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PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

- OF THE

AQVOLOGICAL SOCIETY

Oe ON aEOUNE

1915, pp. 299-712,

with 11 PLATES and 121 TEexr-FIcuREs.

PRINTED FOR THE SOCIETY,

SOLD AT ITS HOUSE IN REGENT’S PARK.

LONDON:

MESSRS. LONGMANS, GREEN, AND CO.,,

PATERNOSTER ROW.

Los 6

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL SOCIETY OF LONDON,

15.

Patron.

fis Masesty Tur Kine,

COUNCIL.

His Grace Tas Duke or Beprorp, K.G., F.R.S., President.

RicHarp H. Burne, Esg., M.A.| Guy <A. K. MarRsHALL,

AtrreD H. Cocks, Esq., M.A. Esq.

Tre Rr. Hon. Tur Ear or

Cromer, G.C.B., F.R.S., Vice-

President.

F. G. Dawrrey Drewirt, Esa.,

M.A., M.D.

CHarLes Drummonp, Esq,

Treasurer.

Tue Earu or Dunmore, V.C.,

M.V.O.

ALFRED Ezra, Hsq.

SrpNbEY Freperic Harmer, Hsq.,

Wits iNe, SCoIDE, Jae Sh [ee

President.

Srr EpmMunpD Gites Loner, Bt.,

Vice-President.

Prof. Ernest W. MacBrips,

M.A., D.Sc. F.R.S., Vice- |

President.

PY pene

‘

H. G. B. Meapm-Watpo,

Ksq.

P. CHALMERS MircHewy, Esq.,

Ee, IDSie,, IIb Dy, E.R.S.,

Secretary.

W. R. Oairvie-Grant, Ese.

ALBERT Pam, Esq.

THe Karu or Porrsmouru.

THE Marquzss or Suico,

WS.A., Vice-President.

AuBYN TreEyor-Barrye, Lisq.,

M.A.

ANTHONY

Ksq.

ArraurR Smira Woopwarp,

Hsq., LL.D. F.B.S., Vice-

President.

WIHINGFIELD,

PRINCIPAL OFFICERS,

P. Cuatmers Mircnetn, M.A., D.Sc., IEJURID),, TDLIES.

Secretary.

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

R. I. Pococs, F.R.S., F.L.S., Curator of Mammals and

Resident Superintendent of the Gardens.

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

Epwarp G. BouLencer, Curator of Reptiles.

Prof. H. Maxwenn Lerroy, Curator of Insects.

Hueyry G. Prurmmer, F.R.S., M.R.CS., Pathologist.

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

Joun Barrow, Accountant.

W. H. Coin, Chief Clerk.

LIST OF CONTENTS.

1915, pp. 299-712.

EXHIBITIONS AND NOTICES.

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

Menagerie during the month of April 1915

Core re seseee

Prof. H. Maxwex Lerroy, M.A., F.Z.S8., Curator of Insects.

The House-Fly Exhibition

Bilshsimls) «j,e/ ee |siieisiel/a leirale siecle /nlo/s\sle) elieie) sifctelere

Mavis: ho Pacotn, HSK. S.ehalusS. F.Z.8., Curator of

Mammals. Exhibition of skins of Wild Boar and

Poreupines

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

Menagerie during the month of May 1915

Mr. EH. T. Newron, F.R.S., F.Z.8. Exhibition of horns of

the Sabre-horned Antelope (Oryx UNICO HE eae eone nck

Mr. Grorer Junnison. A “ nest ”-naking Chimpanzee ...

Dr CC. Cunisry, MB. CM. ZS. »Remarkss on the

habits of Chimpanzees in African Forests

Cee esesssoce

Mr. R. KE. Hotpive. Exhibition of horns of West-African

humped Cattle. (Text-figure 1.)

Pease eee esersessecsseeroee

My. Aurrep Ezra, F.Z.8. Exhibition of a living specimen

of Ricordi’s Humming-bird (Sporadinus ricordi) and

remarks upon these birds in captivity

Pereeeeer ser osease

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

Menagerie during the months of June, July, August,

PROC Sejoinsa ee, INET sajosscoganaasepgodsabogposaepsodesc G0"

Mr. R. I. Pocock, F.R.S., F.Z.8., Curator of Mammals.

Exhibition of burrows of Trapdoor Spiders ............

Mr. D. Seru-Smrrn, F.Z.S., Curator of Birds. Exhibition

of a living male hybrid Swinhoe’s and Silver

Pne@asaints W, 2 eee Pleenn emaes 2S es atenins steiae gees sod aaa

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

of an abnormal male Mandarin Duck (4x galeri-

goulana)). (AUesxnstkemes Io) esosn.esactesonescncsoe0ac0 73000)

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

Menagerie during the month of October 1915.........

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

Notice of Report on the House-Fly Investigations ...

Mr. C. J. C. Poon, Assistant Curator of Insects. Notes

joa wave) Opnmael Ibs, IEUOMNY ..cdscoseanssosascoscacosesane

Mr. R. W. Haroitp Row, B.Sc., F.Z.S. Exhibition of

photographs oman abnormal iio s wea eects eeeeeer

PAPHRS.

24. The Head Cavities and Development of the Hye

Muscles in Trichosurus vulpecula, with Notes on

some other Marsupials. By ExizaperH A. FRASER,

B.Sc. (Lond.), F.Z.S., Embryological Laboratory,

Department of Zoology, University of London,

University College. (Plates I. & II., and Text-

fieumes 126.3) isi des teh. 126 2 eee en ee ee eee ene

25. On the Organ of Jacobson and its Relations in the

‘“‘ Insectivora.”—Part II. Talpa, Centetes, and Chryso-

chloris. By Lieut. R. Broom, M.D., D.Sc., C.M.Z.S.,

eA MG@. i) GPlaites duLNocs Jl Ve), eae scanner ee

708

709

=r)

28.

ie)

se)

30.

oA.

._ On the Anomodont Genera, Pristerodon and Trepido-

stoma. By Lieut. R. Broom, M.D., D.S8c., C.M.Z.5.,

R.A.M.C. (Text-figures 1-4.) .........2- cece

_ On the Triassic Stegocephalians, Brachyops, Bothriceps,

and Lydekkerina, gen. nov. By Lieut. R. Broom,

M.D., D.Se:,C.M.Z.S8., R.A.M.C. (Text-figures 1-3.)

A list of the Snakes of Madagascar, Comoro, Mas-

earenes, and Seychelles. By G. A. BouLencsr,

TBIR SL) UIASS (Gee cegonecosncbacsrocb coUsandapne shoes. 3oncebr

Cn a Blood-sucking Gamasid Mite (/choronyssus ser-

pentium, sp. n.?%), parasitic on Couper’s Snake. By

Srawuny Hirsr, F.Z.S. (Text-figures 1 & 2.) .........

On the Feet and Glands and other External Characters

of the Paradoxurine Genera Paradoxurus, Arctictis,

Arctogalidia, and Nandinia. By R. 1. Pocock,

F.R.S., F.LS., F.Z.S., Curator of Mammals. (Text-

figures 1-10.) ........2 cence teense eee ete tener etree ences

_ Anatomical Notes on the Gruiform Birds dramas

giganteus Bonap. and Rhinochetus kagu. By P.

CuAtMers Mrrcueit, D.Sc., LL.D., F.RB.S., F.Z.S.,

Secretary to the Society. (Text-figures 1-5.).........

_ On the Skull of an extinct Mammal related to Aluropus

from a Cave in the Ruby Mines at Mogok, Burma.

By A. Suir Woopwarp, LL.D., F.R.S., V.P.Z.8.

(Plate I. and Text-figure 1.) ........-.:.seeeeeeeeeeeee ees

. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—X VII. On Tenia tawricollis

of Chapman and on the Genus Chapmama. By

Frank E. Bepparp, M.A., D.Se., F.R.S8., F.Z.S.,

Prosector to the Society. (Text-figures 1-6.) .........

Minchinia: A Haplosporidian. By Heten L. M.

PrxeLt-Goopricu, B.Sc., Beit Memorial Research

Fellow. (Plates 1. GIL.) .,.........:eceees eee ee ete ee ees

. The Early Development of the Heart and Anterior

Vessels in Marsupials, with Special Reference to

Perameles. By Kavnartne M. Parxer, B.Sc.(Lond.),

Assistant in the Department of Zoology, University

of London, University College. (Plates I & se

and Text-figures 1-25.) ...........scececesene eset ee eeeneene

383

387

413

A45

36.

40.

4].

43.

44,

aval

On Spiders of the Family Salticidee collected by the

British Ornithologists’ Union Expedition and the

Wollaston Expedition in Dutch New Guinea. By

H. R. Hoge, M.A., F.Z.S. (Text-figures 1-11.)......

. Some Alcyonaria and a Stylaster from the West Coast

of North America. By Sypney J. Hickson, M.A.,

D.Se., F.R.S., F.Z.8., The University of Manchester.

(Gelato st vanade Mex hi sumecel(—o}) see ansec Sere ne eee

. On Specimens of Cuvier’s Whale (Ziphius cavirostris

is i:

from the Irish Coast. By Srpney F. Harmer, Sce.D.,

F.R.S., F.Z.8., Keeper of Zoology in the British

Museum

fee reat oes secon esse ee scrceseres esses aees corse nseesesed

39, Fauna of West Australia.—IIT. A new Nemertean,

Geonemertes dendyi, sp. n., being the first recorded

Land Nemertean from Western Australia. By W.

J. Dakin, D.Se., F.Z.8., Professor of Biology, Uni-

versity of W. Australia. (Text-figure 1.)

cree cco nese sees

Fauna of West Australia.—IV. Palemonetes australis,

Sp.n., being the first record of the genus in Australia.

By W. J. Dakin, D.Sc., F.Z.S., Professor of Biology,

UniversttysonaW. eNtuciraliaye (elace dl) len eee

The Keeping and Breeding of Tasmanian Devils (Sarco-

philus harrisi). By Mrs. Mary G. Rossrrs, C.M.Z.8.,

M.R.A.O.U. (Text-figure 1.)

. Notes on the Feeding of Snakes in Captivity. By E.

G. Boutencer, F.Z.8., Curator of Reptiles

eee cee e scone

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.— X VIII. On Tenia struthionis

(Parona) and Allied Forms. By Frank E. Bepparp,

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

(Text-figures 1-6.)

Some Notes upon the Anatomy of Rana tigrina. By

Guo. E. Nicuonts, D.Sc., late Professor of Biology,

Agra College, India. (Text-figures 1-3.)....:..........

A List of the Snakes of East Africa, North of the

Zambesi and South of the Soudan and Somaliland,

and of Nyassaland. By G. A. Bounmnerr, F.R.S.,

F.Z.S. (Text-figures 1-3.)

Page

501

589

603

611

46.

48.

49.

A List of the Snakes of North-East Africa, from the

Tropie to the Soudan and Somaliland, including

Socotra. By G. A. Boutencsr, F.R.S., F.Z.8. ......

. Descriptions of a new Amphisbena and a new Snake

discovered by Dr. H. G. F. Spurrell in Southern

Colombia. By G. A. Bovunrnesr, F.R.S., F.Z.8.

(exer iieuves ya, 2a) js raates stn tess acer cleenistelen aaccelacne acer

The Distribution of Secondary Sexual Characters

amongst Birds, with relation to their Liability to the

Attack of Enemies. By J.C. Morrram, M.B.(Lond.)

Some Observations on Pattern-Blending with reference

to Obliterative Shading and Concealment of Outline.

By J.C. Morrram, M.B.(Lond.). (Text-figures 1-5.)

. On some Land-Planarians collected in Western

Australia and Tasmania by Members of the British

Association for the Advancement of Science. By

ArtHuR Denpy, D.Sc., F.R.S., F.Z.8., Professor of

Zoology in the University of London (King’s College).

AlpiabeuicalMist: of€ ont ubOrsy) \cudesedees + asim eases

fliar Cle sania vee sane acre erste ere cies decche iat ieanctae etal noua se nceis cial merc tNsistos

Page

641

663

679

653

Ae BoA elk CrAcke ins

OF THE

CONTRIBUTORS,

With References to the several Articles contributed by each.

(1915, pp. 299-712.)

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

to the Society.

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—XVII. On Venia tauricollis of

Chapman and on the Genus Chapmania. (Text-figures

EOS) dae acs err e icuitusn oc Senta s uit wich sn ches ae uenas taaie

XVIII. On Tenia struthionis (Parona) and Allied

Romine, » (lesniatetnes (0, 28804 sbadizns Shasboeesboncasanoosonc

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

Notes on the Feeding of Snakes in Captivity............

Boutencer, GEoren A., F.R.S., F.Z.S.

A List of the Snakes of Madagascar, Comoro, Mas-

CAREERS, BuNGl Say OMNEIIEY goose Hobeocdessarnoo doe cpuscoobeaudodoaes

A List of the Snakes of East Africa, North of the

Zambesi and South of the Soudan and Somaliland, and

on Whgasseleinl (bersisiennees ll =Gis))neenoceseocnapancens saa

Page

583

369

611

Page

Boutenesr, Grorce A., F.R.S., F.Z.8. (continued).

