oviduct,

and open into the renal portal vein.

2. ii. The Posterior Parietal Veins (p.-spinal veins, Parker)

arise from the myotomes of the region of the body along the

side of the kidney, and open into the renal portal vein on its

dorsal side.

VII. Tur Coronary Verns (Text-fig. 4).

The Coronary Veins are situated one on each side of the heart.

Each of them is formed at the posterior end of the furrow

separating auricle and ventricle, and enters the sinus venosus

just behind the corresponding flap of the sinu-auricular valve.

The posterior smaller vein collects the blood from the caudal end

of the ventricle, while the remaining one collects blood from the

anterior part of the ventricle, and, after receiving a well-marked

tributary from the conus arteriosus, runs backwards in the

groove between auricle and ventricle. - Small factors from the

auricle probably join the coronary veins, but are difficult of

injection.

VIII. Tue Ducrus Cuvier.

The Ductus Cuvieri (Parker) [Pls. I. & IJ., D.C.] convey to

the heart all the venous blood save that brought by the hepatic

veins. Their anatomical relations in the adult have already been

briefly noted. The outer end forms a spout-like structure with an

oval end projecting into the posterior cardinal sinus. Each passes

CIRCULATORY SYSTEM OF ELASMOBRANCHS. 453

inwards almost horizontally on the ventro-lateral sides of the

cesophagus through a very conspicuous notch in the posterior

border of the fifth cerato-branchial cartilage up to the lateral

wall of the pericardium. It is continuous through this with the

sinus venosus.

Text-figure 4.

acer enn Ned &,

Co.

Lateral view of the heart to show the arrangement of the coronary veins.

For explanation of lettering see p. 454.

1X. List or REFERENCES.

1. Banrour, F. M.—‘“‘ The Development of Elasmobranch

Fishes.” Journ. Anat. & Physiol., 1876-1878.

2. Diamare, V.—“ Sw’ rapporti della vena porta e delle arterie

splaneniche in Seylliwm catulus e Torpedo marmorata.”

Anat. Anzeig. Bd. xxxiv., 1909.

2a. GRossER, O.— Die Elemente des Kopfvenensystems der

Wirbeltiere.”- Verh. Anat. Ges. Wiirzburg, 1907, Anat.

Anz. Erganzungsh. zu Bd. xxx.

3. Hocusterrer, F.—‘‘ Vergleichenden Anatomie und Entwick-

lung des Venensystems.” Morph. Jahrb. Bd. xiii., 1888.

4. Hocusterrer, F.—‘‘ Die Entwicklung des Blutgefisssystems.”

In Hertwig’s Handbuch der Entwicklungslehre der

Wirbeltiere, 1901—06.

5. Horrmann, O. K.—“‘ Zur Entwickelungsgeschichte des Venen-

systems bei den Selachiern.” Morph. Jahrb. Bd. xx.,

1893.

6. Kinestry.—Guides for Vertebrate Dissection. New York,

Ore

7. O'DonocHuE, C. H.—‘‘ The Circulatory System of the

Common Grass-Snake.”+ Proc. Zool. Soc. 1912.

8. O’'Donoeuur, C. H.—‘‘ Two Cases of Abnormal Heart and

One of an Abnormal Anterior Abdominal Vein in the

Frog.” Zoolog. Anzeig. Bd. xxxvii., 1911.

30*

45 MR. C. H. O'DONOGHUE ON THE

9. Parker, T. -J.—‘‘ On the Venous System of the Skate (Raia

nasuta).” Trans. & Proc. New Zealand Inst. vol. xiii.

1881.

10. Parker, T. J.—‘‘ The Blood-Vessels of Mustelus antarcticus.”

Phil. Trans. Roy. Soc., 1886

11. Rasr, C.—“ Ueber die Entwickelung des Venensystems

der Selachier.” Festschr. z. 70. Geburtstag R. Leuckarts,

1892.

12. Rasy, C.—‘‘ Theorie des Mesoderms.” Morph .Jahrb. Bd.xix.,

1892.

13. Rano, H. W.—“ The Skate as a Subject for Classes in

Comparative Anatomy.” American Naturalist, vol. xxxix.,

1905.

14. Rann, H. W., and Utricu, J. L.—‘ Posterior Connections

of the Lateral Vein of the Skate.” American Naturalist,

WOlls sconbx,, OS.

15. Tanpier, J.—‘t Mikroskopische Injectionen mit kaltflussiger

Gelatin.” Zeitsch. f. wiss. Muikros. u. Mikros. Tech.,

NGO:

EXPLANATION OF THE PLATES.

Lettering.

A.C. Anterior Cerebral Vein. L.M. Lateral Myelonal Vein.

A.Ca. Anterior Cardinal Sinus. M. Myelonal Vein.

A.Ch. Anterior Choroid Plexus. M.G. Median Gastric Vein.

A.Co. Anterior Coronary Vein. N.S. Nasal Sinus.

A.F. Anterior Factor of A.C. O. Opening of the Ductus Cuvieri.

A.J. Anterior Intestinal Vein. O.C. Olfactory Capsule.

A.L.G. Anterior Lieno-gastric Vein. O.N. Orbito-Nasal Vein.

An. Anastomosis between Posterior Or. Orbital Sinus.

factors of A.C. P.An. Pelvic Anastomosis between

Au. Auricle. the two L.A.

B. Brachial Vein. P.C. Posterior Cerebral Vein.

Ba. Basipterygium. P.Ca. Posterior Cardinal Sinus.

C. Caudal Vein. P.Ch. Posterior Choroid Plexus.

Cl. Cloacal Vein. Pe. Pectoral Fin.

Co. Conus Arteriosus. P.I. Posterior Intestinal Vein.

D.A.G. Dorsal Anterior Gastric Vein. Pl. Pelvic Fin.

D.C. Ductus Cuvieri. | P1.C. Pelvic Cartilage.

D.G. Dorsal Gastric Vein. | P.2.G. Posterior Lieno-gastric Vein.

D.O. Dorsal (sophageal Vein. P.O. Post-Orbital Sinus.

HK. Eye. P.S. Posterior Splenic Vein.

¥. Femoral Vein. R. Portion of the PJ. on the

G.I. Gastro-Intestinal Vein. Rectal Gland.

G.S.2. 2nd Gill-cleft. Re. Rectal Vein.

H. Hyoidean Sinus. R.P. Renal Portal Vein.

H.P. Hepatic Portal Vein. R.S. Right Spermatic Vein.

H.S. Hepatic Sinus. | S.C. Sub-Clavian Vein.

1.1. Intra-Intestinal Vein. | Sp. Spiracle.

L.J. Inferior Jugular Sinus. S.S. Sub-Scapular Sinus.

J]. Ihac Vein. | S.V. Sinus Venosus.

I.M. Intestino-Mesenteric Vein. T.G. Thyroid Gland.

1.0. Inter-Orbital Vein. | V. Ventricle.

KX. Caudal Mesonephros. | Va. Valve between A.Ca. and P.Ca.

L.A. Lateral Abdominal Vein. | V.A.G. Ventral Anterior Gastric Vein.

L.C. Lateral Cutaneous Vein. | V.F. Ventral factor of A.C.

Le. Lateral expansion of the Pos- | V.G. Ventral Gastric Vein.

terior Choroid Plexus. | V.O. Ventral @sophageal Vein.

CIRCULATORY SYSTEM OF ELASMOBRANCHS. 455

Puate I.

Diagram showing the general disposition of the main venous trunks in Seyllium

canicula. The more dorsally situated vessels are stippled and tie wore ventral ones

black. For the sake of clearness, the ventral cutaneous vein has been omitted.

Puate II.

Fig. 1. Sketch of the arrangement of the Hepatic Portal factors seen from the dorsal

side. The vessels were injected and the gut hardened and removed whole

from the body-cayity.

Fig. 2. Diagram of the main vessels of the anterior end of the Dogfish viewed from

the side in order to give their dorso-ventral relations.

A aasee Tt aahe i

2 eg

Gee tes Fearne Ahi SEN r ter ta

xe huet spice WAP ade OT pa

vist eet

: Pi eee

ON COLOUR DEVELOPMENT IN THE INDIAN WOOD-STORK, A457

31. Notes on Colour Development in the Indian Wood-

Stork. By Grorce Jeynison, Belle Vue Gardens,

Manchester *.

{Received April 17, 1914: Read May 19, 1914. }

On June 2nd, 1908, a specimen of Pseudotantalus leucocephalus,

which I judged to be six months old, came to Belle Vue Gardens

with other stock. It was at that time not more than half its

present weight, the beak was straight and rather slender, pale

yellow in colour. The pinions were of a dull black, the rest of the

plumage a yellowish white. The bird ate fish freely, and has not

had a day’s illness or check of any kind, so we may assume its

subsequent development to be normal. The first change was

noted in February 1909, when the feathers on the larger wing-

coverts were slightly tinged with pink. During the next three

years, that is until the spring of 1912, there was a continual

change of the yellowish to ever whiter body-feathers, for the

moult is extremely gradual and the new plumes make quite a

checkered pattern with the old: the dull black of the pinions and

tail took on a lustrous greenish tinge, and the bare skin of the

head became red where it meets the feathers of the neck.

These changes might pass unnoticed by a casual observer, but

in January 1912 a drastic alteration supervened which could not

fail to attract attention. The white feathers of the smaller and

median wing-coverts of the chest and underparts of the wings

were slowly replaced by black feathers, with a narrow edge of

white. The perfection of plumage was reached in May, when the

whole of the shoulder was a wavy pattern of brilliantly contrasted

broad black and narrow white, and the bird in flight showed the

same beautiful coloration on the chest and beneath the wings.

The beak changes slowly, first thickening and afterwards taking

a pronounced downward curve, which continues to develop long

after the plumage-change is completed ; its colour changes mean-

while to a deep shiny wax-like yellow. During this time the

skin of the head, which can be drawn back an inch or more, also

continues to develop until the forehead and cheeks are a deep

yellow-umber and the hinder part of the head becomes a rich

purple.

The legs are now (1914) showing signs of a remarkable

alteration ; for five years they were grey with a shght tinge of

pink on the toes, the upper half of the tarsus is now a deep

magenta-red.

* Communicated by the SECRETARY.