A List of the Snakes of North-East Africa, from the

Tropic to the Soudan and Somaliland, including Socotra. 641

Descriptions of a new Amphisbena and anew Snake

discovered by Dr. H. G. F. Spurrell in Southern

Clolloremlone., | (beruanisionees) We Me) 555 econ st otssosacs asiea stn 659

Broom, Lieut. Ropert, M.D., D.Sc., C.M.Z.8., R.A.M.C.

On the Organ of Jacobson and its Relations in the

‘“‘ Insectivora.”—Part IIT. Valpa, Centetes, and Chryso-

Giultonuss,” (HEE NaeS IE Ler iskse IAe ru koe of bop quasodeocasgadposanes 347

On the Anomodont Genera, Pristerodon and Tropido-

stoma mu (lext-noures al 43) ix, cma ees ncasas ner eee 395

On the Triassic Stegocephalians, Brachyops, Bothriceps,

and Lydekkerina, gen. nov. (Text-figures 1-3.) ......... 365

Curisty, Dr. Curnpurt C., M.B., C.M., F.Z.8.

Remarks on the Habits of Chimpanzees in African

IO ESES Preticcrche cc ogee hathon ie ete ee ter ea en Se Nees Tomine rsloyse eae 536

Daxin, Prof. Witiiam J., D.Sc., F.Z.S.

Fauna of West Australia.—III. A new Nemertean,

Geonemertes dendyt, sp. n., being the first recorded Land

Nemertean from Western Australia. (Text-figure 1.)... 567

Fauna of West Australia.—lV. Palemonetes australis,

sp. n., being the first record of the genus in Australia.

(CENGG sa EO We eve een ORR GEC SER eo hme, Sods acca canon D7 1

Denpy, Prof. ArtHur, D.Sc., F.R.S., F.Z.8.

On some Land-Planarians collected in Western Aus-

tralia and Tasmania by Members of the British Associa-

OM now tne ANGlwATAGErMETNG Oi SYONETO 2245 ceccnecceccennooce 695

Ezra, ALFRED, F.Z.S8.

Exhibition of a living specimen of Ricordi’s Humming-

bird (Sporadinus ricordt) and remarks upon these birds

TTA (CR) ONS Ssonoaencatanbe db ane daneone Soespsue docs: GRebORncheeasooon

Fraser, Miss Exizaseru A., B.Sc. (Lond.), F.Z.S."

The Head Cavities and Development ofthe Eye

Muscles in Trichosurus vulpecula, with Notes on some

other Marsupials. (Plates I. & Il., and Text-figures 1—

Goopricn, Hreten L. M. Pixeri-. See PixeLu-Goopricu,

eave

Harmer, Sipney F., Sc.D., F.R.S., V.P.Z.S.

On Specimens of Cuvier’s Whale (Ziphius cavirosiris)

irom the. lish: Coasts. dscts-cas sossa esse sadee saeco stewie

Hickson, Sypnry J., M.A., D.Sc., F.R.S., F.Z.8.

Some Aleyonaria and a Stylaster from the West Coast

of North America. (Vext-figures 1-5.) .....................

Hirst, Stantey, F.Z.S8.

On a Blood-sucking Gamasid Mite (/choronyssus ser-

pentium, sp. n.(?)), parasitic on Couper’s Snake. (Text-

howe SMUG eerste s.wintn aac Nce es eek a Seon ese aemenr ae

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

On Spiders of the Family Salticide collected by the

British Ornithologists’ Union Expedition and the Wol-

laston Expeditron in Dutch New Guinea. (Text-figures

Ue MLN RR ce sea ce ein ore etnias Same sahara MGR one RelA lrtnage ders eu atiep storas

Hotpine, R. E.

Exhibition of horns of West African humped Cattle.

pe Chetetroances Ibs )i oak, eames aa scnen orca Te OME are ths Ss

54]

385

Page

JENNISON, GEORGE.

A “nest ”-making Chimpanzee.............:.0-2ss2ees seco D3D

Lerroy, Prof. H. Maxwent, M.A., F.Z.8., Curator of

Insects.

Mine Islormgealoihy Woeslniloslinom 25 scocbosacasocececcdnassesenn0e 529

Notice of Report on the House-Fly Investigations ... 709

Mirouent, P. Caaummurs, M.A., D.Sc., LL.D., F.R.S., F.Z.S.,

Secretary to the Society.

Anatomical Notes on the Gruiform Birds dAranwus

giganteus Bonap. and Rhinochetus kagu. (Text-figures

EB, sadesedacndasgasnnevddonesc gdodoscdqecandoasaaoyaasuensaspacssosce 413

Report on the Additions to the Society's Menagerie

during the month of April 1915)... 1c... settee 529

Report on the Additions to the Society's Menagerie

during the month of May 1919.............. steer eeeeee eee D34

Report on the Additions to the Society’s Menagerie

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

TQS. act sae ee MERE RRR Cicaa.ban8% spb asopcon Snomnarusnemuacds ac 705

Report on the Additions to the Society's Menagerie

during the month of October L915 ..........eeeeeeeeeeeee ees 708

Morrram, JAmus C., M.B. (Lond.).

The Distribution of Secondary Sexual Characters

amongst Birds, with relation to their Liability to the

INTHiaOLS Os IMMETINIOS 4c ssagdo sees ses sebdkaous suabesadsodadcasnoco500% 663

Some Observations on Pattern-Blending with reference

to Obliterative Shading and Concealment of Outline.

(Text-figures 1-5.) ......ccecerceceeeee ree ec nese eee enees eens aan iG 19

Newton, Epwarp T., F.B.S., F.Z.8.

Exhibition of horns of the Sabre-horned Antelope

(Oiled Wy Cachet seine comincie noi. Ge au sstneeere antl

NicHotts, Grorce E., D.Sc.

Some Notes upon the Anatomy of Rana tigrina.

(Wextettonces) Lae 5.2 hoe. acadas aeeeasior or SOR ahs tee

Parker, Miss KarHarine M., B.Sc. (Lond.).

The Early Development of the Heart and Anterior

Vessels in Marsupials, with Special Reference to Pera-

meles. (Plates I. & I1., and Text-figures 1-25.) .........

Pixety-Goopricu, Mrs. Heaven L. M.

Minchinia: A Haplosporidian. (Plates I. & IL) ......

Pocock, Reeinaup I., F.R.S., F.L.S., F.Z.8., Curator of

Mammals.

On the Feet and Glands and other External Characters

of the Paradoxurine Genera Paradoxurus, Arctictis,

Arctogalidia, and Nandinia. (Text-figures 1-10.) ......

Exhibition of skins of Wild Boar and Porecupines ......

Exhibition of burrows of Trapdoor Spiders ............

Poot, C. J. C., Assistant Curator of Insects.

Notes from the Caird Insect House...... Feet Be ale

Rogerts, Mrs. Mary G., C.M.Z.8., M.R.A.0.U.

The Keeping and Breeding of Tasmanian Devils

(Sancopiuil ds) Wants). (Wert Meurenln)) erase cece.

Row, R. W. HARo.wp, B.Sc., F.Z.S.

Exhibition of photographs of an abnormal Frog ......

603

459

709

X1V

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

Exhibition of a living male hybrid Swinhoe’s and

Shull eoured BA NWerIC PY aU A eaattacstys orice a 8 Ee PaO nS sr tros. i te

Exhibition of an abnormal male Mandarin Duck (4a

Galeruconlamms” (ALep<-tieavige) 5): sone naBoundonanencs> cshacntescco

Woopwarp, A. Smiru, LL.D., F.R.S., V.P.Z.8.

On the skull of an extinct Mammal related to dlu-

ropus, from a Cave in the Ruby Mines at Mogok, Burma.

(Platem@ rand: Mextotigume dc)s o.0. oi )...ceaeeeee enema ee

Page

707

708

TN DEX.

1915.—Pages 299-712.

[New names in clarendon type.

Systematic references in italics.

(z.8.1.) indieates additions to the Society’s Menagerie. |

Ablabes rhodogaster, var. mayottensis, |

3708.

Acrantophis dumerili, 371.

Agithina viridissima (z. s. u.), 706. |

#Zlureidopus baconi, gen. et sp. n.:

structure (Pl. I. fig. 1), 420. |

Aluroglena cucullata, 648. |

luropus melanoleucus : structure

(Pl. I. fig. 1), 425. |

Aix galericulata: variation (Fig. 1), |

708.

Ahetulla enint, 622, 648.

—— heterolepidota, 625.

— hoplogaster, 622.

Alcyorium arboreun, 548.

Allopora californica, 544,

oculina, H44.

Alluaudina belly, 379.

Amphiophis angolensis, 631.

Amphisbena spurrelli, sp. n.

(Rig. 1), 659.

Amplorhinus giintheri, 629, 652.

—— nototenia, 629, 652. |

——. teniatus, 629.

Anatomy. See STRUCTURE.

Anguis platura, 382, 635, 69d.

Anthropopithecus calvus :

555, 536.

Aotes felinus (Z. 8. L.), O34. |

Aparallactus capensis, 634.

christyi, 634,

concolor, 634, 699.

—— guentheri, O34, |

ethology,

Aparallactus jucksonit, 633.

—— lunulatus, 634.

—— nigriceps, 634.

punctatolineatus, 634.

—— wernert, 633.

ARACHNIDA :

Salticidse from Dutch New Guinea :

systematic, 501; Ichoronyssus

serpentium: ethology, systematic,

3Q°

JOU.

Arachnothera magna (z. s. L.), 709.

Aramus giganteus :

413.

Arctictis binturong : structure (Figs. 2,

4, 8), 387.

Arctogalidia: structure (Higs. 3, 4, 5,9),

387.

Artamus superciliosus (z. s.u.), 705.

structure (Hig. 5),

Artioposthia diemenensis, 699.

Ascyltus penicitlatus, 522.

Aspidelaps lichtensteinii, 6357.

Asthenophis ruspolit, 649.

Atheris ceratophorus, 639.

—— nitschei, 639.

squamiger, 639.

Atractaspis aterrima, 640.

hipostocularis, 640,

conradst, 640.

engdahlii, 658.

- hildebrandti, 632.

irreqularis, 640.

hatange, 640.

—— /eucomelas, 658.

XV1

Atractaspis microlepidota, 640, 658.

phillipsi, 658.

rostrata, 640.

Attus @urvillii, 513.

paykulli, 512

AVES :

Secondary sexual characters, 668 ;

pattern - blending and _ conceal -

ment of outline, 679; Aix galeri-

culata : 708;

giganteus, Rhinochetus

structure, systematic, 413: Spora-

dinus ricerdi : ethology, 938.

variation, Aramus

kagu :

Bathippus montrouziert, var. papuanus,

522.

Barracnta:

Rana tigrina :

Betta pugnax (z.

Britis

584,

gabonica, 638.

nasicornis, 638.

Boa constrictor: ethology, 584.

dumerilii, 371.

dussumicri, 371.

madagascariensis, 301.

regia, G44.

Boethoportia ocellata,

sp. n. (Fig. 1), 501, 502.

Bolieria multicarinata, 372.

Boodon erlangeri, 646.

Fuliginosus, 646,

geometricus, 377.

infernalis, 620; ethology, 584.

lemniscatus, 646

— lineatus, 620, 646.

olivaceus, 620.

virgatus, 620.

Bos bubalus: horns, 538.

Bothriceps australis: structure (Mig. 2),

364.

Bothrophthalmus lineatus, 619.

Boulengerina stormsi, 639.

Brachyophis revoili, 655.

Brachyops laticeps: structure (skull)

(Fig. 1), 368.

Bucephalus typus, 631, bot.

structure, 6053.

8. L.), 034.

638, 657 ;

arietans, ethology,

gen. et

INDEX.

Calamaria meleagris, 649,

unicolor, 632.

Calamelaps polylepis, 632.

unicolor, 632.

Caligorgia fraseri, sp. n

fig, 2); Hig. 4), 553.

Callimico goeldii (z. s. .), 529.

Casarea dussumieri, 371.

Causus defilippii, 637.

lichtensteinti, 637.

resimus, 637, 657

rhombeatus (Fig. 1), 637, 657.

Centetes: structure (Organ of Jacobson)

(Pl. IIT. figs. 8-14), 849.

Cerastes cornutus, 657.

Chalcomitra amethystina (z. 8. u.), 709.

Chamepelia cruziana (2. s. L.), 707.

Chametorius aulicus, 628.

Chapmania tauricollis :

1-6), 429.

Chilorhinophis butleri, 654.

648.

(Pl. I.

structure (Figs.

Chlorophis emint, 622,

gracilis, 625.

—— heterolepidotus, 628,

hoplogaster, 622.

—— irreygularis, 623,

macrops, 620.

neglects, 623, GAS.

schubotzi, 622.

Chrysochloris: structure (Organ of

Jacobson) (PI. LV.), 3851.

Ciconia ciconia (Zz. 8. L.), 705.

Cinnyris chalybeus (z. s. u.), 705.

Crvettictis civetta: structure (Figs.

4, 5), 397

Clavularia moresbii, sp. n. (PI. I.

fic, 4; Hig. 1), 546.

Ca@LenreRAvA

Krom North America ;

systematic, O41.

Celopeltis moilensis, 652.