ERIE, RMT TSS. APS QUAI RE tetas Cee Suer eee gy, Tara aa ewe bis

ep Ma i heh eee THe ner ab fades crib vend A pada

eal

eh PLT kn hee, oT]

‘ - ‘

(a's Eaaee a Sa? 5 B

at pete: Ps ah ae a Ra

ay he P

hoe} i We ane sei wee |

; Sab uae, sacs PRI D ay i) ER ae ee ty

Shu ainp wank Perens, MPL) Ve tpt Oates t we yess "hee eke Holt 4 ea

ee: RVR Neorg Pg SES ee cau 8 Wy aI «One TO ER “yay rt

Mie ‘Wee ee eta Si Ae eet | ; ’ ec aa Yt OAR: Pee

seek Bye SMT: et oi eye PEL ee ele He Serta Aes

tree eee Ps oe cae evtng: LEER Goober ‘ehiiaahes edgy

iidbiihn eis £008 A yamine eile ae alee

bylh sd ih an , \ SOA ee See re og » ey tae erhy Ver

MR ROG Head ny’ Hae: be +: Dera a

Pee iats Meee a rede adh ig d.. F Ave at +o.

evieig s ey Fart

Pd Gots be anes

POLI eres ur aie

4 a

a tees Atte = Set eunees

bh athe el tt (itll) ane oe

Le i See iies eh Te ee ie Wrdkeen

aT tee Poult Pine es «Terie

f iar ee ee ae lesht Rie “ot +a y

oa fe ie

eee arg pias Ma, i # tee

At iy

{iste ee See be sy peerie

Ce a ae ait ee PHL) y i hicp os; 7 Last ty fs ; ee ue Bs alt

Rees yore: : Waals ee TY ETA Gaius . .

(Deut heh aa ee Re ire Gok vides divin pat ote ee

He a tenon. A inebhe, Tee gp cs 4 hae iat: Pe ta

th NaS girticath ‘eet; veh ri a get

AOR 9 gaara pate: ean a eR a LEE

Ae a be sth SS vgs Beas ss bri oa a rk

Find

ON SCENT ORGANS IN TRICHOPTERA. 459

32. Scent Organs in Trichoptera. By Brucn F. Cummines,

Baan Museum (Natural History) *.

[Received April 3, 1914: Read May 19, 1914. |

(Text-figures 1—8.)

InpEx.

Page

wal GHAROLC KDVELOIEY So netobbosceroocadodoccs dad eRpaueHer Ree EERE REE eto co nes 459

i. Historical . PI hte EO APE a te ath ead ans MR Re Cana Sn AO)

i. Scent Crema f in incecte af eneansieeoe a WOO),

ii. The Maxillary Palpi oo ho. TMleltes of s ericostoma

personatwm ......... a Nasr Tem et TER Mona AA

iv. The Scent Organs of Sop ioostome Pronsonaam eee AGS

We laiberatire ees eens Se cen reat nae mck Stes ara Hl AS

INTRODUCTION.

Sericostoma personatum Spence is a tolerably common caddis-

fly in Great Britain, and immediately attracts attention on account

of the enormous development of the palpi of the first pair of

maxille in line male. Unlike the maxillary pee of the female,

which are 5-jointed and quite normal (text-fig. 1), the maxillary

palpi of the male consist of but a single segment very much

enlarged and shaped like a half-moon. These two palpi are

placed together and held vertically so as to mask the front of the

head (text-fig. 2, p. 463).

Text-figure 1.

Sericostoma personatum.

Palpus of the first maxilla, 2. » 17.

Despite their bizarre shape, these palpi have not obtained,

so far as I am aware, that amount of enquiry into their

nature and function which they deserve, and a résumé of their

uneventful entomological history therefore will not detain us

long.

1, HisToricat.

Kirby & Spence (1) regarded them as the genz or cheeks of

the skull. Pictet (2), as McLachlan gravely points out, ‘ scarcely

* Communicated by the SecRETARY, and published by permission of the Trustees

of the British Museum.

4.60 MR. B. F. CUMMINGS ON

committed himself” to any direct statement as to the number of

segments, though it seems more than likely, from an expression on

p. 20 of his ‘ Recherches,’ that he regarded three as probable.

Stephens (3), p. 148, assumes two to be the number. Burmeister

(4) says that according to his experience the segments of the

maxillary palpi are only two in number. Rambur(5) agrees with

this, and makes the observation regarding the “ fluff ” on the inner

surface of the two palpi that it is perhaps not produced until some

time after the insect has emerged, as all specimens do not possess

it. Kolenati (6) arrives at the astonishing conclusion that there

are four segments, the second being galeate and the third and

fourth more slender. McLachlan (7) is cautious, and says ‘“‘ they

are probably 3-jointed, but the basal joint is scarcely separable

from the sides of the face and the second joint... . is ill-defined

and transverse.” A very brief and somewhat inaccurate descrip-

tion of the palpi follows. MclLachlan’s account appears to have

set the matter at rest in the minds of Trichopterists, for Ulmer

(8) in 1907 echoes the former’s opinion that three segments are

present, the first and second being ill-defined.

It is necessary to point out at once that, as will be seen on

examination of text-fig. 2, McLachlan mistook the cardo and

stipes of the first maxilla for segments of the palpi. A paper by

W. Miller (9) in 1887 appears to have been overlooked, for here

not only is the maxillary palpus described correctly as one-jointed,

but convincing evidence is brought forward showing that these

palpi serve as scent organs. Miller observed the large tuft of

elongate hairs (MclLachlan’s inelegant but expressive “ fluff”)

on the inner surface of the palpi and compared them with similar

hair-tufts in the males of some Lepidoptera.

During copulation, one individual was seen ‘seine Palpen

auseinander zi spreizen und die in denselben liegenden Haar-

biischel zu entfalten,” so that they surrounded the head ‘ wie

ein Heiligenschein.” A strong smell of vanilla was emitted.

ll. SCENT ORGANS IN INSECTS.

By “scent organ” the entomologist usually means those glands

which, secreting an attractive odour and being confined to the

male sex, are supposed at mating time to charm or stimulate the

females. Such scent glands, well known in the Lepidoptera, aie

situated at the bases of hairs arranged in tufts or at the bases of

specially modified scales (called androconia) (10). Typical scent

organs like these occur not only in the Lepidoptera (11) (12), but

also in Coleoptera (Blaps mortisaga) (13), Blattide (15), and, as

now appears, in Trichoptera (Sericostoma personatum).

Another type of scent gland occurs in insects. This assumes

the form of fairly long eversible tubular filaments, usually two

in number, and is known in the males of some Lepidoptera,

Spilosoma virginica, Arctia virgo, Haploé clymene (10), and in

the males of the Cricket, Hadenwcus subterraneus (17). Similar

SCENT ORGANS IN TRICHOPTERA. 461

retractile tubular filaments have not yet been described from

the Trichoptera.

Under the term “scent glands” it is necessary also to include

those hypothetical organs the emanation from which, in the females

of certain moths, is supposed, in the well-known phenomenon of

“assembling,” to attract males from long distances. These alluring

glands have not, I believe, been actually located, nor has the

nature of such emanation been ascertained. Of course odoriferous

glands undoubtedly do exist in many female Lepidoptera, and

recently Ernst Urbahn (21) has made a detailed study of these

glands, which are restricted to the abdomen and occur as

intersegmental sacs, folds, and so on.

Then, again, in insects like the Musk Beetle (Aromia mos-

chata), scent glands of still another type are found. These are

pluricellular and open to the exterior by an aperture. In Aroma

they are present in both sexes, though the aroma is stronger in

the female, while the male is the more active organism.

It is often difficult to distinguish scent glands of this type from

stink or repugnatorial glands used in self-defence. The resem-

blance is increased by the fact that some organs which have been

described as stink glands, e.g., in the Cockroach, Phyllodromia

germanica (16), are nevertheless limited to the male. On the

other hand, sac-like glands at the end of the abdomen of a

Cricket, Ceuthophilus maculatus [(10) p. 393], occurring only in

the male, are regarded as scent glands *.

There is, indeed, a widespread confusion in the literature of

the subject, and it is difficult or impossible, in the present state

of our knowledge of the natural history and mating habits of

these insects, to say whether glands, occurring as they may in

one sex only or in both, are of sexual import or are used in self-

defence. A priori, stink glands if used in self-defence, one would

expect to be either common to both sexes or, if limited to one sex,

to occur in the female rather than the male 7.

The occurrence of typical unicellular scent glands at the bases

of hairs in Trichoptera as well as in Lepidoptera is interest-

ing, and in view of the close relationship of these two Orders,

not wholly unexpected. But in consequence of the common

occurrence of scent glands in other insects besides Lepidoptera

and Trichoptera, this cannot be taken in itself as evidence of

phylogenetic affinity any more than can the presence of scales,

which are also present in Trichoptera and Lepidoptera, but

which occur also in Thysanura and other insects. Further,

in Sericostoma, as will presently be shown, they occur on the

* The organs, called by their discoverer, Kraus, “ duft organe” in Aphlebia

bivittata, ave named by Berlese “ ghiandole repugnatorie.”

+ The stink gland, of course, must have been independently acquired very many

times, for it is a device for self-defence adopted by many animals in very different

phyla of the Animal Kingdom (e.g. Myriapods and Mammals). Scent or alluring

glands are also common ‘The mammalian anal, preputial, and inguinal glands

are doubtless of sexual importance on account of their odoriferous secretions.

Odoriferous glands occur also in crocodiles and snakes.

462 MR. B. F. CUMMINGS ON

maxillary palpi, a position in which, I believe, they are unrecorded

in the Lepidoptera.

Although Kellogg (18) bas described and figured special

plumules and scale-lke hairs on the wings of 'Trichoptera,

é.g., Mystacides punctata, which function probably as andro-

conia, scent organs have not, I think, been hitherto examined

anatomically in this Order, for even in the paper in which

Miller gives us an account of his discovery of the true function

of the extraordinary palpi of Sericostoma, no account is included

of the structure either of the palpi or of the glands.

In Sericostoma they differ in position from the scent glands of

other insects. Scent glands are found on the abdomen or on

the thoracic appendages. According to Berlese [(12) p. 525]

odoriferous scales have been found on the palpi of some Satur-

nine butterflies, but he does not say whether they are the labial

palpi or the maxillary palpi. Probably they are the former.

Well-developed and characteristic scent organs occur in the

labial palpi of an Indian butterfly, Bertula chalybialis; this,

and the case of Sericostoma personatum, form the only well

authenticated instances known to me of scent organs in the

head, in the one case on the labial palpi and in the other on

the maxillary ‘palpi.

When not in use, the hair-tufts on the legs of the Lepidoptera

with scent organs are often concealed in cavities, just as the hair-

tuft of Sericostoma is concealed within the cavity of the imner

surface of the maxillary palpi.

I anticipate that among the many strange modifications of

the maxillary palpi of the males of many genera in the Family

Sericostomatide, sometimes densely clothed with large black

striated scales (Gara and Lepidostoma), scent organs will be

commonly found and their occurrence in the Trichoptera be

very generally recognised.

It is evident that scent glands in insects are of fairly common

occurrence and have been independently acquired over and over

again. According to Berlese, hairs like androconia have been

described from the wings of Diptera.

i. Tar MAxituAry Paupt or SERICOSTOMA PERSONATUM.