——- virgata, 620.

Coendu insidiosus (z. s. u.), 709.

—— prehensilis (z. s. u,), 707.

Coluber aulicus, 877

structure,

—— cans, 622,

—— cornulus, 657.

—— florulentus, 647.

—— haic, 636, 655.

INDEX.

Coluber irregularis, 625.

lutrix, 625, 650.

miniatus, 378.

motlensis, 652.

nasicornis, 638.

obtusus, 651.

scaber, 626, 650.

schokari, 658.

sebe, 617, 644.

sibilans, 631, 658.

smythit, 625.

Compsophis albiventris, 574.

Contia africana, 649.

Corallus madagascariensis, 371.

Coronella hotambeia, 628, 651.

nototena, 629, 652.

olivacea, 619, 645. |

——— scheffleri, 624. |

semiornata, 624, 649. |

torquata, 373.

Corvus scapulatus (z. s. u.), 706.

Croeodilus cataphractus (z. 8. u.), 706.

Crotalus atrox: ethology, 585.

CRUSTACEA :

Palemonetes australis: geographi-

cal, systematic, O71.

Cytzxa laodamia, sp. n. (Fig. 11),

520.

sylvia, sp. n. (Fig. 10), 524. |

Dasypeltis scabra, 626, 650.

Dasyprocta agouti (2. 8. L.), O34.

structure, 339 ;

development, (heart, ete.) (Pl. I. |

fivs. 2,3; Figs. 4, 5, 24), 463. |

Dasyurus viverrinus :

1-6), |

Davainea sp.: structure (Figs.

589. |

Dendraspis angusticeps, 636, 656.

antinoril, 656.

jamesonit, 636.

spoestedti, 636.

Dendrophis smaragdina, 625. |

DuyrLorMENT.

Manmarta : Marsupials : eye-mus-

cles, 299; heart and anterior vessels,

459.

Protozoa: Minchinia, 445.

Didelphys marsupialis: structure, de-

velopment (Fig. 26), 537,

XVil

Diolenius albopiceus, sp.n. (Hig. 2),

504.

Dipsadoboa maculata, 380.

Dipsadomorphus blandingit Givgess=2)5

628.

reticulatus, 628.

Dipsas betsileana, 381.

—— blandingii, 628.

colubrina, 379.

—— gaimardii, 380.

—— (Heterurus) gaimardi, var. granu-

liceps, 380.

Dipsina rubropunctata, 630, 652.

Dispholidus typus, 631, 654.

Ditypophis vivax, 651.

Dolichotis magellanicus (z. Ss. u.), O34,

Dromicodryas hernieri, 374.

quadrilineatus, 374.

Dromicus dolicocercus, 379.

sexlineatus, 373.

——— stwinpffid, 374.

Dromophis lineatus, 630, 653.

Dryophylax lineatus, 630, 653.

Dumetia hyperythra (z. s. 1.),

TOG.

Echidna gahonica, 638.

Echis carinatus, 657.

———- ¢oloratus, b6d7.

squamigera, 039.

Elapechis boulengeri, 635.

gueutheri, 639.

moebii (z. 8. u.), 706.

niger, 63).

Elaphis (Bothrophthalmus) lincatus,

619.

EHlapomorphus gabonensis, 633.

Hlaposchema vaillanti, 654.

Elaps trregularis, 640.

Jamesonii, 636.

Elapsoidea boulengeri, 635.

—— guenthert, 639.

—— niger, 635.

Enneoctonus erythronotus (2. s. 1.),

706.

Ephippus @urvitle:, 513.

Kquus chapmanni X HE. zebra (z. s. .), °

70d.

Hrithacus akahige (4, s. 1.), TOY.

Xvill INDEX.

Eryx braminus, 370. GHOGRAPHICAL (con.).

—— muelleri, 644. | Africa, 369, 611, 641; Amphis-

multocarinata, 372. | bena, Herpetodryas: Colombia,

Hteirodipsas colubrina, 379. Aracunipa: Salticide: Dutch New

thebaicus, 618, 644. | 659.

7 Ah -0

Evnonogy. | Guinea, 501.

MAMMALIA: Concealing powers of | Crustacea: Palemonetes australis:

patterns, 679; Anthropopithecus | West Australia, 571.

calvus : (‘nests’), 535, 536; Sarco- | Vuermrpea: Land Planarians: West

philus barrisi, 575. | Australia, Tasmania, 693; Geo-

Aves: Secondary sexual characters | nemertes dendyi : West Australia,

and relation to enemies, 663 ; 567.

concealing powers of patterns, Ca@uyterata: North America, 541.

679; Sporadinus ricordi, 538. Geonemertes dendyi, sp. n.:

Reprinia: Feeding of Snakes in | structure, geographical (Fig. 1), 567.

captivity, 583. Geoplana alba, 701.

Insecta: Concealing powers of pat- baifouri, 697.

terns, 679; species bred in the | —— comitatis, sp. n., 696.

Society's Gardens, 709. | —— dakini, sp. n., 694.

Aracunipa : Ichoronyssus serpen- diemenensis, 699.

tium, 383. | —— flavilineata, sp. n., 695.

Prorozoa: Minchinia, 445. eee flynni, sp. n., 701.

Euhydrina valakadien, 382. mortont, TOL,

Kulampis jugularis: ethology, 538. —— nichollsi, sp. n., 699.

Eunectes murinus: ethology, 583. —— quinquelineata, 698.

tasmaniana, 697.

typhlops, 701.

Felis concolor (z. s. L.), 706. Gisella harrisi (z. s. u.), 707.

eyra (z. Ss. L.), 706. Glauconia algeriensis, 643.

manul (Zz. s. L.), 708. boulengeri, 617.

pardalis (z. s. u.), 707. —— hraueri, 617.

pardus (z. s. L.), 534. —- cairi, 643.

—— conjuncta, 617.

—— dissimilis, 643.

Gastropyxis orientalis, 624. -—— distanti, 617.

smaragdina (Fig. 2), 623. —— emint (Fig. 1), G17, 648.

Gazella rufifrons (z. s. 1.), 706. —— filiformis, 643.

Genetta poensis (z. s. u.), 706. — longicauda, 617.

Gennxus nycthemerus X G. swinhoil —— macrorhynchus, 648.

(isa) Oe —— mucrura, 648.

Geodipsas houlengeri, 378. —— merkeri, 617.

—— infralineata, 378. —— reticulata, 643.

vaueroceg@, 627. Glypholycus bicolor, 619.

GEOGRAPHICAL. whytti, 619.

Mammaria: Aélureidopus baconi: Gonyylophis muelleri, 644.

Upper Burma, 425; Ziphius cavi- Gowionotophis degrijsii, 622.

rostris in British waters, 559. Grayia giardi, 619.

ReptiviA: Snakes of Madagascar, smythii, 625.

to) Y )

etc, and East and North-East ©—— ¢hodloni, 625, 649.

. INDEX.

Hupsidophrys lineata, 624.

Hasarius glaucus, sp. n. (Hig. 9),

523.

Hemirhagerrhis kellert, 629, 651.

Herpetodryas bernierti, 374.

goudoti, 378.

quadrilineatus, 374.

—— rhodogaster, 373.

vicinus, sp. n. (Fig. 2), 660.

Fleterodon defilippii, 637.

—— diadema, 648.

madagascariensis, 376.

modestus, 376.

FHleterolepis capensis, 621.

poensis, 21.

Heteroliodon torguatus, 376.

Heterophis resiinus, 637, 557.

Heterurus arctifasciatus, 381.

Holwropholis olivaceus, 620.

Homalocephalus heterurus, 379.

Homatosoma abyssinicwm, 625, 650.

——- lutrix, 625, 650.

shiranum, 625, 650.

Hloubara undulata (z. s. u.), 529.

Hydrus platurus, 382, 635, 655.

— valakadyn, 382.

Ichoronyssus serpentium, sp. n.

(Figs. 1, 2), 383.

Idiophis vaillanti, 376.

INSECTA :

Pattern-blending and concealment of |

outline, 679; Species bred in the

Society’s Gardens, 709.

Tthycyphus goudoti, 378.

—— miniatus, 378.

Ixulus flavicollis (z. s. u.), 709.

Jotus igneus, sp. n. (Fig. 5), 510.

Lachesis alternatus: ethology, 585.

—— mutus: ethology, 584.

Lamprophis abyssinicus, 645.

—— rogeri, 646.

Langaha alluaudi, 379.

—— erista-galli, 379.

intermedia, 379.

Langaha nasuta, 379.

Leptodira atiarensis, 628.

—— degeni, 628.

hotambeia, 628, 651.

torniert, 628.

wernert, 628.

Leptophis kirtlandi, 631, 6538.

—— lateralis, 374.

Letheobia pallida, 616.

Lioheterodon geayi, 376.

madagascariensis, 376.

—— modestus, 376

voeltzkovit, 376.

Liophidium gracile, 374.

trilineatum, 374.

Liophis imerine, 375.

quinguelineatus, 375.

Liopholidophis grandidieri, 373.

Lophophorus impeyanus (z. s.t.), T05-

Lophura rufa (z. s. .), 705.

Lycodon aulicus, 377.

capensis, 621, 646.

—— fuliginosus, 646.

—— geometricus, 377.

Lycodryas sancti-johannis, 881.

Lycognathophis sechellensis, 377.

Lycophidium abyssinicum, 646.

—— acutirostre, 620.

capense, 621, 646.

jacksonii, 621.

—— meleagris, 620.

-—-— semiannulis, 621.

—— semicinctum, 621.

Lydekkerina, gen. n., 366.

huxleyi: structure (Fig. 3), 866.

Lytorhynchus diadema, 648.

Macropus ruficollis : structure, deve!op-

ment (Figs. 25, 24), 354; (Figs. 21,

22), 483.

MAMMALtIa :

Marsupialia :

ties), development

299; development (heart, etc.),

459; Insectivora: Organ of Jacob-

son in Talpa, Centetes, Chryso-

chloris, 347: °Paradoxurine :

structure (head-cayi-

(eye-niuseles),

Proc. Zoou. Soc.—1915, No. X LIX. 49

MAMMALIA (€on.) :

systematic, 387; Atlu-

reidopus baconi: structure, sys-

425; Anthropopithecus

calvus: ethology, 535, 536; Zi-

phius geographical,

structure, systematic, 559; Sar-

cophilus harrisi: ethology, 575.

Manucodia atra (z. s. L.), 707.

Mehelya scmaliensis, 647.

Meleagris ocellata (z.s. L.), 705.

Menemerus culicivorus, 512.

—— paykulli, 513.

Micrelaps bicoloratus, 632.

boetigeri, 654.

——- nigriceps, 654.

-—— vaillanti, 654.

Micropisthodon ochraceus, 377.

Millepora norvegica, 544.

Mimophis mahfalensis, 381.

Minchinia: development :

(Pls. I., I1.), 445.

Miodon christyi, 633.

—— gabonensis, 633.

—- graueri, 633.

Mopsus mormon, 522.

MorpnoLocy. See Srructure.

Mus sylvatieus wintoni (z.s. 1.), 5384.

Musca domestica: ethology, 529.

structure,

tematic,

cavirostris,

Naia angusticeps, 636, 656.

haie, 636, 655.

, var. melanoleuca, 636.

-—— melanoleuca, 636.

nigricollis, 636, 656.

Nandinia binotata: structure (Figs. 3

4, 5, 10), 387.

Neocrex erythrops (z. s. 1), 709.

Oligolepis macrops, 623.

Onychocephalus arenarius, 370.

dinga, 615.

—— mossambicus, 615.

—— mucruso, 616.

—— tettensis, 615.

—— (Letheobia) lumbriciformis, 616.

Ortygospiza atricollis (z. s. u.), 706.

ethology |

INDEX, :

Oryx leucoryx: horas, 538.

Otaria californiana (z, s. u.), 706.

Palemonetes australis,

geographical (Pl. I.), 571.

Paradisea rubra (z. s. u.), 707.

Paradoxurus: structure (Figs. 1, 4-7),

387.

Paragorgia arborea (Fig. 2), 548.

nodosa, 548.

regalis, d48.

Pararhadinea melanogaster, 376.

Pelophilus madagascariensis, 371.

Perameles nasuta: structure, develop-

ment (Fig. 25), 336; development

(heart, ete.) (Pl. I. figs. 4, 5; Figs.

6-15, 25), 466.

—— obesula: development (heart, etc.)

(Pls. I., I1.; Figs. 1-3, 16-20, 25),

387, 460.

Pericrocotus roseus (z. s. u.), 706.

Phascolarctos cinereus : structure,

development (Figs. 20, 21), 330.

Phascolomys mitchelli ;

development (Fig. 22), 333.

Philothamnus neglectus, 623, 648.

semivartegatus, 623, 648.

Phlogeenas rufigula (z.s. u.), 707.

Phyllomedusa sauvagii (z. s. u.), 709.

Planaria tasmaniana, 697.

Plexippus montrouzieri, 522.

——- paykullii, 512.

Polyodontophis mayottensis, 373.

— rhodogaster, 373.

torguatus, 373.

Potos caudivolvulus (z. s. u.), 707.

sp.

Moy

structure,

| Primnoa willeyi, sp. n. (Fig. 3),

551.

Pristerodon agilis: structure, 358.

brachyops, 358.

mekayt: structure (Fig. 1), 355.

raniceps: structure (Fig. 2),

307.

Prosymna ambigua, 6295.

bocagii, 625.

meleagris, 649.

variabilis, 626.

vasset, 625.

INDEX. XX1

Protozoa : ReEPrILia (con.):

Haplosporidia: Minchinia, develop- Lydekkerina: structure, syste-

ment, ethology, systematic, 445, matic, 363; Snake-feeding in

Psammogorgia teres (Pl. 1., fig. 1;

Fig. 5), 554.

Psammophis angolensis, 631.

biseriatus, 631, 653.

— mahfalensis, 381.

oxyrhynchus, 630, 653,

-—— pulcher, 654.

—— punctulatus, 630, 653,

schokari, 663.

seychellensis, 377.

captivity, 583; Amphisbena,

Herpetodryas: geographical, sys-

tematic, 659.

Rhabdotophis subcaudalis, 375.

Rhagerrhis tritentata, 629, 652.

Rhamnophis jacksonti, 624.

Rhamphiophis oxyrhynchus, 630, 653.

rubropunctatus, 630, 652.

Rhea americana (z. s. L.), 706.

Rhinocalamus dimidiatus, 682.

—— sibilans, 631, 653. — meleagris, 632.

, var. subteniata, 631. Rhinochetus kagu: structure (Fig. 5),

subteniatus, 631. 413.

trinasalis, 631.

Psammophylax variabilis, 629.

Pseudaspis cana, 622. Salticus culicivorus, 512.

Pseudoboa carinata, 657. Sarcophilus harrisi: ethology (Fig. 1),

' Pseudoboodon gasce, 645. 579.

Pseudois nahura (z. s. L.), 709. Seaphiophis albopunctatus, 626, 649.

Pseudoayrhopus ambreensis, 375.

—— dubius, 375.

— heterurus, 375.

Schlegelia wilsoni (2. s..), 707.

Sciurus stramineus (2. s. L.), 707.

Sepedon rhombeatus, 637, 657.

Simia satyrus (z. s. u.), 706.

Simocephalus butleri, 647.

—- capensis, 621.

chanleri, 621.

nyasse, 622.

—— poensis, 621.

unicolor, 621.

Spermestes nigriceps (z. 8. L.), 707.

Sphodromantis gastricus (4. 8. L.):

ethology, 710.

guttata (4. s. L.): ethology, 709.

Sporadinus ricordi: ethology, 538.

Stenophis arcti fasciatus, 381.

—— betsileanus, 381.

—— gaimardii, 380.

granuliceps, 380.

twmering, d79.

—— microps, 375.

eccipitalis, 375.

—— quinquelineatus, 375.

—— subcaudalis, 379.

Ptilopachys fuseus (z.s. u.), 706.

Ptinus pusillus (z. s. u.), 711.

-—- tectus (z.s.L.): ethology, 711.

Python regius, 644; (2. 8. u.), 706.

reticulatus: ethology, 583.

— sebe, 617, 644.

Pythenodipsas carinata, 627.

Rana temporaria: structure, 712.

tigrina: structure (Figs. 1-3),

603. guentheri, 380.

Rangifer tarandus (z. s. L.), 534. inornatus, 380.

Reerrwia : —— longicauda, 381.

maculatus, 380.

variabilis, 381.

Stenostoma cairi, 645.

conjunctum, 617.

-—— dissimile, 643.

Snakes of Madagascar, etc., East and

North-East Africa: systematic,

369, 611, 641; Pristerodon, Tro-

pidostoma: structure, systematic,

355 ; Brachyops, Bothriceps,

Stenostoma longicaudu, 617.

macrorhynchum, 643.

STRUCTURE,

Mamaia: Head-cavities and eye-

muscles in Marsupials, 299; Or-

gan of Jacobson in Talpa, Centetes,

Chrysochloris, 347; Paradoxurus,

Arctictis, Aretogalidia, Nandinia

(feet, glands, etc.), 387 ; Aflurei-

dopus baconi (skull), 425; Ziphius

cavirostris, 559; pattern-blend-

ing: experimental analysis, 679.

‘Aves: Aramus giganteus, Rhino-

chetus kagu, 413; secondary sexual

characters, 663; pattern-blending:

experimental analysis, 679.

Reprinta: Pristerodon, Tropidosto-

ma, 355; Brachyops, Bothriceps,

Lydekkerina, 363.

Barracnta: Rana tigrina, 603.

Insecra: Pattern-blending :

mental analysis, 679.

experi-

Vermipua: Trenia tauricollis, 429 ;

Geonemertes dendyi, 569; Da-

vainea sp., 589.

Ca@Lenrerata: Stylasteride (gono-

phores), 545,

Sturnia sinensis (z. s. L.), 706.

Stylaster (Allopora) norvegicus (Pl. J.

fie. 5), 544.

Sus scrofa: skin, 534.

Sylvicapra coronata (z. s. 1), 706.

Sylvilagus superciliaris (z. s. u.), 534.

Tuchymenis boulengeri, 378.

infralineatus, 378.

Tenia struthionis: structure, 589.

tauricollis: structure (Figs. 1-6),

429.

Talpa: structure (Organ of Jacobson)

(Pl. III. figs. 1-7), 348.

Tarbophis guentheri, 627, 651.

—— obtusus, 651.

senvannulatus, 627.

Taurotragus oryx (z. s. u.), 706.

Tayassu tajagu (2. s. L.), 534.

Telamonia mandibulata, sp. n.

(Fig. 4), 508.

INDEX.

Telamonia vidua, sp. n. (Fig. 3),

505.

Telescopus semiannulatus, 6217.

Testudo galapagoensis (z. 8. u.), 706.

—— nigrita (z. 8. u.), 706.

Thaumaglossa bimaculata (z. s. 0.) :

ethology, 710.

Thelotornis kirtlandii, 631, 653.

Thrasops jacksonit, 624.

—— rothschildit, 624.

Tigrisoma salmoni (z. s. u.), 707.

Trichosurus vulpecula: structure, de-

velopment (head-cavities, eye -mus-

cles) (Pls. I., II.; Figs. 1-19), 299.

Trimerorhinus tritentatus, 629, 652.

vartabilis, 629,

Tropidonotus dolichocercus, 373.

grandidiert, 373.

lateralis, 374.

—— olivaceus, 619, 645.

sexlineatus, 373.

stumpfit, 374.

Tropidostoma microtrema :

(Figs. 3, 4), 358.

Typhlops acuttrostris, 642.

— adolphi, 615, 642.

—— arenarius, 370.

blanfordii, 642.

hocagit, 615.

structure

— hoettgeri, 370.

braminus, 370, 614,

comorens?s, 370.

cuneirostris, 642.

decorset, 370.

| ——- dinga, 615.

gierre, 615,

—— gracilis, 614.

| —— grandidiert, 371.

hottentotus, 616.

humbo, 616.

Ne Abe, Kleebergi, 616.

—— latirostris, 616.

| -—— luinbriciformis, 616.

—- madagascariensis, 371.

mandensts, 615.

microcephalus, 370.

mossambicus, 615.

—— mucronatus, 370.

—— mucruso, 616.

INDEX. XX111,

Typhlops obtusus, 610. Vipera arietans, 638, 657.

-_— pallidus, 616. —— hindii, 638.

—— platyrhynchus, 615. superciliaris, 638.

—— punctatus (Fig. 1), 615, 642. Viverricula malaccensis: structure

—— schlegelit, 615. (Big. 5), 399.

—— socotranus, 642.

—— somalicus, 642.

—— tettensis, 615.

——- torniert, 615.

—— uniteniatus, 616, 642. Xenocalamus bicolor, 632.

Xiphosoma madagascariense, 371.

Walterinnesia egyptia, 656.

Uriechus concolor, 634, 655.

jacksonit, 633.

: Zamenis brevis, 647,

—— lunulatus, 634. |

ee — citernii, 647.

nigriceps, O34, —— florulentus, 647.

—— rhodorhachis, 647.

VARIATION. —— smithii, 647.

Avus: Aix galericulata, 708. —-— socotre, 648.

VERMIDEA: —— somalicus, 647.

Cestoda: Twnia tauricollis, Chap- , Zenodoris dane, sp. n. (Fig. 6),

mania: structure, systematic, 429 ; ol4.

Teenia, Dayainea: structure, sys- | —— d@wrvillii, 518.

tematic, 589; Nemertinea from | ——— rhodope, sp. n. (Fig. 7), 517.

Western Australia and Tasmania :

Syrinx, sp. n. (Fig. 8), 519.

systematic, 693; Geonemertes | Ziphiws cavirostris: structure, geogra-

dendyi: structure, systematic, 567. phical, 559.

Proc. Zoou. Soc.—1915, No. L. 50)

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EXHIBITIONS AND NOTICES.

The Szcrerary. Report on the Additions to the Society’s Menagerie during the month

OF AVAL VOLS | sisie ele sie ve 60's s -u.e ais cre ime IR eT REARS REO “cote vse eeitere te sre gete ie cae

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Mr. Acrrep Ezra, F.Z.8. Exhibition of a living specimen of Ricordi’s Humming-bird

(Sporadinus ricordt) and remarks upou these birds in captivity..........-..0ee eee

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EYE-MUSCLES IN TRICHOSURUS.

PROCEEDINGS

Or THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

Or THE

ZOOLOGICAL SOCIETY OF LONDON.

PAPERS.

24. The Head Cavities and Development of the Hye

Muscles in Trichosurus vulpecula, with Notes on some

other Marsupials. By Hnizasera A. Fraser, B.Sc.

(Lond.), F.Z.S., Embryological Laboratory, Department

of Zoology, University of London, University College.

[Received April 23, 1915: Read May 11, 1915.]

(Plates I., II.* and Text-figures 1-26.)

INDEX. - Page

TG ROM UG ET OTR ES ae kee Ee ak eee hice eee ROD)

Material . ES, 2 Rene ey ee OU)

The Eye Muscles of the Adult Shins Met ee ee eo eS (SS

Description of Material ....... scosuspen GY

Summary of Events in Tr ichosurus s vulpec THE Nawab 328

NobessonoghersMansupialsi-ceey. teeter eee eee oO.

TP LGUSCOLELRCLOS MC ie eon a ho ee SO)

BRASCOLOMGYS HE tame isaeeche ces shae:c qosd eae, OOO

MIG CR OMUS Brn scadenins Sah asiite sree sie tem nL eye seaeees Ayete ea Oe

TACT ONVEVES NPR oe Re is ene rk oi LR Ce See EO ‘

BDC NOU Sp cechidencpe Pose ce aneneder ub sto saa OCCA EC, LE

YD ASU OURS os ake sap couse cocoon dina SPE SUCH EH ORCA DORA 5)

Guneludice Remarks Boe rE acide Meu nGung GAC OB ET OMB RCD OCORR OLSTEA

Bibliography .. NE ane aA Sheek Gar Mea ee Re orn eee liry 2 12)

INTRODUCTION.

The following investigation of the head-cavities and develop-

ment of the eye muscles of Zrichosurus vulpecula was undertaken

at the suggestion of Professor J. P. Hill, whose help and advice

I gratefully acknowledge. All the material examined was

* For explanation of the Plates see p. 346.

Proc. Zoot. Soc.—1915, No. XXII. 22

300 MISS E. A. FRASER ON THE DEVELOPMENT

obtained from Professor Hill’s excellent collection of young stages

of marsupials.

Cavities in the mesoderm of the anterior (preotic) region of the

head, bounded by more or less definite epithelial walls, have been

described in many vertebrates, and they are now considered by

the majority of observers to represent somites comparable with

those of the trunk. They were first observed in Selachians

by Balfour (’78), later by Milnes Marshall (81), van Wijhe (83),

Dohrn (90) (04), and since then by many other investigators

in different fishes.

In Amphibians, cavities have been seen by Scott and Osborne

(79) in the newt and in the Gymnophionan Hypogeophis by

Marcus (’09). They appear to be absent in WVecturus (Julia

Platt, ’94); in the frog (Corning, 99) and in the toad (Kdge-

worth, 799) the anterior somites in the head are apparently

solid, although Goette (’75), in his classical work on the develop-

ment of the toad, appears to refer to them as cavities.

Van Wijhe (’86) was the first to point out the occurrence of

head-cavities in birds and reptiles. In birds, Rex has more

recently studied them in the duck (97) and in the gull (01)

(05), Edgeworth (07) in the fowl, and Professor Hill has directed

my attention to the occurrence of large premandibular cavities

in sections of early emu chicks in his collection.