Each palpus in outline is something like a half-moon. The

outer surface is convex and strongly chitinous, dark brown in

colour, with scattered short black hairs. The inner surfaces of

the two palpi, which are carried closely apposed to each other,

are concave, but their concavities are filled up to a level surface

with an extremely thick felting of very long golden-yellow silky

hairs. Until the palpi are separated with a needle the one from

the other, the yellow felting lining the inner sides cannot be

seen, or seen only with difficulty by looking at them edgeways

from in front.

Both the palpi are held vertically with their broad surfaces facing

SCENT ORGANS IN TRICHOPTERA. 463

laterally (text-fig. 2). Together they entirely cover up the front

of the head or clypeal region. Their tips are received each into a

slot or excavation of the inner side of the enormously enlarged

basal joints of the antenne (text-figs. 4 & 5); emerging from

underneath the palpi may be observed the tip of the white fleshy

haustellum, the white tip of the labium and the lobes of the first

maxille.

Text-figure 2.

Sericostoma personatum, 6.

Head, side view (diagrammatic). 22.

E. Eye. Aj. First segment of the antenna. F. Genal flap. C. Cardo.

ST. Stipes. H. Haustellum. JP. Palpus of the first maxilla. LP.

Palpus of the second maxilla. SW. Subocular wart.

In order to understand the modifications which have been

drawn in the train of the enormous enlargement and porrec-

tion of the maxillary palpi, a few words are necessary upon

the maxillary palpi and the structure of the head in other

Trichoptera. In this Order, with the exception of Plectrotarsus,

Chimarrha, and Ptilocolepus*, the lower side of the skull is

incomplete, the gular region from the occipital foramen to the

labiam being soft and membranous. On either side the two

* The species examined were Plectrotarsus gravenhorstii Kolenati, Chimarrha

argentinea Ulmer, and Ptiloculepus granulatus Pictet. P. gravenhorstii is the

only species known belonging to its genus, and in the other genera it has been

assumed that the observations following, recorded for the particular species, apply

as well to the rest of the species of the geuus.

AGA MR. B. F. CUMMINGS ON

gene are drawn out ventrally into flaps, usually triangular in

shape, which hang down as strong perpendicular walls beside

the soft gular region (text-fig. 3, F).

Text-figure 3.

Phryganea, 3.

Side view of the head (diagrammatic). Much enlarged.

E. Eye. O. Ocellus. A. Antenna. #'. Genal flap. C. Cardo. STZ. Stipes.

LAB. Labrum. H. Haustellum. JP. Palpus of the first maxilla.

The cardo and stipes are not free like the four joimts of the palpus, but are

attached by their inner surfaces to the sides of the haustellum. Maxillary lobes

not shown.

For the sake of convenience it is easy to divide the mouth-parts

of adult Trichoptera roughly into those with a very elongate

haustellum*, and with no mandibles or with the mandibles much

reduced, and those with a shorter haustellum and a pair of

powerful well-developed mandibles. In the former case, for

example in Phryganea and Limnophilus, the cardo of the first

maxilla is articulated to the head in a small angle or niche

formed at the point where the gena curves down as the trian-

gular flap (text-fig. 3, F). In these genera, both cardo and stipes

are elongate and lie along each side of the stalk or peduncle of

the haustellum and carry off the maxillary palpi at their distal

end some distance away from the head. In the latter case, for

* Haustellum was the name given by Lucas (19) to what he regarded as an

enormously developed fleshy labium projecting from the head as a sort of proboscis

by means of which caddis-flies obtain their nourishment. In reality the haustellum

is a modification of the region of the hypopharynx.

SCENT ORGANS IN TRICHOPTERA. 465

example in hyacophila, the cardo and stipes are much shorter,

so that the maxillary palpi are carried close to the head instead

of at a distance from it. In such cases the angle or niche

in which the cardo, is articulated is much larger, and in this

angle the cardo is attached to the head by the whole of its

inner surface, whereas in Phryganea and others this angle or

Text-figure 4.

LAB

Sericostomea personatum.

Head from in front with the palpi and lobes of the first maxillee dissected off

(diagrammatic). >< 36.

E. Eye. Aj. First segment of the antenna, excavated to receive tips of the

palpi. P. Upturned process. LAB. Labrum. SC. Soft chitin. C.SC.

Cly peal sclerite.

niche receives only the proximal end of the cardo, which is

provided with a stout condyle for the attachment of a powerful

muscle arising from the tentorium. ‘The stipes is carried at

right angles to the cardo, so that the general direction of the

palpi is dorsal. The female of Sericostoma personatum agrees

466 MR. B. F. CUMMINGS ON

with this general description; but in the male the angle or

niche has developed into a deep somewhat rectangular exca-

vation of the gena, which is carried right back as far as the -

lower part of the eye and the subocular wart. Above it. is

bounded by the lower margin of the clypeus. The whole of this

area (text-fig. 2) is reserved for the cardo and stipes which are

very closely attached along their inner surfaces. The stipes has

been forced back close to the head and back upon the cardo,

both cardo and stipes being almost vertical in direction (text-

fig. 2, ST & C). The stipes is a long sclerite, narrow at the base,

broadening out gradually towards the distal end, where the

palpus is inserted. It apparently gives only the slightest support

to the maxillary lobe, as the stipes always comes away from

the latter in dissection with great ease, and the chitinous bar,

which in other Trichoptera the stipes sends in as a supporting

connection with it, cannot be discovered. The cardo is much

reduced, being visible at the side as a narrow chitinous splint.

Thus the two maxillary palpi completely cover the clypeus and

front of the head, and their chitinous, convex, outer surfaces

form a kind of mask or false front, beneath which the chitin of

the clypeus has become thin and in places delicate and trans-

parent (text-fig. 4, SC).

The labrum is fairly large, with a knob or enlarged basal] piece.

The mandibles, though present, are very much atrophied, but may

be detected in a careful preparation one on either side of the

base of the labrum, as a pair of thin pointed splint-like pieces of

chitin. In most other Trichoptera well-developed mandibles are

commonly found,

The basal joint of the antenna requires further description.

Text-fig. 5 is a sagittal section of the head and gives a view of

the relations between the maxillary palpus and the antenna.

Text-fig. 4 gives a view from in front after both the palpi and

lobes have been removed.

From an inspection of these figures it should be clear that

these enlarged basal joints meet each other behind, but are

excavated in front on their inner surfaces to receive the tips of

the ascending maxillary palpi. The inner front part of each

basal antennal joint is scalloped out into a slot into which the

palpus tip is neatly fitted and locked by means of the little

upturned process (text-figs. 4&5). The tip of the palpus also

is modified to serve this end, as is mentioned and illustrated

further on (see p. 468, Section iv.).

In addition to the mask or false front to the head formed by

the palpi there is also a false top to the head formed by these

two greatly enlarged basal segments of the antenne, which

extend backwards nearly as far as the occiput, covering nearly

the whole of the top of the head. The whip-like remainder of

the antenna springs, as if from the head itself, from the extreme

anterior corner of this large basal segment. Hach basal segment

on the inner excavated side is white in colour and composed of

SCENT ORGANS IN TRICHOPTERA. 467

two “ windows” of thin delicate chitin divided by a median bar

of thicker chitin running dorso-ventrally. The segment itself is

almost immobile, but its enlarged size allows for the presence

of muscles by which the long whip-like remainder of the antenna

can be moved.

Text-figure 9.

Sericostoma personatum, 3b.

Sagittal section of the head, with the palpus of the first maxilla and the first

joint of the antenna seen from the inside (diagrammatic). 27.

The shape of the tentorium appears to indicate limited anten-

nary movements, as the two columnar endosternites or supporting

pillars, which run from the cross-bar at the occipital foramen

across the inside of the head to the clypeus, are simple, without

wings or lateral expansions from which, when present in

Phryganea and other genera, arise numerous powerful muscles

to the antenne. In the female of S. personatwm, where the

antenne have much smaller basal segments, the tentorium is,

however, similar to the tentorium of the male in the absence

of its wings.

The head of the female is more or less normal. The maxillary

palpi are fairly long 5-jointed appendages; the chitin of the

clypeus is dark brown in colour and of equal thickness to that

of the rest of the head. The mandibles are aiso much longer,

Proc. Zoou. Soc.—1914, No. XX XI. 3l

468 MR. B. F. CUMMINGS ON

being visible under a strong power without the need of special

microscopic preparation. The first segment of the antenna,

though much larger than the succeeding segments, is separated

from the first segment of the other antenna by a wide space,

and bears no sort of resemblance to the eccentric form of these

segments in the male. The cardo is also much larger than the

cardo in the male, and instead of being practically vertical beside

the vertical stipes it is at right angles to the latter. The stipes

is shorter and broader than the same piece in the male, and the

palpi are so hinged upon it as to point outwards and away from

the head.

iv. THe Scent ORGANS OF SERICOSTOMA PERSONATUM.

More detailed examination of a detached palpus of a male

reveals the following additional facts.

The deep brown, comparatively thick chitin of the convex

outer side ceases abruptly in a transverse line towards the tip

so as to leave the soft white tip of the palpus (composed of

thinner chitin of a lighter colour) to fit more easily into the

base of the antenna. On the inner surface at the base is

a small lobe with two or three short black bristles at its base

(text-fig. 6). These bristles in cross-section are seen to be

fluted.

In side view the outer margin is very convex, the inner

margin, which is applied to the head, more or less straight.

The outer surface is very convex, and the inner surface applied

to the inner surface of its fellow is concave. But the concavity

is full of very long silky hairs, which curve up and around in a

semicircle to protrude near the tip into the hollow in the base

of the antenne. In the figures only a few of these hairs are

sketched in. This huge hair-tuft is bounded on the outside by

a rather pronounced lip, bare except for a row, on the margin,

of small black bristles, in cross-section seen to be fluted (text-

fig. 7, FH), and on the inside by a flap (text-fig. 6, FL), which

projects and keeps the hairs tucked in so that they are prevented

from straying in an untidy mass on both sides. Further refer-

ence to this flap is given below. It runs down towards the base

of the palpus, where it is continuous with the “lip” of the convex

outer margin. Higher up it carries a fringe of hairs much

shorter and stouter than the silky hairs of the scent-gland tuft.

The ends of these hairs are somewhat swollen. The flap ends

abruptly higher wp so as to leave a channel between it on one

side and the lip of the palpus opposite on the other side.

Between these two promontories the long hairs sweep out beyond

the edge of the palpus, and are sometimes visible as “ fluff”

projecting from between the two basal antennal segments behind.

Under the flap is the opening into a large sac which occupies

the whole of the interior of the palpus from the top to its extreme

bottom near the stipes.