The conditions in reptiles concern us more closely and may be

considered in greater detail. In this class three somites have

been observed in the preotic region of the head, of which the

first or premandibular develops into a large and conspicuous

cavity surrounded by epithelial walls, the cavities of either side

at some time of their development being usually connected across

the middle line by a transverse canal as in fishes. The second

and third somites are small and more difficult to determine, and

observers differ considerably in their accounts of them. Van

Wijhe (86) did not discover a second somite in Lacerta and he

describes the third somite as solid; Hoffmann (90), on the other

hand, finds three hollow somites behind the premandibular, but

whether the last two both belong to the third head somite or

whether they correspond to somites three and four of fishes, he

was unable definitely to decide. Again in Lacerta, Corning (’99)

recognises the third somite but no similar second somite, the

m. obliquus superior, which usually develops from the latter, here

being described as arising from the dorsal part of the primordium

of the trigeminal musculature which grows out over the eye. In

Anguis fragilis, both the second and third somites were seen by

Oppel (90), each of which contained a small central cavity, but the

second was much less distinct than the third. In the Chelonia

the same somites are present, but whereas in Hmys lutaria,

according to Filatoff ('07), the third is solid, in Chelydra serpen-

tina, described by Johnson (13), both second and third possess

a distinct cavity round which the cells are arranged in a radial

manner.

OF THE EYE MUSCLES IN MARSUPTALS. 301

The occurrence of large anterior head-cavities followed by

corresponding structures behind them, has led to much discussion

on the metamerism of the vertebrate head. As one advances

in the scale of vertebrates, however, the segmentation in the

mesoderm of the preotic region of the head becomes more and

more obscure, until in the Mammalia only traces remain of a

former metameric condition. No definite head-cavities have

ever been observed in mammals, although probable remnants of

these have been found in man by Zimmermann (98). In the

rabbit, although the first head somite arises muchas in reptiles

from the fore-end of the alimentary canal as a solid outgrowth

(Corning ’99), or as a hollow one soon becoming solid (Edgeworth

11), it never acquires a cavity as in lower vertebrates but in

later stages forms a small solid mass of cells behind the optic

cup on each side, difficult to distinguish from the surrounding

tissue. Reuter (97), who has studied the pig, sees there no signs

of primitive segments in the head and considers the eye muscles

as a completely independent formation arising from an accumu-

lation of mesenchyme cells; his observations, however, begin at

a late stage and no early embryos are described.

It was first noted by Milnes Marshall (’81) in Elasmobranchs

and is now well established, that the walls of the premandibular

cavity give rise to four muscles of the eye, the mm. recti superior,

inferior and internus and the m. obliquus inferior, all innervated

by the oculomotor nerve; that the second head somite contributes

the m. obliquus superior supplied by the trochlear nerve, while

the third furnishes the m. rectus externus and, in reptiles and

mammals, the m. retractor bulbi, both these muscles being supplied

by the abducens nerve. We have an exception to the usual

conditions in Petromyzon, where the innervation of the muscles

is peculiar. According to Fiirbringer (75, p. 70), ‘ Die Inner-

vation der Augenmuskeln anlangend besteht bei den Petromy-

zonten die Kigenthiimlichkeit, dass, wihrend der Oculomotorius

der Fische simmtliche Augenmuskeln mit Ausschluss des Rect.

ext. und Obliq. sup. versorgt, hier der Rect. inf. nicht vom Oculo-

motorius, sondern vom Abducens innervirt wird, so dass also der

Abducens 2 Augenmuskeln versorgt. Dieses Verhalten ist

vielleicht dahin zu deuten, dass der Rect. int. den vereinigten

Rect. int. und Rect. inf. entspricht und dass in gleichem Masse,

wie diese beiden Muskeln sich spiaterhin in 2 gesondert, der

Rect. inf. mit dem Rect. ext. verschmolzen, einen einzigen durch

den Abducens innervirten Muskel vorstellend.”

The conditions in Amphibia also do not seem to coincide with

those in other groups and the views of different authors are very

conflicting. Marcus (10) has studied the Gymnophiona in some

detail, and according to his account of Hypogeophis the mandibular

somite, at first connected by a transverse bridge with its fellow

of the opposite side, develops into a large cavity surrounded by a

single layer of epithelial cells lying postero-dorsally to the eye,

and from which a cell-strand grows down into the mandibular

29%

fel el

302 MISS E, A. FRASER ON THE DEVELOPMENT

arch (see his fig. V, p. 139, Marcus, 09). The premandibular

cavity, which is small and has also in early stages a connecting

median portion, partly degenerates, but a smal] part moves up to

the mandibular cavity and completely unites with the rostral side

of the latter. All the eye muscles, except the m. rectus externus

derived from the third somite, take their origin from the

mandibular cavity which is innervated by the oculomotor and

perhaps also by the trochlear nerve. Marcus (’10) further states

(p. 409) :—‘“ Daher glaube ich, das, wenn auch Pramandibular-

zellen sich mit der Mandibularmasse vereinigt haben, nicht

deswegen der Oculomotorius die Mandibularhohle innervirt,

sondern dass er ganz primar der zugehérige Nerv ist. Dies

ergiebt sich, ausser durch die naive Betrachtung, auch durch

folgende Ueberlegung. Aus dem rostralsten Teil der Mandibu-

larhohle entwickelt sich der Muse. obliquus superior, also ein vom

Trochlearis innervierter Muskel. Die Zellmasse, woraus er sich

entwickelt, glaube ich aus der Priimandibularmasse ableiten zu

konnen. Exackt beweisen kann ich diese Behauptung freilich

nicht, weil bei der Muskelbildung die Mandibularhéhle schon

mesenchymatés war, also die Muskelbildung nicht so scharf wie

bei Selachiern verfolgt werden kann; aber aus der Topographie

der Gebilde ergibt es sich, das zur Bildung des Muse. obliquus

superior wie bei Selachiern so auch bei Hypogeophis dieser

Abschnitt der Mandibularhohle verwendet wird.”

Although many authors have given accounts of the origin of

these preotic somites and of the first development of the eye

muscles from their walls, few have traced in detail the further

growth of these muscles until their adult position is reached.

In reptiles and mammals this latter growth is more difficult to

follow owing to the development of the m. retractor bulbi. The

most complete account in reptiles is that of Chelydra serpentina

by C. E, Johnson (13), and the only description in mammals is

that of Reuter’s paper (97) on the pig, above mentioned. The

following observations on the marsupial Z'richosurus vulpecula

will therefore be of some interest.

. MATERIAL.

The material examined comprises twelve stages of Trichosurus

vulpecula, eight embryos and four pouch young of Phascolarctos

cinereus, five embryos of Phascolomys mitchelli, an embryo and a

foetus of Jacropus, five embryos and one pouch young of

Perameles, one embryo of Didelphys and a good series of

Dasyurus.

Two wax-plate models of the optic cup and primordia of the eye

muscles were made and are reproduced on Plates I. & Il. Tam

much indebted to Miss EK. A. Steele for her excellent coloured

drawings of these models. The terminology of the arteries and

veins in the head has been adopted from Grosser’s account of the

development of the vascular system in the Chiroptera (01) and

OF THE EYE MUSCLES IN MARSUPIALS. aU

also from the description of the development of the veins of the

head in reptiles by Grosser and Brezina (’95).

THe Kye Muscuss or THE ADULT.

Adult specimens of 7'richoswrus and Dasyurus were examined,

and the usual eye muscles, including a well-developed m. retractor

bulbi, were found to be present. In 1902, Corning (’02)

described in some detail the eye muscles of the cat, and his

account is generally applicable to the recti and oblique muscles

of the marsupial. The only noteworthy difference isin regard to

the form of the m. rectus externus which in marsupials has a

double origin, the two portions uniting to have a common

insertion on the eyeball.

The m. retractor bulbi shows considerablevariation in different

mammals. When present, it always arises further caudally than

the m. rectus externus but enters the orbit together with the

mm. recti and the m. obliquus superior, and its insertion in the

bulbus lies nearer the optic nerve than the other muscles.

According to Motais (87), who has studied this muscle in many

mammals, the m. retractor bulbi attains its maximum develop-

ment in ruminants ; he also observed its relatively large size in

the opossum. He further states, p. 56: ‘Le muscle choanoide

peut offrir un ou plusieurs interstices celluleux qui le divisent en

deux ou plusieurs parties. Ces interstices sont larges dans les

carnivores et séparent le muscle en quatre faisceaux bien distincts.

Ordinairement, les lignes de separation sont moins nettes; dans

les solipédes et les ruminants, les bords des deux divisions du

muscle s’envoient réciproquement des fascicules.... Dans le

pore, on ne trouve qu'un seul interstice assez large au milieu du

muscle droit inférieur.” Similar conditions to those in the pig

are found in rodents; the same author, in his description of the

rabbit, p. 206, says: “Il [le muscle choanoide] forme un céne

régulier qui n’est interrompu que par un seul interstice celluleux

situé au-dessous du muscle droit supérieur. Cet interstice se

prolonge jusqu’a l’insertion bulbaire du nerf optique et le long du

nerf lui-méme, jusqu’au trou optique.” In the cetaceans (Weber,

’86) the same variations occur; here again, the muscle may be

present as an almost complete circular sheet surrounding the

optic nerve or it may be divided into four parts, each more or less

united by connective tissue.

In Trichosurus and Dasyurus, aS in other mammals, the

m. retractor bulbi has a more deeply seated origin than the other

muscles. It arises from the basisphenoid and passes forwards to

enter the orbit between the two portions of the m. rectus externus.

Surrounding the optic nerve, it extends outwards as a muscular

sheet which gradually increases in circumference up to its insertion

round the inner side of the eyeball within the recti muscles.

Although appearing as a completely closed cone, the two edges of

the sheet are not fused along the anterior side ‘of the nerve just

behind the m. rectus internus.

304 MISS BE. A. FRASER ON THE DEVELOPMENT

Descriprion oF MATERIAL.

Trichosurus vulpecula. Kmbryo of G.L. 5 mm. (=Stage I.a,

6 &c)*. (Text-figs. 1, 2.)

This stage is represented by three embryos, a (6 ’97), 6 (LA.’01),

and ¢ (1.01), of whieh (a) and (c) are cut horizontally and

(6) longitudinally.

The cavity of the optic vesicle is flattened by the thickening

and partial invagination of its lateral wall, and it is connected

with the brain by a hollow stalk. The ectoderm opposite the

vesicle is thickened.

Text-figure 1.

Trichesurus vulpecula, -G.L: 5mm. (6°97).

Horizontal section through the head (S 3-4-5), somewhat oblique, passing through

the premandibular head-cavity of the left side. XX 75 and reduced by +.

a.c.a.=arteria cerebri anterior, #'B.=fore-brain. G.V.=trigeminal ganglion.

U.h.c.=lett head-cavity. ».=portion of cavity constricted off from the rest.

op.v.=optic vesicle. v.c.m.=vena capitis medialis.

There are two large and conspicuous head-cavities, one on each

side immediately posterior to and on the medial side of the optic

vesicle, situated ventro-laterally to the fore-brain; between the

cavity and the brain runs the anterior cerebral artery. The

cavity is lined by a single layer of epithelial cells which is more

attenuated on the side. farthest from the brain, where it cannot

everywhere be distinguished in the sections. The shape of the

cavity varies slightly in the three embryos, being obliquely

* Vide Fraser and Hill: “The Development of the Thymus, Thyroid and

Epithelial Bodies in Trichoswrus vulpecula,” Phil. Trans. Roy, Soe, 1915.

OF THE EYE MUSCLES IN MARSUPIALS. 305

rectangular in cross-section in (a) (text-fig. 1, /./.c.), where on the

left side it measures -26 mm. in a dorso-ventral direction, °31 mm.

antero-posteriorly and -32 mm. transversely ; but it is more oval

in (5), and tapers somewhat towards the eye (text-fig. 2,7.h.c.). On

both sides in (@) a small medial portion is marked off from the

main body of the cavity by a constriction, the upper part of which

can just be seen in text-fig. 1, m. No such condition is present in

(6) or (ce), although on the left side of the former there is an

incomplete partition across the middle of the cavity ; the division

in this case, however, may not have the same significance.

Trichosurus vulpecula, G.L. 5mm. (LA. ’01).

Longitudinal section (S 2-1-13) passing through the premandibular head-cavity

of the right side. XX 75 and reduced by 3.

FB.=fore-brain. HB.=hind-brain. MB.=mid-brain. op.v.=optic vesicle.

r.h.c.=right head-cavity. v.c.m.=vena capitis medialis.

In embryo (¢), lying obliquely along the postero-lateral wall of

the cavity, on the inner side of the vena capitis medialis, is a

shghtly more condensed portion of the mesenchyme, representing

the third or hyoidean somite and constituting the primordium of

the abducens muscle-mass, It runs forwards and laterally just

below the origin of the vena cerebralis anterior into another mass

of loosely connected cells, which extends downwards round the

lateral side of the vena capitis medialis to become united below

306 MISS E. A. FRASER ON THE DEVELOPMENT

with the maxillo-mandibular mesenchyme. On its anterior side

this group of looser cells, to which in following stages I have given

the name of the intermediate mass and which represents the second

or mandibular somite, projects slightly forwards anterior to the

cavity : this projection, as we shall see later, is the first indication

of the primordium of the m. obliquus superior. The intermediate

mass is, at the same time, connected with the postero-lateral

wall of the cavity for three or four sections. The limits of

the complex formed by the intermediate and abducens masses

are difficult to determine accurately, as they are not easily

distinguished from the surrounding mesenchyme and in embryo

(a) are still more indefinite. In the longitudinal series, a distinet

but small elongated collection of cells stretches along the posterior

border of the cavity, apparently representing the abducens mass,

but no connection with any maxillo-mandibular mesenchyme is

recognisable.