SCENT ORGANS IN TRICHOPTERA. 469

Beneath this sac is another sac which opens by a longitudinal

slit along the inner margin of the palpus beneath the opening of

the first sac. At first I thought this sac was only an artefact,

imagining that the membrane lining the hard chitinous convex

surface of the palpi had come away. But sections showed the

existence of a distinct cellular lining to the inside of this chitin,

and there can be no doubt that the sac in fact exists.

Text-figure 6.

Sericostoma personatum, 6.

Palpus of the first maxilla, seen from the inside with most of the silky hairs

removed (diagrammatic). X 39.

ST. Stipes. L. Lip. LB. Lobe. FL. Flap. 8S). Entrance to sac 1.

So. Entrance to sac 2. S. Scent hairs. ZT. Tip of palpus.

A study of transverse sections of the palpus affords us the

following histological information.

Text-fig. 7 is through the line AB in text-fig. 6. It shows

the two sacs and the septum between them, and the layer of

large, deeply stained, glandular cells at the base of the silky

hairs.

The hypodermis beneath the outer, convex cuticle of the palpus

is composed of a double layer of flat squamous cells. The

septum between the two sacs is formed by the innermost cell-

31*

470 MR. B. FE. CUMMINGS ON

Text-figure 7.

Sericostoma personatum, 3.

Transverse section along the line AB in text-fig. 6. Length of section 1:1 mm.

8). Sacl. Sp). Sac 2. FL. The Flap. ZL. Lip. FH. Fluted hairs. GL.E.

Glandular epithelium. C. Cuticle. H¥P. Hypodermis. SC. Scent

hairs. CZ. Chitinous intima. J Muscle-fibres.

SCENT ORGANS IN TRICHOPTERA. 471

layer which runs up to encircle a number of muscle-fibres. This

septum is stiffened by a comparatively thick chitinous intima

(text-fig. 7, CL), which runs in from the entrance to cover about

half of the outside wall of the inner sac.

In the sections, the “lip” (LL) has been accentuated owing to

the sinking in of the tissues beneath it to form a hollow where

the fluted edges of the sections of the dark hairs are seen.

Under this lip the two squamous layers of cells are widely

separated the one from the other and the space between them

traversed by strands. ‘The cells are also larger.

On working round to the inner surface of the palpus one finds

the coneavity full of an immense thickness of hairs seen to be

circular in cross-section and containing a central canal of small

bore. The cuticle supporting these hairs is produced into elongate

papille containing the alveoli in which the scent hairs are fixed.

Beneath the cuticle the hypodermis consists of a glandular

epithelium of elongate cells, specialised formative cells called by

Graber trichogens, in which the scent is secreted [(10) p. 188].

On account of the fact of the immense number of these cells and

of the hairs which they support, it has not been easy, from an in-

spection of sections of unfixed material, to say definitely whether

there is a single trichogen cell to each hair or whether there are

several (text-fig. 8). Usually in the scent organs of the Lepi-

doptera there is one cell—one hair; but Bertkau (20), in the

ease of the Noctuine genera Hadena and Dichronia, points out

that there is not one giant cell to each of the hairs of the scent

tuft (which are of enormous length), but several smaller cells

belong to each hair.

That these hairs may act as scent organs it is necessary for

them to remain in connection with the living hypodermic tissue.

Text-fig. 8 shows how a pore-canal or channel runs up to the

base of each alveolus through the chitinous papilla, thus putting

into communication the cutaneous appendage with the hypodermic

trichogen cell.

No opening pores were found either at the base of or at the

tip of the hairs, and probably the scent secretion runs up the

canal within the hairs by capillary attraction and becomes

diffused by osmosis into the outer air. In the androconia and

scent hairs of the Lepidoptera it is now generally held | Berlese

(12) p. 533] that the secretion reaches the air by osmosis, as

apertures in the integumentary appendages are no longer thought

to exist.

The cuticle of the inner surface is plicate and the hypodermic

cells are much smaller. Strands of tissue run across between

the hypodermis of the inner and outer layers, being very clearly

seen where the two layers of cells diverge from each other in

the “lip.”

It will be remembered that Miiller (supra, p. 460, Section i) de- —

scribes the male during mating as separating the maxillary palpi

and spreading out the hair around the head * wie ein Heiligen-

A472, MR. B. F. CUMMINGS ON

schein.” As a result of my study of these palpi, no very clear

or concise expression of the manner in which this is done can be

offered. No longitudinal muscle-fibres as they occur, for example,

at the base of scent hair-tufts in some Butterflies, were discovered

attached to the bases of the hairs. A circular muscle around

the hair within the alveolus, but at some point above the base,

would serve to erect the hair. No such muscle has been detected.

Text-figure 8.

Sericostoma personatum.

A. Portion of the glandular epithelium, enlarged. Actual width varies from

053 to °03 of a millimetre.

B. Some trichogen cells very greatly enlarged.

Reichert objective ;'y homog. imm., with eyepiece No. 4 was used for the

examination of the cells. ‘The drawings are free-hand.

A, Alveolus. H. Hair. PC. Pore-canal. WV. Nucleus.

It remains to consider the action of the stout muscle-fibres in

the edge of the septum previously described. Their course is

longitudinal from the base of the palpus to near the end. The

result of their contraction would be to draw down the tip of the

palpus and so make the convex outer margin still more convex,

SCENT ORGANS IN TRICHOPTERA. A473

in which event the long silky hairs, which are curved and so fit

the normal amount of the convexity of the outer margin of the

palpus, would slip out of their concavity over the “lip” of

the outer convex edge and project as a “frill.” This action

is often suggested while manceuvring the detached palpus in the

dissecting dish. Again, if we suppose that the palpi by the aid of

their muscular attachments to the stipes can be easily divaricated

the one from the other and their inner surfaces exposed, it seems

probable that the natural resilience of the hairs in the live

animal would account for their erection, as they are inserted at

right angles to the surface on which they stand, but in repose

are flattened down upon it because the two palpi are then held

closely apposed to each other. Perhaps also in the live animal

the natural elasticity of the inner surface of the palpus may result,

when exposed as a free surface, in its becoming swollen and convex,

rather than as in spirit-specimens, concave.

I do not think that the resemblance between the scent glands

of Lepidoptera and those of Trichoptera can very profitably be

carried into histological details. The papers on scent glands in

Lepidoptera which I have consulted all show in one particular

or another considerable differences from those of Sericostoma

personatum, as, for example, in the presence of muscle-bands, in

the arrangement of the trichogen cells, or the shape of the pore-

canals and the position of the hair-tuft.

I have to thank Mr, Martin E. Mosely for kindly giving me

the material for this study, Prof. Maxwell Lefroy for allowing

me to work in the Laboratory at the Imperial College of Science

and Technology, and Mr. HE. Hargreaves for assistance 1n section

cutting.

v. LITERATURE.

(1) Kirpy and Srence.—Introduction to Entomology. London,

1826.

(2) Picter, F. J.—Recherches pour servir a Histoire et

VP Anatomie des Phryganides. Geneva, 1834.

(3) SrepHens.—Ilustrations, of British Entomology, vol. vi.

p. 148. London, 1836-1837.

(4) Burmerster.—Handbuch der Entomologie, vol, ii. (2), p. 928.

Berlin, 1839.

(5) Rampur.—Histoire Naturelle des Insectes: Névropteres,

p. 494. Paris, 1842.

(6) Konenatr.—Genera et Species Trichopteroram, Pt.I. Act.

Reg. Bohem. Soc. Se. Prague, vol. vi. 1851, p. 89.

(7) McLacutan, R.—A Monographie Revision and Synopsis

of the Trichoptera of the European Fauna. London,

1874-80, pp. 223-224.

(8) Umer, G.—Wytsman’s Genera Inseectorum: Trichoptera.

Brussels, 1907.

(9) Mutter, W.—Duftorgane bei Phryganiden, Archiv fur

Naturgeschichte, 1887, p. 95.

474 ON SCENT ORGANS IN TRICHOPTERA.

(10) Packarp, A. S.—Textbook of Entomology. London, 1898,

p- o9 1.

(11) Inure, K. G.—Duftorgane der Mannlichen Schmetterlinge.

Zoologica, Bd. xv. Heft 38, 1902.

(12) Bertese, A.—Gli Insetti, vol. i. p. 533 Milan, 1906.

(13) Branpes, G.—Ueber Duftapparate bei Kiifern. Zeit. f.

Naturwiss. Bd. Ixxii. (ser. v. Band x.), 1899, p. 209.

(14) Haas, E.—Zur Anatomie der Blattiden. Zool. Anz.,

Bd. xii. p. 169.

(15) Kraus, H.—Die Duftdriisen der Aphlebia bivitiata. Zocl.

Anz., Bd. xiii. p. 054.

(16) Mincuin, EH. A.—Further Observations on the Dorsal Gland

in the Abdomen of Periplaneta and its Allies. Zool. Anz.,

Jan. 27, 1890.

(17) Garman, H.—On a Singular Gland possessed by the Male

Hadenecus subterraneus. Psyche, 1891, p. 105.

(18) Kenioce, V. L.—The Affinities of the Lepidopterous Wing.

American Naturalist, 1895.

(19) Lucas, R.—Beitriige zur Kenntniss der Mundwerkzeuge

der Trichoptera. Archiv fur Naturgeschichte, Bd. lix. 1.

TSI), av, 218155

(20) Berrkau, P.—Abstract in Journ. Roy. Micros. Soe. ser. 2,

vol. vill., 1888.

(21) Ursaun, Ernsr.—Jena Zeitschr. Naturw. Bd. 50 (1913).

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 475

EXHIBITIONS AND NOTICES.

March 17, 1914.

Prof, E. A. Mrincutn, M.A., F.R.S., F.Z.8., Vice-President,

in Ae Ghai:

The Secrerary read the following report on the additions to

the Society’s Menagerie during the month of February 1914 :—

The number of registered additions to the Society’s Menagerie

during the month of February was 157. Of these 95 were

acquired by presentation, 14 by purchase, 20 were received on

deposit, 5 in exchange, and 23 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 188.

Amongst the additions special attention may be directed to :-—

1 Pigmy Antelope (Neotragus pygmeus), from Accra, Gold

Coast, new to the Collection, presented by E. B. Reece on

February 4th.

2 Unial Sheep (Ovis vignez), from Jhelum, presented by Capt.

T. H. Scott on February 5th.

1 Wild Boar (Sus scrofa), from Antioch, presented by the

Officers of H.M.S. ‘ Duke of Edinburgh’ on February 25th.

1 Colpeo Dog (Canis culpeus) and 1 Salt-Desert Cavy (Doli-

chotis salinicola), from Cordova, presented by Wilfred A. Smithers,

C.M.Z.S., on February 4th.

1 Slow Loris (WVycticebus tardigradus) and 2 Finlayson’s

Squirrels (Sciwrus finlaysoni), from Koh Si Chang, Siam, pre-

sented by Commander Robert E. Buske-Peel on February 27th.

1 Graceful Mocking-Bird (AZimus gilvus), from Central America,

new to the Collection, presented by Hubert D. Astley, F.Z.S.,

on February 3rd.

2 Tooth-billed Tanagers (Pyranga bidentata), from Central

America, new to the Collection, received in exchange on

February 11th.

2 Cinnamomeous Kestrels (Cerchneis cinnamomina), from Cor-

dova, Argentina, new to the Collection, presented by Wilfred A.

Smithers, C.M.Z.8., on February 4th.

Mr. G. C. Rosson, B.A., read a report on Mollusca from Dutch

New Guinea collected by the British Ornithologists’ Union and

Wollaston Expeditions. Im general, the collection appears to

endorse Hedley’s views as to the Oriental affinities of the Papuan

molluscan fauna. Though numerically small in species and in-

dividuals, the collection has yielded two new genera and three

new species, the anatomy of all of which is described. The two

new genera, which were obtained from considerable altitudes,

viz. 10,500 ft. and 14,200 ft. respectively, are of considerable

476 MR. R. H. BURNE ON

interest, though their precise affinities are as yet uncertain. In

any case they cannot be regarded as typical members of the

Zonitide, though an aggregate of anatomical characters exhibits

the characters of that family. An account of the anatomy of

Papuina litwws (Lesson) is given, and discrepaney between the

anatomical and conchological relationships of a new species of

Papuina is discussed.

This paper has been published in the TRANSACTIONS.

Mr. K. G. Buatr, B.Se., read his report on the Heteromerous

Coleoptera collected by the British Ornithologists’ Union and the

Wollaston Expeditions in Dutch New Guinea.

The most interesting feature of the collection, from the point

of view of distribution, is the occurrence of Cissites maxillosa Fab.

iu thisregion. This beetle has been hitherto regarded as peculiar

to the Oriental Region, its range extending from Assam to Java,

Borneo, and the Philippine Islands; it has alse been found in

Ceylon.

The three species of Amarygmus belong to a section of the

genus that makes New Guinea its headquarters ; a few species of

this section are found in the extreme north of Australia, but the

majority of the Australian species belong to other groups.

Of the fourteen species noted, seven are described as new.

This paper will be published in the TRansacrions in due

course,

Palatal growth in mouth of Camel.

Mr. R. H. Burne, M.A., F.Z.S., exhibited a specimen of the

palate of a female Bactrian Camel (Camelus bactrianus) and a

lantern-slide of a dissection of the throat of a male Common

Camel (Camelus dromedarius)* (text-fig. 1), and drew attention to

a pendulous outgrowth from the roof of the mouth.

This outgrowth or palatal appendage is situated several inches

in front of the free posterior border of the palate and is

continuous on either side with the anterior pillars of the fauces.

It is rudimentary in the female, but of large size in the male

where it forms a great flaccid mass of tissue 11 inches (28 cm.)

long, hanging down the throat for some distance beyond the

larynx.

Short accounts of this appendage are to be found in many of

the older anatomical text-bookst, and it has recently been briefly

described by Prof. Lesbre in his monograph on the anatomy of

the Camel =.

It is a secondary sexual organ which during rut, when the

animal is excited, is protruded from the mouth “to the accom-

* R. Coll. Surg. Museum No. 1497, Physiol Series, Hunterian specimen.

+ Buffon, Cuvier, de Blainville, Owen, Milne Edwards.

t Lesbre, Arch. Mus. Hist. Nat. Lyons, ¢. 8, 1903.

PALATAL GROWTH IN MOUTH OF CAMEL. AGT

paniment of an abominable gurgling noise” as a “ thin mem-

branous bladder... . until it is as large as the animal’s head” *.

Text-figure 1.

Pharynx of Camel, opened from the right side.

B.PH., buccal pharynx; £., epiglottis; N.PH., naso-pharynx; CHs., cesophagus;

P., soft palate (cut edge); P.’, posterior free margin of soft palate; P.A., pala-

tine appendage; 7'., base of tongue.

From published descriptions of the process and from remarks

made by Prof. Minchin and others at the meeting, it appears as

though the appendage were protruded by inflation with air, but

a careful examination of the specimen figured above revealed no

hole or passage leading from the naso-pharynx or elsewhere

through which air could be forced into the interior of the organ.

The only indication of anything of the kind was a shallow cleft

or recess in the mid-line of the floor of the naso-pharynx above the

root of the appendage. But the size of this pit (3°5 em. long and

2-5 em. deep) even allowing for considerable powers of distension,

was not great enough to suggest that it could by any possibility

be sufficiently blown out to fill the whole interior of the ap-

pendage.

* Spencer & Gillen, Across Australia, 1912, vol. i. p. 38, fig. 17.

478 MR. R. I. POCOCK ON THE

Judging by feel and by the parts of the interior of the

appendage exposed by cuts, it would appear that the organ is

solid and composed of very delicate, loose, areolar tissue. Injection

of water or air through a cut upon the surface caused immediate

and extensive inflation of this tissue, with the exertion of very

little force. Protrusion may, therefore, he due to the infusion of

fluid (lymph ?) into the substance of the organ, though possibly

inflation of the above-mentioned pit in the naso-pharynx may be

accessory to the protrusion by exerting pressure upon the fluids

contained in the more distal parts of the appendage.

On the Feet of Domestic Dogs.

(Text-figures 1-3.)

Mr. R. I. Pocock, F.R.S., F.L.S., F.Z.S., Curator of Mammals,

exhibited a series of lantern-slides illustrating some points in the

structure of the feet of domestic breeds of dogs (Canis familiaris),

and remarked :—

“‘ According to F. Cuvier and Geoffroy St. Hilaire (Hist. Nat.

Mamm. il. no. 166, 1820), the interdigital integument of New-

foundland dogs extends almost to the claws and widens to such

an extent as to make the feet palmated. In this respect,

according to these authors, the feet of this breed differ from

those of the majority of breeds in which the web in question is

of small extent and reaches only as far as the origin (proximal

end) of the second phalanx ; but, they add, ‘the peculiarity found

in the Newfoundland dog is not restricted to that breed, but is

observable in several of our [French] breeds, and especially in

those not belonging to the category of running dogs.’ St. Hilaire

restated the fact about the feet of the Newfoundland dog in

1862 (Hist. Nat. Gén. iti. p. 450).

In ‘The Variation of Animals and Plants under Domesti-

cation,’ 1. p. 49, ed. 1905, Darwin, after referring to St. Hilaire’s

later work, wrote: ‘In two Newfoundland Dogs which I

examined, when the toes were stretched apart and viewed on the

underside, the skin extended in a nearly straight line between

the outer margins of the ball of the toes, whereas in two terriers

of distinct sub-breeds, the skin viewed in the same manner was

deeply scooped out.’

This description is not very intelligible. It neither confirms

nor contradicts Cuvier’s statement, because the point on the

margin of the balls of the toes to which the skin was attached is

not given. Moreover, no web can extend between the outer

margins of the toes. It must stretch across the middle line

between the third and fourth toes, and from the outer margins

of the latter to the inner margins of the second and third,

respectively. Setting these difficulties aside, however, it will be

seen that Darwin did not allude to any difference between the

FEET OF DOMESTIC DOGS. 479

feet of Newfoundlands and terriers with respect to the forward

extension of the skin between the digits.

He also referred to a record by Mr. Greenhow of the occurrence

in Canada of a peculiar dog with ‘ half-webbed feet’ which was

‘fond of the water’ (Loudon’s Mag. of Nat. Hist. vi. p. 511,

1833). and quoted Mr. C. O. Groom Napier to the effect that the

hind feet of otter-hounds are more webbed than those of harriers

and blood-hounds (‘ Land and Water,’ il. p. 270, 1866).

Darwin, it may be added, attributed this alleged peculiarity in

the feet of water- dogs partly, to unconscious selection by man

and partly to the inherited effects of use. Finally, as suggestive

of the correctness of the above-mentioned statements SES PSU ONS

the webbing of the feet of Newfoundland dogs, we find ‘ well-

webbed feet ’ enumerated amongst the show-points of this breed

(J. J. Cooper, ‘The Kennel Encyclopedia,’ iii. p. 942, 1908).

On the other hand, so long ago as 1861 or thereabouts,

J. G. Wood wrote (Illustrated Nat. Hist. i. p. 271): ‘Some

people fancy that the Water Spaniel possesses webbed feet, and

that its aquatic prowess is due to this formation. Such, however,

is not the case. All dogs have the toes connected with each

other by a strong membrane, and when the foot is wide and the

membrane rather loosely hung, as is the case with the Water

Spaniel, a large surface is presented to the water. Now since it

is generally admitted that the Newfoundland belongs to the

same group of dogs as the Water Spaniel, it is difficult to find

any reason why the larger form should have better-webbed feet

than the smaller, since both are what are called ‘ water-dogs.’

Be it remembered, too, that Cuvier’s statement about the feet of

Newfoundlands does not refer to the ‘looseness’ of the inter-

digital web, but to its extension along the edges of the pads

nearly up to the claws; and, as quoted above, this author asserted

the existence of similar webs in other European breeds excluded

from the category of running dogs.

Being unable to procure the feet either of a Newfoundland dog

or Otter-hound, I wrote to Mr. J. Sidney Turner, M.R.C.S.,

F.L.S., about the former breed and I venture to quote his reply.

‘There is no doubt that Cuvier’s statement that the feet of

Newfoundland dogs are more webbed than those of other dogs is

a pure myth. I have heard the same statement made about

Otter-hounds, but that is of course also wrong. The fact is, that

both these breeds have rather larger or longer feet than usual

compared with Mastiffs and Foxhounds. I mean that the feet

are not so compact and drawn up, and therefore the webbing is

rather more apparent, but it extends no farther along the

phalanges of the toes.’ This is practically what J. G. Wood said

about the Water Spaniel. And to clinch the matter so far as

the Newfoundland is concerned, Mr. Vero Shaw, the only modern

author, as Mr. Turner informed me, to mention the matter,

wrote: ‘The feet must be broad and flat. The vulgar opinion

that the dog is web-footed...has no other foundation in fact

480 MR. R. I. POCOCK ON THE

than that the toes of all dogs are connected by a skinny mem-

brane, but it does not extend to the point of the toes as in

web-footed birds’ (Cassell’s ‘ Book of the Dog,’ p. 69, 1881).