Text-figure 3.

Prichasurus vulpecula. Glu. 45 mm. (3 98).

Transverse section (S3-6-13) passing through the premandibular head-cavity on

each side. he sections are rather oblique, the cavity of the left side being cut

nearer its ventral end. he anterior portion of the abducens muscle-mass lies

posterior to the cavity. X 110 and reduced by +.

@.c.a.=arteria cerebri anterior. FB.=fore-brain. U,h.e,=left head-cavity. 0p.v.=

optic vesicle. 7.e.=anterior portion of abducens muscle-mass. rh.c.=vight

head-eavity. ,c.m.=vena capitis medialis. v.0.i.=vena orbitalis inferior.

The Gasserian ganglion is large and the mandibular branch of

the trigeminal nerve is already well developed. The oculomotor

nerve runs from the mid-brain anteriorly for a short distance but

does not reach the head-eavity.

' OF THE EYE MUSCLES IN MARSUPIALS. 307

Another embryo (8 798) measuring 4:5 mm. (= Stage IT a) at

its greatest length, may be mentioned here. It is cut trans-

versely and almost at right angles to the last and is slightly

older, the optic vesicle being still further invaginated.

The head-cavities are narrow from side to side and have the

form in the sections of an elongated triangle, the long pointed

end of which runs ventrally and medially (text-fig. 3, r ie).

They are lined by a single layer of epithelial cells which can be

distinguished all round the eavity but which, on the side next

the optic vesicle and extending round this side laterally for a

short distance, 1s considerably attenuated. The exact limits of the

abducens muscle complex are again very indefinite (text-fig. 3, 7.¢.).

Embryo of G.L. 7 mm. (a’97) (=Stage IIT.).

(Text-figs. 4-6.)

The cavity of the optic vesicle is very narrow, the outer or

retinal layer being much thickened and well invaginated and the

adjacent ectoderm now forms a rounded solid inbulging, the

prumordium of the lens.

Text-figure 4.

Trichosurus vulpecula. G.L. 7 mm. (4797).

Horizontal section through the head (S 3-2-11), passing above the optic vesicle and

through approximately the middle of the head-cavity of the left side (/.h.c.), to

show the proliferation from postero-medial wall of the cavity (p.m.). The

ventral end of the mtermediate mass is seen postero-laterally to the cavity.

X 110 and reduced by .

a.c.a.=arteria cerebri anterior. I’B.=fore-brain. dnt.m.=intermediate mass.

v.0.4.=Vvena orbitalis inferior.

The premandibular head-cavity has only increased slightly in

size but changes are to be seen in its walls. About half-way

down the cavity on the postero-medial side, the wall has thickened

308 MISS E. A. FRASER ON THE DEVELOPMENT

and is now composed of several layers of cells (text-fig. 4, p.m.);

immediately ventral to this and on the postero-lateral side occurs

a much more extensive proliferation, which extends down as far

as the end of the cavity but does not involve the extreme ventral

wall (text-fig. 5, p.J.). Atits lower end, however, a part of this

thickening stretches out laterally behind the bulbus and forms

the first indication of the m. obliquus inferior (text-fig. 5, o.inf.).

On the left side, the two proliferations appear to be separate

from each other, but on the right the more dorsal one extends

round at its lower end to join the larger ventral. Except where

proliferation is proceeding the wall of the cavity is very thin ;

on the side next the brain it consists of a single but perfectly

definite epithelial layer, but on the side next the optic cup it is

much less definite (text-figs. 4 & 5).

Text-figure 5.

Trichosurus vulpecula. G.L. 7 mm. (@ 97).

Horizontal section through the head (S 3-3-9), passing through the dorsal portion

of the optic vesicle (op.v.) and the ventral portion of the head-cavities (l.h.c. &

r.h.c.), to show the proliferation from the postero-lateral wall (p./.). The

primordium of m. obliquus inferior (0.inf.) is seen on the left side. X 75 and

reduced by 3.

a.c.a.=arteria cerebri anterior. J'B.=fore-brain. hp.=hypophysis.

ph.=pharynx. v.0.i.=vena orbitalis inferior.

The primordium of the abducens muscle-mass consists of a

postero-dorsal portion which lies along the medial side of the

vena capitis medialis (text-fig. 6, 7.b.) and which, at its anterior

end, runs ventrally outwards to join with the larger anterior

portion of the mass (text-fig. 6, 7.e.). The latter, whose dorsal

side surrounds the root of the vena cerebralis anterior, lies

anterior to the vena capitis medialis and extends half-way down

OF THE EYE MUSCLES IN MARSUPIALS. 309

the head-cavity close to the postero-lateral wall of the latter, its

lateral end running into the intermediate mass, the limits of the

two being indistinguishable (text-fig. 6, imtm., and also the

ventral end in text-tig. 4, int.m.), The intermediate mass itself,

which is composed of more loosely connected cells, passes on the

one hand, forwards into a mass of condensed mesenchyme lying

above the optie vesicle on the outer side of the ophthalmic nerve,

this portion representing the primordium of the m. obliquus

superior (text-fig. 6,0.s.); and on the other hand, backwards and

Text-figure 6.

Trichosurus vulpecula. G.L. 7 mm. (a 797).

Horizontal section (somewhat diagrammatic) through the head (S 3-1-11), showing

the posterior (r.b.) and anterior (7.e.) portions of the abducens muscle-mass of

the left side, which extend in front of the vena capitis medialis (v.c.m.), just

ventral to the vena cerebralis anterior (v.c.a.), to unite with the intermediate

mass (int.m.). The primordium of the m. obliquus superior (0.s.) is seen as a

forward outgrowth from the intermediate mass. Owing to the obliquity of

the sections, the right side is cut at a more dorsal level than the left and only

shows the posterior portion (r.b.) of the abducens mass lying medial to the

vena capitis medialis (v.c.m.) and the vena cerebralis anterior (v.c.a.). 75 and

reduced by 3.

a.c.a.=arteria cerebri anterior. J.h.c.=left head-cavity. n¢.=notochord.

downwards lateral to the vena capitis medialis and is loosely

connected with the maxillo-mandibular muscle-mass, the con-

nection here not being well seen. The union of the intermediate

mass with the postero-lateral wall of the cavity is most probably

again present.

The oculomotor nerve is now thicker and its distal end

terminates dorso-laterally to the head-cavity.

310 MISS E. A. FRASER ON THE DEVELOPMENT

From the Gasserian ganglion arise a well-developed mandibular

nerve, a small maxillary branch and an ophthalmic nerve.

The abducens is here seen for the first time. It leaves the

mid-brain by many fine fibres which unite and run forwards

parallel to the side of the brain for a short distance.

Another example of this stage, which measures 7°5 mm.

(XTX. ’04) (=Stage IV.), presents almost the same features.

Although the sections are cut in the same direction, that is

horizontally through the head, the head-cavity in section is more

triangular and less elongated so that its area is larger than in

the 7 mm. embryo above described. The postero-medial and

postero-lateral proliferations form one thickening and the vena

cerebralis anterior appears to be only slightly surrounded by the

anterior portion of the abducens muscle-mass.

The conditions in this embryo, however, are difficult to make

out accurately.

Embryo of G.L. 6 mm. (II. 01) (=Stage V.).

(Text-figs. 7 & 8.)

The cavity of the optic vesicle is still further reduced and the

lens is almost separated off from the ectoderm which is closing

over it.

Text-figure 7.

Trichosurus vulpecula. G.L. 6 mm. (II. 701).

Horizontal section through the head (S 2-4-2), passing just above the middle of the

head-cavity of the right side (7.h.c.), and showing the proliferating bud on the

dorso-lateral wall (d./.) and the thickening on the postero-medial wall (p.m.).

The anterior portion of the abducens muscle-mass (v.e.) lies posterior to the

head-cavity and in front of the vena capitis medialis (v.e.m.). > 110 and

reduced by 3.

@.¢e.¢.=arteria cerebri anterior. 27B.=fore-brain. hp.=hypophysis.

op.v.=optic vesicle.

OF THE EYE MUSCLES IN MARSUPIALS. ail:

The premandibular head-cavity is large and conspicuous on

each side; although at its widest it is only slightly smaller than

that of the 7 mm. embryo, the ventral portion is reduced,

perhaps owing to further proliferation of the walls.

The proliferation of the postero-medial wall is more extensive ;

it begins more dorsally and, as the cavity narrows ventrally, it

extends round on to the postero-lateral wall, the two thickenings

forming one mass (text-fig. 8, pm.+pl.). The thickened area

is continued on to the extreme ventral wall from which a solid

mesodermal mass, the m. obliquus inferior, hangs down below

the cavity posterior to the optic cup and tapers at its lower

end.

Text-figure 8.

ee ay :

: & 8D 9 bie ‘“ ha

a a; a0 5 ed f

SEES: s oe Ba ze & ’ Go \ Eo

5 a or 4s wt

PR BAG Row

a woop Ff

VW. O.f- é pm.+pl.

Trichosurus vulpecula. G.L. 6 mm. (II.’01).

Horizontal section through the head (S 2-4-8), passing through the dorsal region

of the optic vesicle (op.v.) and the ventral portion of the left head-cavity (/./.c.),

to show the now united postero-medial and postero-lateral proliferations from

the wall (pm.+pl.). 110 and reduced by +.

a.c.a.=arteria cerebri anterior. #'B.=fore-brain. hp.=hypophysis.

v.0.1.=vena orbitalis inferior.

At this stage for the first time, on the dorso-lateral region of

the cavity on both sides, the wall shows a distinct, though not

very extensive thickening which originates as a bud growing out

of the cavity (text-fig. 7, d./.). This bud is the primordium of

the m. rectus superior.

The cavity is lined by a single layer of cells except where the

wall is proliferating.

The abducens muscle complex has increased considerably in

size and is more distinct than in our earlier stages. The much

larger anterior portion, still connected on its lateral side with

Sl MISS E. A. FRASER ON THE DEVELOPMENT

the intermediate mass, lies very close behind the head-cavity and

in front of the Gasserian ganglion, being separated from the

latter by the vena orbitalis inferior (text-tig. 7, 7.e.); it extends

down along the upper two-thirds of the cavity to the region

where the postero-lateral proliferation begins. The smaller

postero-dorsal portion runs back as before on the medial side of

the vena capitis medialis.

The primordium of m. obliquus superior has now grown further

forwards from the intermediate mass and reaches anteriorly

beyond the distal end of the ophthalmic branch of the trigeminal

nerve, on the outer side of which it lies. The connection of the

intermediate mass with the maxillo-mandibular mesenchyme is

most probably present but could not definitely be made out in

this embryo; the mass is again joined with the dorso-lateral

corner of the cavity for about three sections.

The oculomotor nerve is stronger and, running down ventrally

on the medial side of the vena capitis medialis, now reaches the

head-cavity and appears as a small nerve between the wall of the

latter and the anterior portion of the abducens muscle-mass ; its

distal end soon disappears and can only be followed for a short

distance alongside the cavity.

The abducens nerve is now well developed and runs forwards

on the inner side of the Gasserian ganglion, but it is not very

much longer than at the last stage.

Embryo of G.L. 7°75 mm. (XX. ’04) (=Stage VI.).

As regards the development of the eye and the muscle-masses

this stage is very similar to the last, and only a few observations

need be made upon it.

The optic vesicle is not so flattened as at 6 mm. and the lens

is still joined with the ectoderm.

The premandibular head-cavity is smaller and pear-shaped, its

wider side lying next the eye, and the posterior proliferations

are more extensive though the thickening on the ventral wall

does not appear to be so marked. No dorso-lateral proliferation

was recognised.

The abducens nerve now runs into the pointed posterior end

of the dorsal portion of the abducens muscle-mass.

Embryo of G.L. 7:25 mm. (=Stages VII. & VIIl.a & 6).

(Text-figs. 9-13.)

In this next stage there are three embryos, a (III. ’01),

6 (XIT a. 02) and c(XII. ’02), each measuring 7:25 mm. at its

greatest length. Although in many particulars (a) is the youngest

of the three and (c) is slightly older than (6), in respect of the

eye-muscle primordia very little difference exists between them.

The invagination of the optic vesicle has advanced still further,

the cavity being reduced to a narrow slit, but the optic stalk still

contains a wide lumen. ‘The lens is closed off from the ectoderm

OF THE EYE MUSCLES IN MARSUPIALS. ole

and forms a hollow vesicle with thickened walls and a central

cavity.

In (a) the premandibular head-cavity has attained its maximum

size and, at its widest, has roughly the form in horizontal

section of a right-angled triangle (text-fig. 9, /./.c.) measuring

-48 mm. transversely x °37 mm. antero-posteriorly x *23 mm. dorso-

ventrally. In (6) and (c) the cavity is smaller than in (@), but is

still large, the difference being due mainly to a reduction in the

antero-posterior width, which here is only -12 mm.

Text-figure 9.

sg Wapasegh

ned \eerezhe

5 OP, : oe

m Meee

Trichosurus vulpecula. Gl. 7°25 mm. (IIT. ’01).