Since, however, the authoritative testimony of Cuvier and

Geoffroy St. Hilaire can hardly be dismissed in the summary and

concise manner adopted by Mr. Shaw, and since zoologists are

sure to consult Darwin for information on a point of this kind,

and to accept as true the uncontradicted statements of others

contained in his volume, I think it may be useful to publish in

our ‘ Proceedings’ figures and descriptions of the feet of some

of our breeds of dogs, to show the actual extent of the inter-

digital web. To ascertain this the hairs, long or short, clothing

this web and growing between the pads in all domesticated dogs,

have been cut away, and the figures here published are taken

from the paws after clipping. The drawings are partially dia-

grammatic in the sense that the digital pads are represented as

lying in the same plane as the rest of the lower surface of the

foot, whereas, naturally, they incline upwards in a plane of

varying steepness, according to the breed. One or two additional

points in which the feet have been modified by selective breeding,

or in correlation with other features, have also been referred to.

I was induced in the first instance to look into the question of

the alleged palmation of the feet of Newfoundland dogs and

of some European breeds, by finding that the feet of various wild

species of the family belonging to several valid and nominal

genera of so-called wolves, jackals, and foxes, only differ to a

small degree, inter se, in the extension of the web along the edges

of the digital pads*. The feet of some species, it is true, are

more webbed than others in the sense that the toes are more

widely separable, but this is attributable to the greater width of

the integument connecting adjacent toes and permitting their

wider separation.

For the material examined for the purpose of this notice, I am

indebted partly to Mr. A. J. Sewell, M.R.C.V.S., partly to

Mr. R. E. Holding, but mainly to Mr. B. Gorton, M.R.C.V.S.,

the Society’s Veterinary Surgeon. The series comprising, I

think, the extremes of modification met with in the dogs, with

the exception possibly of the Dachshund, which I have been

unable to procure, shows that the feet differ remarkably in

length from the wrist to the digital pads, in the length of the

digits, the width, length, and shape of the plantar pad, the width

across from the second to the fifth toes and the length of the hair

between the pads, and other minor features; but very little in

the extent to which the hair spreads over the pads, and scarcely

at all in the extension of the web along the margins of the

digital pads.

The web passes between the inner proximal angles of the third

and fourth digital pads, and from the inner proximal angle of the

* There is only one exception to this, which I shall refer to in a subsequent

publication.

FEET OF DOMESTIC DOGS. 481

second and fifth to the outer proximal angles of the third and

fourth. The edges of the web are thick and elastic, and the

elasticity keeps the toes in a compact mass when the foot is slack,

but permits their separation under the weight of the standing

Text-figure 1.

Wesssccrcnl ld 1 yy

AY LOANS

A. Left fore paw of Clumber Spaniel, from below.

c., carpal pad; p., plantar pad; d2, 13, d4, @5, digital pads of second

third, fourth, and fifth digits; 2w., web.

B. The same, from above.

gl., glandular pocket at base of digits.

C. Piece of integument of the interdigital web, showing tufted growth of

hair (diagrammatic, the hairs cut quite short).

animal and particularly under the pressure of running. The dog

eannot voluntarily expand its toes by overcoming this elasticity

when the foot is lifted and, as Mr. Sidney Turner pointed out to

me, pressure against the water in swimming 1s insufficient for the

482 MR. R. I. POCOCK ON THE

purpose. Hence the webs themselves can be of little, if any,

service for aquatic progression.

Text-figure 2.

VV)

SHEN Nos

ASS HTL (|

WA&

iN)

\y\

Ss <—

TEAS s

iit

Ze YOU

A. Left fore paw of Aberdeen Terrier, from below.

B. The same of Bulldog.

C. The same of Greyhound.

Both the under and upper sides of the web are highly glandular,

and on the upper side there is a little naked pocket in the angle

formed by the junction of the third and fourth and fourth and

fifth toes. This is sometimes absent, and always much shallower,

between the second and third toes. The hair clothing the web

grows in little tufts from crater-like depressions. The plantar

pad varies considerably in length and width, according to the

FEET OF DOMESTIC DOGS. 483

length and width of the foot. It is comparatively long, for

example, in the Greyhound, and broad in the Aberdeen Terrier.

The foot of the latter is exceedingly short, owing mainly to the

extreme abbreviation of the third and fourth toes, in accordance

with the modern fashion for shortening the feet and aligning the

metacarpus with the radius. This fashion has also changed the

slope of the digital pads, so that they incline nearly at right

angles to the long axis of the foot, and has thickened or deepened

the plantar pad. These points may be seen by comparing the

Text-figure 3.

A. Left fore paw of Aberdeen Terrier, from its inner side.

B. The same of Bulldog.

C. The same of Clumber Spaniel.

D. The same of Airedale Terrier.

rofile figures of the feet of the Airedale and Aberdeen Terriers

and the Bulldog with that of the Clumber Spaniel, which, in the

inclination of the digital pads and the depth of the plantar pad,

shows more resemblance to the paw of a wolf. The largest and

the smallest feet examined were respectively those of a St. Bernard

and of a Manchester terrier. The former do not differ materially

from those of the Clumber Spaniel, except that the median digits

- are shorter and approach those of the Aberdeen Terrier.

Proc. Zoou. Soc.—1914, No. XXXII. 32

A84 , ON THE FEET OF DOMESTIC DOGS.

A feature of the Greyhound’s foot is the length of the third

and fourth digits and the width of the intervening web which

permits their wide separation. In this sense, the fore foot* of

this breed is more fully webbed than that of any dog examined ;

and the wide divarication and length of these digits make the

web between them and the second and fifth digits appear to be

more deeply and widely scooped out or emarginate than in other

breeds. In the Bulldog the web between the third and fourth

digits appears to extend farther along the inner edges of the pads

than in other dogs; but this is due to the circumstance that

a small triangular area of the skin adjoining the inner angles of

these pads behind the margin of the web is naked and, simulating

the integument of the pad, makes the pads appear to meet across

the middle line.

The subjoined table giving some of the principal measurements

will show how the feet of the specimens examined vary in relative

and actual proportions, the difference between the figures in the

first two columns giving the length of the median toes beyond

the plantar pad.

If the Greyhound’s foot be compared with the Bulldog’s it will

be seen that the former, although actually very much longer, is

considerably narrower across the toes and a little narrower across

the plantar pad than the Bulldog’s. Again, the foot of the

Aberdeen Terrier, which is very short, is much wider than the

Pomeranian’sand almost as wide as the Greyhound’s, and whereas

the median toes of the Aberdeen project less than 20 mm. beyond

the plantar pad, those of the Greyhound project about 40 mm.

beyond it.”

Length from | Length from ; Wadithueneraee

carpal to ay carpal to tip Width of digitall pads

of digital! of plantar | plantar pad and 6:

pads. pad.

St. Bernard ...... .. 150 mm. 110 mm. 65 mm. 70 mm.

Greyhound ......... 125 mm. 84 mm. 35 mm, 48 mm.

Retriever ............ 118 mm. 80 mm. 38 mm. 50 mm.

Clumber Spaniel ... 95 mm. 68 mm. 38 mm. 50 mm.

Sheepdog ............ 100 mm. 70 mm. 38 min. 48 mm.

Pomeranian ......... 75 mm. 63 mm. 27 mm. 35 mm.

Airedale Terrier ... 90 mm. 63 mm. 35 mm. - 47 mm.

TBHCUNGNO) 35 oc ooe oadnne 83 mm. 60 mm. 38 mm. 55 mm.

Aberdeen Terrier ... 63 mm. 45 mm. 35 mm. 45 mm.

Manchester Terrier. 63 mm. | 37 mm. 18 mm, 28 mm.

* I was unable to get the hind foot of this dog.

ON LONGEVITY IN AN ORANG-UTAN. 485

April 7, 1914.

Prof. E. W. MacBripz, M.A., D.Sc., F.R.S., Vice-President,

in the Chair.

The Srcrerary, Dr. P. Cuaumers Mrircuett, F.R.S., exhibited

the photograph of a female Orang-utan (Simia sabyr ws), kindly sent

to him by Mr. W. H. D. Le Souéf, the Director of the Zoological

Gardens at Melbourne. According to the statement of Mr. Le

Souéf, this Ape had lived in the Gardens at Melbourne for twelve

years in an open-air enclosure attached to a shelter without any

artificial heat. Orangs were notoriously difficult to keep alive in

captivity, and even in Singapore they seldom lived for two years

after capture. Mr. Le Souéf’s example was certainly extremely

interesting. In the Society’s own Gardens, a fine male Orang,

obtained on Sept. 7, 1905, was still alive, and it was reported to

have been in captivity for eight years before it came to London,

so that it was still older than the Melbourne example and had

shown the cheek-plates for the last two years. Chimpanzees

were less delicate, but the average duration was not good. The

Chimpanzee known as ‘“ Mickie,” which had been purchased by

the Society on April 6, 1898, was still living, and certainly was

the Anthropoid Ape known to have lived longest in captivity.

The almost universal experience with Gorillas was that they lived

only a few weeks after reaching Hurope, and, in consequence of

this high mortality, the Secretary had for some years declined to

encourage importers by refusing to buy. In one Continental

Collection, however, a Gorilla had lived for several years.

The Secretary also exhibited two photographs recently sent to

him by Surg.-Major George Henderson, M.D., F.L.8., showing

a number of specimens of the large-tailed variety of Punjab

Domestic Sheep. The tails of some of these animals are so large

that they trail on the ground, and a small cart is provided to

carry the tail and enable the sheep to move about. One of these

carts, harnessed to a sheep, was shown in one of the photographs.

Dr. W. T. Cauman, F.Z.S., read a report on the River-Crabs

(Potamonide) collected by the British Ornithologists’ Union and

Wollaston Expeditions in Dutch New Guinea, containing the

descriptions of two new species.

This paper has been published in the TRANSACTIONS.

Mr. Ouprietp Tuomas, F.R.S., F.Z.S., read a report on the

Mammals collected by the British Ornithologists’ Union and

Wollaston Expeditions 1 in Dutch New Guinea.

The species obtained numbered 31, of which the apes of 12

had been brought home by the Expeditions,

486 REPORT ON AFTERNOON SCIENTIFIC MEETINGS.

The two Expeditions had obtained a very valuable series of

ground-animals, notably of the genus Uromys, but there seemed

to be, in the part of New Guinea explored, a remarkable absence

of arboreal species, these forming in other parts of New Guinea

a large proportion of the mammal fauna.

No species were now described as new, as the novelties had

been already published in previous papers.

This paper has been published in the Transactions.

April 21, 1914.

Dr, Henry Woopwarp, F.R.S., Vice-President,

in the Chair.

The Secrerary read the following report on the additions to

the Society’s Menagerie during the month of March 1914 :—

The number of registered additions to the Society’s Menagerie

during the month of March was 109. Of these 20 were acquired

by presentation, 29 by purchase, 32 were received on deposit, 16

in exchange, and 12 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 217.

Amongst the additions special attention may be directed to :—

2 Grévy’s Zebras (Hquus grevyi) 2 2, from Abyssinia, pur-

chased on March 2nd.

3 Indian Antelopes (Antilope cervicupra), presented by H.M.

Tue Kryé on March 2nd.

1 Eland (Yaurotragus oryx) 2, born in the Menagerie on

March 3rd.

1 Ibean Potto (Perodicticus ibeanus), new to the Collection,

deposited March 2nd. x

2 Blue-cheeked Amazon Parrots (Chrysotis versicolor), from

St. Lucia, presented by E. J. Cameron, C.M.G., on March 2nd.

The SEcrETaRy announced that the following recommendation

from the Committee of Publication was considered by the Council

at their meeting held on April 15th last, and adopted :—

AFTERNOON Sctrentiric MEETINGS.

The Publication Committee beg to report to Council the result

of a post-card ballot onthe question of afterneon meetings for

Scientific Business.

Notices and ballot-cards were sent to attendants at the

Scientific Meetings during the last two years, and to all the

members to whom the ‘ Abstracts’ are regularly sent, the total

number being 195.

ON THE EGG OF MANTELL’S KIWI. A487

150 replies have been received, and of these 109 are in favour

of the change, 39 are against it, and 2 are indifferent.

The Committee therefore recommend that, commencing with

the new Session in October 1914, the Meetings for Scientific

Business shall be held on Tuesdays at 5.30 p.m., the usual

refreshments being served from 5 to 5.30 p.M., and the Publi-

cation Committee meeting at 5 P.M.

Mr. D. Seru-Smuiru, F.Z.S., Curator of Birds, exhibited the

egg of Mantell’s Kiwi (Apteryx mantelli), laid in the Gardens

on April 12th. It weighed 113 oz., and measured 4:75 x

2°75 inches. The bird that laid the egg weighed only 654 oz.,

considerably less than the weight of a domestic hen, which laid

eges of only 27 oz.

Dr. P. L. Sclater had recorded the laying of eggs by this species

in the Gardens in 1853 and 1860, the weights of which were

greater than that of the present specimen, one being 144 oz. and

the other “‘ somewhat larger” (P. Z. S. 1853, p. 350, and 1860,

p- 194), and Sir Walter Buller had recorded the weight of an

egg taken in the wild state as 15 oz. 90 grs. (Trans. New Zealand

Inst. 1892, p. 85).

Mr. Sranitey Hirst, F.Z.S., reported on the Arachnida (other

than Spiders) and Myriopoda obtained by the British Ornitho-

logists’ Union and Wollaston Expeditions in Dutch New Guinea.

The collection is only a small one, but contains two new species

of Acari parasitic on mammals and three new species of

millipedes. A new species of parasitic mite collected by Prof.

F, Forster on various mammals in German New Guinea 1s also

described.

This paper has been published in the TRansacrions.

May 5, 1914.

Dr. Henry Wcopwarp, F.R.S., Vice-President,

in the Chair.

Surgeon G. Murray Luvicx, R.N., gave an interesting account

of the manners and customs of Adélie Penguins (Pygoscelis

adelic), which he had observed at the Cape Adare rookery while

with Scott’s Antarctic Expedition. He described their mating

habits, the making of their “nests,” hatching of the eggs, and

rearing of the young, and illustrated his remarks with a series

of lantern-slides prepared from his photographs.

488 ON THE ANTLERS OF RED DEER.

Antlers of Red Deer.

Str Epmunp G. Lover, Bt., F.Z.S., exhibited four pairs of

antlers of Red Deer (Cervus elaphus) and made the following

remarks :—

‘““The Red Deer antlers which I am exhibiting are exception-

ally fine specimens. ‘The one from the Carpathians was obtained

during the present generation. The horns are very massive and

heavy, with long points, and have 20 tines. With a small piece

of the frontal bone they weigh 244 lbs.

The most celebrated collection of Red Deer antlers is at

Schloss Moritzburg, belonging to the King of Saxony. The

antlers have been collected during the last 300 years and some of:

them are certainly older than 1611, Only twelve pairs of antlers

weigh more than 243 lbs.

Two very fine pairs of antlers which I exhibit were found in

a morass in Hungary. It is not easy to guess from the appear-

ance of the bone, etc., how long they had been there, but I do

not take them to be prehistoric or of any very great age.

One pair measures 51 in. in length, 112 in. round the burr,

and 50 in. in greatest outside spread.

The longest Red Deer horns known measure 533 in.

The other pair found in the morass has the great outside

spread of 58? in., and measure 102 in. round the burr; with

18 tines.

The fourth pair comes from Germany, and the animal was

probably killed many years ago. The horns measure 48 in. in

length, 92 in. round the burr, and 512 in. in greatest spread ;

with 20 tines.

We have always known that the Red Deer of Persia, Asia

Minor, and the Caucasus had longer faces than those of Germany,

France, and Britain; and I thought that the faces of the Deer

would be longer the farther Hast one found them, and that in

the Carpathians they would be intermediate between those of

Germany and Persia—from which country comes the type

of Cervus elaphus maral.

Quite lately I have had the opportunity of measuring several

skulls of deer which had been killed in the Caucasus and in the

Carpathians, and I found them practically identical in their

proportions and with equally long faces.

The measurements taken are the distance between the lower

edges of eye orbits, and from occipital crest to end of pre-

maxillaries.

Roughly, I find in the short-faced type the ratio is 1 to 3:3,

and in the long-faced type 1 to 3°6. Ido not attach any great

weight to these figures as the material has been so limited.

I have hunted up the skulls of Red Deer in the Natural

History Museum and measured those which I found, but there

are no specimens from France, Germany, Austria, Hungary, or

the Carpathians.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 489

It would appear that there may be some place in Hungary

where the short-faced type and long-faced type may be found

close together.

A book has lately been published by an Austrian gentleman on

the Management of Deer Forests, and in this book he mentions

that in some part of Hungary two types of Stags are to be met

with; he describes one as of a grey colour and the other of a red

colour. He says nothing about the shape of the skulls, but I

think it is quite probable that the grey-coloured stag may turn

out to be the long-faced Cervus elaphus maral.

It is much to be desired that more specimens of skulls (with or

without horns) should be obtained from different localities.”

- The following are the weights and measurements (in inches) of

the specimens exhibited :—

g : a

5 2 Se 5 7) nd) he

oS BI 2 3S cS) a 5 5 or.

oe : = pS one a) ia ° ~ m

Sere = O to) i af Qs » » or

ect = >S uu fo) 172) wm =|

oh he S| fi |] StS & By Es roe

ate Ze = 22 | 28 S “s) ze

Se 5 Oo oe) a= | Be A= S i= =

= 4 6 |S S) a = es ||

| ——————| — —

| | | |

Carpathians ...) 243 lbs.| 444 | 113 104 7 14¢ 283 442 20

2 2 2 4 4 2 4

Found in a

morass 1n 21 lbs. BIL 113 9s qi 28% | 394 | 50 14

Hungary.

Found ina

morass in | 23} 1bs.| 48 | 102 Ox if 38 43 582 | 18

Hungary.

_——.

Germany ...... | 182 lbs.| 48 93 82 63 833i | 414 | 612) 20

May 19, 1914.

R. H. Burne, Esq., M.A., Vice-President, in the Chair.

The Sucrerary submitted the following report on the additions

to the Society’s Menagerie during the month of April 1914.

The number of registered additions to the Society’s Menagerie

during the month of April was 244, Of these 172 were acquired

by presentation, 35 by purchase, 12 were received on deposit,

3 in exchange, and 22 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 153.

Amongst the additions special attention may be directed to :—

2 Elephant-Seals (JMJacrorhinus leoninus) § 2, from the

490 ON HORNLESS ANTELOPES FROM THE DINDER RIVER.

Antarctic Seas, presented by H.G. The Duke of Bedford, K.G.,

Pres. Z.S., on April 6th.

2 Indian Hlephants (Hlephas maximus) $ 3, from India, pre-

sented by ‘The Daily Mirror’ on April Ist.

2 Tigers (felis tigris) $ 2, from Burma, presented by Major

F. Bigg Wither on April 27th.

1 Binturong (Arctictis binturong), from Malacca, purchased on

April 6th.

A Collection of Mammals and Birds, including 2 Bonda’s

Squirrels (Sciurus saltwensis bond«), 1 Collared Peccary (Dicotyles

tajacu), 2 White-browed Hares (Sylvilagus superciliaris), new to

the Collection ; 1 Pileated Heron (Pilerodius pileatus), new to the

Collection, 1 Prince Albert’s Curassow (Crax alberti), and 1 Banded

Tinamou (Crypturus noctivagus), from Rio César, Colombia, pre-

sented by W. K. Pomeroy, F.Z.S., on April 29th.

A Collection of Small Birds from Chili, including Chilian

Starlings (Cureus aterrimus), Little Saffron Finches (Sycalis

minor), and others, presented by George H. F. Duncan, F.Z.5.,

on April 20th.

3 Sharp-nosed Terrapins (Vicoria nasuta), from Colombia, new

to the Collection, presented by Dr. H. G. F. Spurrell, F.Z.8., on

April 28th.

1 Merrem’s Xenodon (Xenodon merremii), and 1 Neuwied’s

Viper (Lachesis newwiedii), both new to the Collection, from

Cordova, Argentina, presented by Wilfred A. Smithers, C.M.Z.8.,

on April 28th.

Mrs. R. Hate Tuomas, F.Z.S., exhibited a number of skulls,

head-skins, and photographs of hornless antelopes found by

Mr. A. W. Haig in 1903 on the Dinder River, a tributary

of the Blue Nile. There were two varieties, one larger than the

other. On his return Mr. Haig submitted the skulls, skins,

and photographs to the authorities at the British Museum, who,

while admitting a difference in the formation of the skulls,

stated their view that the evidence for the existence of hornless

antelopes was insufficient and that the specimens shown might

have been females.

In ‘The Nile Tributaries of Abyssinia,’ first published in

1867, Sir Samuel Baker tells us he met with and shot hornless

antelopes on the Royan, a tributary of the Atbara, and we read

farther on that the animal was already known to science and

classified by Riippell. Thus it is shown that Baker’s and Haig’s

hornless antelopes were found on the same watershed, in a

geographical position not a hundred miles apart.

Mr. D. M.S. Watson, M.Sc., F.Z.8., exhibited two specimens

of Procolophon trigoniceps, a Cotylosaurian Reptile from South

Africa, and drew attention to certain sexual differences in this

species.

No. 180.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

March 17th, 1914.

Prof. KE. A. Mincury, 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 made to the

Society’s Menagerie during the month of February, 1914.