Horizontal section through the head (S 4-4-1), passing through the dorsal region

of the optic vesicle (op.v.) and through the middle of the left head-cavity (Z.h.c.).

The ventral portion of the proliferating bud on the dorso-lateral wall (d/.) and

the thickening on the postero-medial wall (pm.) are shown, and also the

anterior portion (r.e.) of the abducsns muscle-mass lying posterior to the

cavity. Between the latter and abducens muscle-mass runs the oculomotor

nerve (III.). > 110 and reduced by +.

a.c.a.=arteria cerebri anterior. 'B.=fore-brain. hp.=hypophysis. g.V.=Gas-

serian ganglion. .c.V. = Naso-ciliary. branch of the trigeminal nerve.

v.c.m.=Vena capitis medialis.

The thickening of the posterior wall is still more marked than

in the preceding stage and, joining with the prominent ventral

proliferation which now forms a conspicuous outgrowth (text-

figs. 10, 12, & 13, o.2mf.), extends out ventro-laterally to taper off

behind and somewhat below the eye. ‘This portion, already

314 MISS E. A. FRASER ON THE DEVELOPMENT

indicated in the 7 mm. and 6 mm. embryos, develops into the

future m. obliquus inferior.

_ On the medial side of the primordium of the m. obliquus

inferior at the lower end of the cavity is a darker patch of cells

(text-fig. 10, r.2nf.), the first indication of the m. rectus inferior,

which therefore apparently arises from the lateral side of the

postero-ventral wall just medial to the m. obliquus inferior.

Text-figure 10.

. :

Cee)

a? BEOS SOF

$ ES eee ge ce

og Pie © Bn 8. o, 3

oS PEG Se Pe at, 2°

"ee" 3

v.c-m. r.inf. Lhe.

Trichosurus vulpecula. G.L. 7°25 mm. (III. 01).

Horizontal section through the head (S 5-1-2), passing through the optic cup (op.c.)

above the optic stalk and through the extreme ventral end of the lett cavity

(L.h.c.). to show the primordium of the m. obliquus inferior (0.inf.) and, just

medial to the latter, the region from which the m. rectus inferior (7.inf-) takes

its origin. X 110 and reduced by 3.

a.c.a.=arteria cerebri anterior. 'B.=fore-brain. hp.=hypophysis.

v.c.m.=vena capitis medialis.

The muscle-bud from the dorso-lateral wall, representing the

primordium of the m. rectus superior, is less conspicuous than

the other proliferations. It is present on both sides, projecting

behind the dorsal part of the optic cup (text-fig. 9, d/.); on the

cht side in (c) the bud is less distinct, probably owing to the

further thickening of its posterior wall and to the commencing

separation of the dorso-lateral portion of the cavity from the

rest.

A distinct epithelial layer can only be distinguished on the

side next the fore-brain.

The abducens muscle-mass still consists of two portions. The

dorsal portion (text-fig. 11, r.b.) stretches still further back

on the medial side of the vena capitis medialis as a narrow

OF THE KYE MUSCLES IN MARSUPIALS. 315

prolongation into which runs nerve VI and becomes continuous

with the anterior part. The latter turns round almost at right

angles and runs out laterally immediately behind the postero-

lateral wall of the head-cavity (text-figs. 9 & 12, 2.¢.). It tapers

ventrally, its lower outer end lying close against the inner

portion of the m. obliquus inferior, the two being connected

through two or three sections.

Text-figure 11.

= tod a nS.

: oes 2 a Ay

ae OS te o Sag 9 B®

9 o a Ck ae De & g?

6) : ° ef oe 0%

Peg Pingtro eet Sh

Set — ¢ oe, Bes

Trichosurus vulpecula. GL. 7:25 mm. (XII a. 02).

Longitudinal section (S 9-3-9), passing through the medial side of the right head-

cavity (v.h.e.) to show the proliferation from the posterior wall (pm.+ pl.)

and the postero-dorsal portion (7.b.) of the abducens muscle-mass. X 110 and

reduced by [.

a.c.a.=arteria cerebri anterior. 2#'B.=fore-brain. a.c.i.=arteria carotis interna.

HB.=hind-brain. IL.=oculomotor nerve.

The trigeminal nerve has now both a supra-orbital and a

naso-ciliary branch, running forwards above the bulbus, the

nago-ciliary lying on the outer side of the dorsal end of the

cavity (text-figs. 9 & 13, 2.c.V.).

The primordium of the m. obliquus superior is best seen in the

longitudinal series (text-fig. 13, 0.s.). It begins as a collection

of cells above and anterior to the optic cup and, extending along

the ventral and ventro-lateral sides of the supra-orbital nerve,

Proc. Zoou, Soc.—1915, No. XXIII. 23

316 MISS I. A. FRASER ON THE DEVELOPMENT

it passes backwards and downwards as a solid band of mesen-

chyme through which runs the naso-ciliary branch of the tri-

geminal just where this branch joins the ophthalmic. This band

continues ventrally, anterior to the Gasserian ganglion, to join

the intermediate mass, the latter being connected posteriorly by

a thin stream of cells round the antero-ventral side of the same

ganglion with the maxillo-mandibular muscle-mass. No union

with the maxillo-mandibular mesenchyme can be traced in either

Text-figure 12.

CoRR

RRA OBE

yt PARQ YS

Lig fe,

Liz g &

gig ig

» “ee

ist 2

Wg Be

Trichosurus vulpecula. G.L. 7°25 mm, (XIT a. ’02).

Longitudinal section (S 10-3-1), passing about half-way through the right head-

cavity (2.c.), to show the primordium of the m. obliquus inferior (o.inf.) and

the anterior portion (7.e.) of the abducens muscle-mass lying behind the cavity

and immediately in front of the vena capitis medialis (w.c.m.) and the vena

cerebralis anterior (v.c.a.). The oculomotor (III.) runs down between the

cavity and the abducens mass. XX 110. ;

a.¢c.a,=arteria cerebri anterior. op.c.=optic cup. v.0.7.= vena orbitalis

interior.

of the transverse series, but the intermediate mass in all three

embryos is still jomed with the dorsal outer end of the anterior

portion of the abducens complex. In the transverse series (c)

the intermediate mass appears to have lost its connection with

the m. obliquus superior and is apparently degenerating.

The oculomotor nerve, descending almost directly ventrally

OF THE EYE MUSCLES IN MARSUPIALS. Si

from the mid-brain, runs close to the postero-lateral wall of the

cavity between the latter and the abducens mass (toxt-figs. 9 & 12,

TIT.). At its distal end it breaks up into many fibres which

extend into the postero-lateral proliferation near the point of

origin of the m. obliquus inferior.

Text-figure 135.

Trichosurus vulpecula, (XII A. 02.)

Longitudinal section (S 11-1-4), passing through the lateral wall of the cavity and

the primordium of the m. obliquus inferior (o.inf.). ‘Lhe primordium of the

m. obliquus superior (0.s.) extends forwards from the intermediate mass

(iné.m.) as a solid band of cells dorsal to the optic cup (op.c.). x 110.

g-V.=Gasserian ganglion. maa.md.=maxillo-mandibular mesenchyme. 2.c. V.=

naso-ciliary branch of the trigeminal nerve. v.0.7.=vena orbitalis inferior.

Embryo of G.L. 8°5 mm. (97 & LV.’01) (=Stage IX.a & b).

(Text-figs. 14-16.)

This stage, which measures 8°5 mm. and of which we have

two embryos, (@) and (6), is a good deal more advanced than the

preceding one.

The primary optic vesicle still shows a well-marked cavity in

the region of attachment of the optic stalk which is still hollow,

and pigment is beginning to form in its outer wall. The inner

wall of the lens is thickened, its cavity being reduced to a narrow

23"

318 MISS E. A. FRASER ON THE DEVELOPMENT

semicircle, and the eyelid has begun to develop as a fold of the

ectoderm on the ventral and posterior sides of the optic cup.

The premandibular cavity is now represented by a.mass of

mesoderm in which are situated a few cleft-like spaces lying

obliquely on each side of the fore-brain (text-fig. 16, /./.c.). On

this side the cells have lost almost all traces of their former

epithelial character and the spaces are bounded by a thin

degenerating layer only a few cells thick; the remaining meso-

dermal mass, except those parts which are giving rise to the

eye muscles, is also breaking down.

Text-figure 14.

Trichosurus vulpecula. Gl. 85 mm. (797).

Longitudinal section (S 7-1-8), showing the ciliary ganglion (eél.g.), the anterior

portion (7.e.) of the abducens muscle-mass and the primordium-of the m. rectus

superior (7.s.). > 110.

t.a.=arteria cerebri anterior. g.V.=Gasserian ganglion. op.c.=optic cup.

v.c.a.=vena cerebralis anterior. v.e.m.=vena capitis medialis. v.0.i.=vena

orbitalis inferior. III.=oculomotor nerve.

The dorsc-lateral proliferation is now well marked and its

cells are undergoing distinct differentiation to form. the future

m. rectus superior. It has moved dorsally and anteriorly as

compared with the last stage, and now lies medial to the dorsal

region of the bulbus (text-figs. 14-16, 7.s.) and is showing signs

of becoming independent.

OF THE EYE MUSCLES IN MARSUPIALS. 319

Further ventrally the cavities have completely disappeared and

from the now solid mass of cells the m. obliquus inferior runs

outwards and downwards behind the eye (text-fig. 15, 0.inf.).

Immediately medial to the m. obliquus inferior the differentiation

of cells destined to become the m. rectus inferior extends down-

wards, its ventral end turning anteriorly, medial to the bulbus

and below the optic stalk.

Text-figure 15.

Trichosurus vulpecula. G.L. 85 mm. (97).

Longitudinal section (S 3-2-8), showing the m. obliquus superior (0.s.) with its

narrow posterior prolongation and the m. obliquus inferior (0.inf.) extending

below the optic cup (op.c.). The distal end of the m. rectus superior (7.s.) and

the lateral side of the anterior portion (7.e.) of the abducens muscle-mass are

also seen. X 110.

g-V.=Gasserian ganglion. .c.V.=naso-ciliary branch of V._ s.o.V.=supra-

orbital branch of V. ma. V.=maxillary branch of V.

The primordium of the m. obliquus superior is now established

as an independent structure. Its main portion forms a con-

densed mass of cells lying anterior and dorsal to the eye, and

from this a narrow prolongation projects posteriorly above the

bulbus, the higher pointed end of which comes very close to the

m. rectus superior (text-fig. 15, 0.s.). This backward prolongation

is obviously the last remnant of the former connection with the

intermediate mass (compare Johnson, °13, p. 150). The last

320 MISS E. A. FRASER ON THE DEVELOPMENT

vestiges of the latter can only be recognised in the longitudinal

series in front of the Gasserian ganglion and just posterior to

the ventral side of the anterior portion of the abducens muscle-

mass, with which it is no longer connected; it passes back on

the inner side of the root of the maxillary branch of the tri-

geminal nerve to join with the maxillo-mandibular muscle-mass.

The abducens muscle-mass has not changed in position, except

that the posterior dorsal portion does not seem to extend so far

backwards, and that the anterior portion has grown out still

further laterally (text-figs. 14-16, r.e.).

Text-figure 16.

: s

Lae

Cais

Trichosurus vulpecula. G.l. 85 mm. (IV. 01).

Horizontal section through the head (S 7-1-4), showing the last vestiges of the left

head-cavity (J.4.c.) and the developing m. rectus superior (7.s.). The anterior

portion (7.e.) of the abducens muscle-mass lies in front of the vena capitis

medialis (v.c.m.) and the Gasserian ganglion (g.V.), and in it lies the abducens

nerve (VL.). The ventral edge of the m. obliquus superior (0.s.) is just seen

anterior to the optic cup (op.c.). X 110 and reduced by. 4.

a.c.i.=arteria cerebri anterior. I’ B.=fore-brain. .c.V¥.=naso-ciliary branch of

the trigeminal nerve. III.=oculomotor nerve.

The oculomotor nerve (text-figs. 14 & 16, III.) again divides

at its distal end close to the point of origin of both m. obliquus

inferior and m. rectus inferior into many fine fibres, some of which

penetrate into the m. obliquus inferior itself (see text-fig. 17,

ING)

OF THE EYE MUSCLES IN MARSUPIALS. ayalh

The ciliary ganglion first appears at this stage as a trans-

versely elongated mass of cells, situated between the ventral end

of the anterior portion of the abducens complex and the point of

origin of the m. obliquus inferior and the m. rectus inferior

(text-fig. 14, cil.g.).

The trochlear nerve was observed for the first time in the

longitudinal series but it could not be made out in the horizontal.

It arises in the usual way from the dorsal side of the brain and

runs ventrally almost as far as the level of the dorsal border of

the Gasserian ganglion.

The first indication of the formation of muscle cells is seen in

series (>) and is most marked in the mm. recti superior, inferior

and externus, less definite in the m. obliquus inferior, and not

recognisable at all in the m. obliquus superior.

Embryos of G.L. 9°5 mm. (V.’01) & 10 mm. (VI.’01)

(=Stages X. & XI.). (Pl. I. figs. 1, 2 & text-fig. 17.)

These two embryos, measuring 9°5 and 10 mm. respectively,

resemble in many respects those of the last stage but show in

some details a slight advance in development.