Mr. R. H. Burne, M.A., F.Z.8., exhibited a specimen” of the

palate of a female Bactrian Camel and a lantern-slide of a

Hunterian preparation of the throat of a male Common Camel

from the Royal College of Surgeons Museum, showing in each a

fleshy appendage attached to the palate some inches in front of

its free posterior border on a level with the anterior pillars of the

fauces.

_ This appendage, though rudimentary in the female, is of great

size in the male, and during the rutting season can be distended

and protruded from the mouth, and is in some way secondarily

connected with the sexual functions.

Mr. R. J. Pocock, F.R.S., F.Z.S., Curator of Mammals,

exhibited a set of lantern-slides illustrating the structure of the

feet in Domestic Dogs, with special reference to the extension of

the interdigital integument.

Two papers were received from Mr. L. N. G. Ramsay, M.A.,

B.Se., entitled: (1) On the Annelids of the Family Nereide

* 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 Sir Shillings per annum, payable in advance.

collected by Mr. F. A. Potts in the N.. Pacific in 1911, with a

Note on Micronereis as a Representative of the Ancestral Type

of the Nereide ; (2) On the Genera Ceratocephale Malmgren and

Tylorrhynchus Grube.

Mr. A. Kyyvert Torron contributed an account of the

Structure and Development of the Caudal Skeleton of the

Teleostean Fish Pleuragramma antarcticum.

Mr. G. C. Rosgox,.B.A., read a report on Mollusca from

Dutch New Guinea collected by the British Ornithologists’

Union and Wollaston Expeditions. In general, the collection

appears to endorse Hedley’s views as to the Oriental affinities of

the Papuan molluscan fauna. Though numerically small in

species and individuals, the collection has yielded two genera

and three new species, the anatomy of all of which is described.

The two new genera, which were obtained from considerable

altitudes, viz. 10,500 ft. and 14,200 ft. respectively, are of

considerable interest, though their precise affinities are as yet

uncertain. In any case they cannot be regarded as typical

members of the Xonitide, though an aggregate of anatomical

characters exhibits the characters of that family. An account of

the anatomy of Papwina litwus (Lesson) is given, and dis-

crepancy between the anatomical and conchological relationships

of a new species of Papuina is discussed.

This paper will be published in the TRANsacrions in due

course.

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

communicated Mr. P. R. Awati’s paper on “The Mechanism of

Suction in Lygus pabulinus Linn.,” a Capsid bug injurious to

the foliage of the potato, on which it feeds. A detailed de-

scription of the morphology and anatomy of those organs of the

head concerned in sucking the plant-juices is followed by an

account of their mode of action, in part deduced from their

structure and arrangement, in part derived from observation

of the living insect. The potato-leaf is pierced by the conjoined

maxillary and mandibular stylets, the labrum and labium serving

to direct and steady them at the point of puncture. The inner

(maxillary) stylets are grooved along their apposed surfaces in

such a way as to form two canals, one conducting to the cavity of

the pharynx, the other to a salivary pump in connection with the

salivary glands. Suction is effected by muscles which raise the

flexible dorsal wall of the pharynx.

The investigation was conducted by means of serial sections, in

the making of which attention to certain details of technique was

found to be essential. An historical summary is given of work

on the homologies of the mouth-parts of the Rhynchota.

Mr. K. G. Buarr, B.Se., read his report on the Heteromerous

Coleoptera collected by the British Ornithologists’ Union and the

Wollaston Expeditions in Dutch New Guinea.

The most interesting feature of the collection, from the point

of view of distribution, is the occurrence of Cissites maxillosa Fab.

in this region. This Beetle has been hitherto regarded as peculiar

to the Oriental Region, its range extending from Assam to Java,

Borneo, and the Philippine Islands; it has also been found in

Ceylon.

The three species of Amarygmus belong to a section of the

genus that makes New Guinea its headquarters ; a few species of

this section are found in the extreme north of Australia, but the

majority of the Australian species belong to other groups.

Of the fourteen species noted, the following seven are described

as new :—

SETENIS COSTIPENNIS, sp. n.

Atra, parum elongata, capite prothoraceque dense rugoso-punctatis, hoe antice et

postice bisinuato, lateribus crebre crenatis, disco medio longitudinaliter impresso

utrinque obsolete bi-impresso, angulis anticis rotundatis, posticis acutis; elytris

subtiliter punctato-striatis, intervallis opacis plus minusve costulatis, costis nitidis,

intervallo tertio et 5° et 7° basi magis elevatis; corpore subtus pedibusque nitidis

parce punctulatis, tibiis omnibus extus late sulcatis, sulcis opacis. Long. 18-21 mm.

AMARYGMUS VIRIDIZNEUS, Sp. nl.

Ovalis, viridi-wneus, nitidus, corpore subtus pedibusque rufescentibus; sulcis

ocularibus nullis, antennis rufo-piceis, prothoracis lateribus arcuatim angustatis,

dorso vix perspicue punctulato ; elytris striato-punctatis, intervallis vix convexis, vix

punctatis. Long. 11, lat. 6 mm.

AMARYGMUS UTAKWENSIS, sp. nl. -

Elongato-ovalis, seneus sat nitidus, corpore subtus cum pedibus rufo-piceis.

Preecedenti similis, magis elongatus, omnino fortius punctatus, striarum punctis

confertioribus. Long. 103, lat. 53 mm.

AMARYGMUS WOLLASTONT, sp. n.

Elongato-ovalis, nitidus, cyaneus ; prothorace subtiliter punctulato; elytris striatis,

striis sat subtiliter punctulatis, punctis et striis ipsis griseo-nigris, intervallis antice

planis postice convexis; corpore subtus pedibusque nigris subnitidis, tarsis rufo-

setosis. Long. 12, lat. 6 mm.

STRONGYLIUM WOLLASTONI, sp. n.

Nigrum, nitidum, elytris obscure czeruleo-nigris, corpore subtus pedibusque obscure

piceis ; capite inter oculos foveolato, clypeo medio transverse suleatulo, prothorace

transverso, lateribus leviter sinuatis tenuiter marginatis, angulis anticis oblique

truncatis, posticis rectis, disco inzequali medio obsolete sulcato, ante basin trans-

verse impresso, impressione crebre punctata ; elytris basi prothorace valde latioribus

post scutellum leviter impressis, striato-punctatis, punctis suturem et apicem

adversus minoribus, intervallis parum convexis; pedibus sat gracilibus, femoribus

leviter clavatis, tibiis omnibus leviter sinuatis. Long. 15 mm.

MorpDELLa SERICEOBRUNNEA, Sp. n.

Grandis, elongata, brunnea parum iridescens, omnino setulis fulvis sericeis induta ;

capite post oculos expanso angulis fere rectis, palpis brunneis, articulo ultimo tri-

angulariter elongato, antennis gracilibus prothoracis medium haud superantibus,

articulis 4-10 intus serratis vix transversis; prothorace valde transverso elytris

latiore, antice et postice medio valde lobato, disco inter lobos rotundato-elevato,

lateribus arcuatis; scutello elongato-triangulari, apice rotundato; elytrise longatis,

transverse subtiliter rugulosis, sutura marginata, humeris obtuse elevatis; abdominis

apice acute sat breve caudato, tarsis posterioribus compressis. Long. 16, lat. 43 mm,

SESSINIA STOTHERTI, Sp. n.

Fusco-testacea, elytris fuscis; prothorace elongato, apicem versus parum angus-

tato, punctis disci ante medium majoribus et parcioribus ; elytris fuscis, subtilissime

sat dense punctulatis et sericeo-pubescentibus, obsolete tricostatis, ong. 9-

105 mm.

This paper will be published in the Transactions in due

course.

A paper on the Malay race of the Indian Elephant was received

from Mr. R. Lypexxer, F.R.S., F.Z.S. The author made the

young Negri Sembilan Elephant, formerly living in the Society’s

Gardens, the type of a new race, Hlephas maximus hirsutus,

subsp. n., characterized by the square, instead of triangular,

form of the ear, the early date at which its upper margin is bent

over, and the presence in the young condition—at least, in some

cases—of a thick coat of black and in part bristly hair,

Prof. W. J. Daxty, D.Sce., F.L.S., F.Z.8., contributed two

papers on the Fauna of Western Australia, dealing with the

Onychophora and the Phyllopoda respectively.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, April 7th, 1914, at half-past Hight

o’clock p.M., when the following communications will be made :—

EXHIBITIONS AND NOTICES.

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

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.— XIII. On Two new Species belonging to the

Genera Oochoristica and Linstowia, with Remarks upon those

Genera.

EK. W. Suany, B.Sc.

On the Nature of the Lateral Muscle in Teleostei,

W. T. Catan, D.Sc., F.Z.S.

Report on the River-Crabs (Potamonide) collected by the

British Ornithologists’ Union Expedition and the Wollaston

Expedition in Dutch New Guinea.

OLprietD THomas, F.R.S., F.Z.S.

Report on the Mammals collected by the British Orni-

thologists’ Union Expedition and the Wollaston Expedition in

Dutch New Guinea.

Guy DouuMaNn.

Notes on a Collection of East African Mammals presented

to the British Museum by Mr. G. P. Cosens.

The following papers have been received :—

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

Further Contributions to the Anatomy of the Ophidia.

Miss ALBERTINA CARLSSON.

On the Fossil Carnivores Cynodictis intermedius and Cynodon

gracilis from the Phosphorites of Quercy.

The Rev. T. R. R. Stespine, M.A., F.R.S., F.L.S., F.Z.8.

Crustacea from the Falkland Islands collected by Mr. Rupert

Vallentin, F.L.8.—Part 2.

Stanuey Hirst, F.Z.S8.

Report on the Arachnida and Myriopoda collected by the

British Ornithologists’ Union Expedition and the Wollaston

Expedition in Dutch New Guinea.

J.S. Huxtey, F.Z.8.

The Courtship-habits of the Great Crested Grebe (Podiceps

cristatus) ; with an Addition to the Theory of Sexual Selection,

Major J. Stevenson Hamirton, C.M.Z.S.

nn en ST CE RR EE I RE EE RR

The Coloration of the African Hunting Dog (Lycaon pictus).

C. Tarr Reaan, M.A., F.Z.8.

Notes on Aristeus goldiei Macleay, and on some other Fishes

from New Guinea.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL SoctEty OF LONDON,

Recent’s Park, Lonpon, N.W.

March 24th, 1914.

No. 181,

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

April 7th, 1914.

Prof. EK. W. MacBrips, M.A., D.Sc., F.B.S., Vice-President,

in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

The Secretary, Dr. P. Caatmers Mircnect, F.R.S., exhibited

the photograph of a female Orang-utan (Simia satyrus), kindly sent