The outline of the eye is more oval and the eyelid has closed

over at the extreme ventral side of it. At 10 mm. the cavity of

the lens is further reduced and the opening in the optic stalk is

narrower.

A slit-like remnant of the head-cavity is still present in both

embryos (text-fig. 17, /.4.c.). Figs. 1 & 2 (Pl. I.) show posterior

and anterior views of a wax-plate model of the embryo of

9°5 mm. The m. rectus superior is now quite conspicuous and

has grown forwards and dorsally above the optic cup (figs. 1 & 2,

r.s.). Below the latter the m. obliquus inferior extends down

from the postero-lateral corner of the solid mass of mesoderm

which has taken the place of the former head-cavity (fig. 1,

o.inf.). The m. rectus inferior is now more marked and stretches

out anteriorly below the optic stalk (fig. 2, r.inf.). The m.

obliquus superior (fig. 2, 0.s.) lies in the same position as in the

last stage; but is larger and more prominent and still retains

the narrow prolongation running backwards towards the m. rectus

superior (fig. 1, 7.s.). The two portions of the abducens complex

are well seen in the model (PI. I. fig. 1 and text-fig. 17, 7.6. &

7.e.) and are very similar to the embryo of 8°5 mm.

At 10 mm. the differentiation of muscle fibres is well ee

lished and can be seen in all the developing eye muscles.

In both embryos, at the point of origin of the m. rectus

inferior and m. obliquus inferior, a narrow band of mesodermal

cells runs directly outwards immediately anterior to the m.

obliquus inferior and becomes united with the ventral side of

the eyelid posterior to the optic cup (PI. I. figs. 1 & 2, +).

Be) MISS E. A. FRASER ON THE DEVELOPMENT

Text-figure 17.

Trichosurus vulpecula. G.L. 95 mm. (V.’01).

Horizontal section through the head (S 4-3-5), passing through the optic cup (op.c.),

optic stalk (op.s.) and lens (/.), and showing the last vestige of tle left head-

cavity (/.h.c.) and the proximal end of the m. obliquus inferior (0.inf.), into

which fibres of the oculomotor nerve (III.) penetrate. The two portions (7.b.

& r.e.) of the abducens muscle-mass are well seen and are penetrated by the

abducens nerve (VI.). X 110 and reduced by ¢.

a.c.a.==arteria cerebri anterior. #'B.=fore-brain. g.V.=Gasserian ganglion.

n.c.V.=naso-ciliary branch of the trigeminal nerve. v.¢.m.=vena capitis

medialis.

Embryos of G.L. 11 mm., 11°5 mm., & 12 mm. (=Stages XII.

& XIII.). (Vext-fig. 18.)

The observations were made from six embryos, one of 11 mm.

(XII. ’04), four of 11:5 mm. (XXIT. 04, 5797, 9’98, & VIT.’01),

and one embryo of 12 mm. (X.’01).

The lumen of the optic stalk is very small and the cavity of

the lens is considerably reduced compared with the 10 mm. stage.

A thick strand of cells running postero-medially towards the

hypophysis on each side of the fore-brain is all that remains of

the unused walls of the former head-cavity. With the lateral

end of this strand is connected the m. rectus superior which,

OF THE EYE MUSCLES IN MARSUPIALS. 323

running dorsally outwards, ends above the eye just behind the

middle of the latter (text-fig. 18, 7.s.).

The m. rectus inferior (text-fig. 18, 7.inf.) arises below the

proximal end of the m. rectus superior with which it is joined

by a slender connection ; it has increased in thickness and passes

out ventrally and anteriorly below the optic stalk. Behind the

middle region of this muscle, towards its proximal end, a small

new offshoot branches out and stretches up dorsally on to the

anterior side of the optic stalk; this is the first appearance of

the m. rectus internus (text-fig. 18, 7.7.).

Text-figure 18.

SEs FE Le .

rY 2

oS e jo 88 ser ae

& ES be <— SO oe

Lr 3 2

& cc et dod 3

SN ved 3 _€e

ae oe 8g

*: ca

* y ate Ss

Ooogs Fg ops Bras ig YP S2e

es seas

Trichosurus vulpecula. G.L. 115 mm. (5797).

Longitudinal section (S 11-3-5), showing the primordium of the m. rectus internus

(7.i.) which develops as an outgrowth from the m. rectus inferior (7.inf.).

The m. rectus externus (7.e.), m. rectus superior (7.s.), and m. obliquus superior

(o.s.) are also seen. X 110.

g.V.=Gasserian ganglion. .c.V.=naso-ciliary branch of trigeminal nerve.

op.s.=optic stalk. III.=oculomotor nerve. WI.=abducens nerve.

The m. obliquus inferior has changed most in position compared

with the last stage. It has moved forwards below the bulbus

and now runs antero-ventrally outwards, lying some distance in

front of the m. obliquus superior above. It is loosely joined to

the outer ventral end of the m. rectus inferior, though this

connection is nothing more than a few strands in the 11:5 mm.

embryo, and its distal end has begun to turn slightly forwards

towards the ali-nasal cartilage from which it finally takes its

origin.

The m. obliquus superior is now well developed; it has

324 MISS E. A. FRASER ON THE DEVELOPMENT

increased in length and runs from the dorsal anterior side of the

bulbus inwards and slightly downwards (text-fig. 18, 0.s.).

The anterior portion of the abducens muscle-mass (text-fig. 18,

7.e.), which we may now call the m. rectus externus, stretches

out laterally in the usual position posterior to the other muscles

and just behind the ciliary ganglion, its pointed lateral end

extending outwards towards the posterior border of the eye.

The inner dorsal portion of the mass, however, is now shorter

and its anterior end, which is united with the m. rectus externus,

has thickened, the thickening being the preliminary stage of the

later forward movement of this portion which is the primordium

of the m. retractor bulbi.

The oculomotor nerve passes ventrally from the mid-brain

anterior to the Gasserian ganglion, gives off a twig into the

proximal end of the m. rectus superior, then goes downwards

through the medial side of the ciliary eanglion immediately

behind the optic stalk. Leaving the ganglion, it gives off a

branch to the m. rectus inferior and runs outwards on the

postero-lateral side of the latter to send an offshoot to the

m. obliquus inferior (text-fig. 18, III.). Thus all the oculomotor

muscles receive their nerve supply at practically the same time.

The ciliary ganglion is slightly larger than at the last stage and

forms a transversely elongated mass of cells lying on a level with

the optic stalk between the ventral end of the m. rectus externus

and the proximal end of the m. rectus inferior.

The trochlear nerve can now be followed from the dorsal

surface of the brain downwards immediately in front of the

Gasserian ganglion, its distal end turning forwards to run into

the m. obliquus superior.

As in the last stage, a strand of mesoderm cells extends

laterally from the proximal end of the m. rectus inferior to the

ventral border of the eyelid, but here, owing to the forward

movement of the m. obliquus inferior below the bulbus, this

strand now lies dorsal and posterior to the latter muscle.

The embryo of 12 mm. only differs in the following respects

from the 11°5 mm. The eyelids have folded completely over the

eye; all the muscles have increased in size and every trace

of the walls of the former head-cavity has disappeared. The

attachment of the distal end of the m. obliquus inferior to

the ali-nasal cartilage has begun and the anterior end of the

primordium of the m. retractor bulbi, only apparent as a

thickening in the last embryo, now projects slightly forwards

towards the optic stalk.

Embryos of G.L. 13 & 13°5 mm. (=Stage XIV.).

The stage consists of three embryos, two measuring 13 mm.

(a) (IX. ’01) & (6) (XXV.), and the third (c) (X XIV.) witha

length of 13°5 mm.

The optic stalk is still hollow, though its cavity is very small

OF THE EYE MUSCLES IN MARSUPIALS. 325

and difficult to see in the mid-region of its extent. In (a) the

eyelids have folded completely over the eye, but in (6) they have

closed over the lower half only and here the lens is solid.

Otherwise the eye resembles that of the last stage.

A vestige of the walls of the former head-cavity is still present

in embryo (a) only, and is seen as a strand of cells running back-

wards and inwards from the medial end of the m. rectus superior.

In (a) the latter is an independent muscle, whereas in (4) and (c)

it is apparently still jomed with the m. rectus inferior.

As the positions of the muscles at this stage are very much

the same as at 15 mm., of which a wax model has been con-

structed, they will be described in detail in the sequel. A few

remarks may, however, be made here. The m. rectus internus

Is as yet very short and at 13 mm. the m. rectus inferior is still

attached to the m. obliquus inferior, but the separation of the

two muscles is complete at 13°5 mm. The m. obliquus inferior

is now definitely attached to the ali-nasal cartilage; it has thus

changed its position, its outer end turning inwards to the point

of attachment, while the proximal end is inserted into the

antero-ventral side of the bulbus.

The part of the abducens muscle complex destined to form

the m. retractor bulbi has grown still further forwards and

outwards towards the medial wall of the eyeball. From its

lateral side the m. rectus externus extends almost directly out-

wards to the posterior lower wall of the bulbus, the two muscles

running out at an acute angle to each other, the m. retractor

bulbi crossing the inner end of the m. rectus externus.

Embryos of G.L. 14, 14:5, & 15 mm. (=Stage XV.).

(Pl. II. figs. 3-5.)

In this stage we have an embryo of 14 mm. (XXIII.), two

pouch young of 14°5 mm., @ (XV. 02) and 6, and one of 15 mm.,

the last three being recently born.

The eyelids are now quite folded over but the optic stalk

still contains a minute cavity.

The m. rectus superior (PI. II. fig. 4, 7.s.) runs from behind

the inner end of the optic stalk just above the proximal end of

the m. rectus inferior slightly dorsally and outwards, its pointed

outer extremity being inserted in the eyeball dorsal to the

posterior side of the latter and at a slightly lower level than the

m. obliquus superior.

The m. obliquus superior (Pl. I. fig. 3, 0.s.) is now a stronger

better developed muscle than at 13 mm. It extends directly

outwards over the anterior side of the bulbus, its broad inserted

end, which is flattened antero-posteriorly, stretching further

laterally over the eyeball than any of the other eye muscles.

The m. rectus inferior (Pl. II. figs. 3 & 5, r.inf) arises from

below the inner end of the optic stalk and passes downwards and

slightly anteriorly below the stalk as a broad muscle which tapers

326 MISS E. A. FRASER ON THE DEVELOPMENT

gradually as it proceeds below the bulbus, its pointed ventral

end being inserted close to the m. obliquus inferior. Near its

wide proximal end, immediately in front of the optic stalk, the

m. rectus internus branches off and runs outwards with a slightly

dorsal trend on to the anterior side of the bulbus close below the

m. obliquus superior (figs. 3 & 5, 7.7.). The m. rectus internus

is as yet a much thinner and shorter muscle than the m. rectus

inferior but has nevertheless grown considerably since our last

stage, now stretching farther laterally and upwards towards its

future point of insertion in the eyeball.

The m. obliquus inferior (Pl. II. figs. 3-5, 0.inf.) is no longer

connected with the m. rectus inferior. It appears as a more or

less dorso-ventrally flattened structure of a somewhat oblong

shape, having a slightly longer axis in an obliquely antero-

posterior direction ; as at 13:5 mm., its now proximal but formally

distal end is attached to the ali-nasal cartilage, the opposite

once proximal end extending towards the lower anterior side of

the bulbus.

As regards the abducens complex, we see at 15 mm. very little

advance compared with the last stage. The portion corresponding

to the m. retractor bulbi (figs. 4 & 5, 7.6.) stretches farther

inwards and posteriorly than any of the other muscles; it

increases in thickness as it runs antero-laterally and downwards

towards the outer end of the optic stalk, its distal end lying just

behind the junction of the oculomotor with the ciliary ganglion.

In the model this end shows the first indications of growth round

the posterior side of the optie stalk, but this growth round the

stalk is, however, more marked in 14mm.and 14:5 mm. The

m. rectus externus (figs. 4 & 5, re.) stretches from the lateral

side of the m. retractor bulbi, not far from the anterior end of

the latter, outwards and downwards, its outer pointed end lying

below the postero-ventral side of the bulbus some distance

behind the outer end of the m. obliquus inferior.

The abducens nerve extends forwards from its origin and then

downwards on the outer side of the m. retractor bulbi, running

into the abducens complex at the region where the two components

of the latter are joined with one another (fig. 4, VI.).

The oculomotor nerve (Pl. II. figs. 3-5, III.) runs forwards

medial to the naso-ciliary branch of the trigeminal into the

posterior end of the m. rectus superior, and continues obliquely

outwards and downwards close against the inner end of the

ciliary ganglion and along the side of the m. rectus inferior next

the eye. It gives off fibres on this same side into the root of the

m. rectus internus. Leaving the m. rectus inferior before the

latter reaches the bulbus, it passes anteriorly and ventrally into

the m. obliquus inferior.

The ciliary ganglion forms an elongated mass of cells stretching

out below the optic stalk ; at 15 mm., as seen in the model (fig. 5,

cil.g.), it is not so elongated as in some of the examples measuring

13 mm. and 14 mm.

OF THE EYE MUSCLES IN MARSUPIALS. SOM

The distal end of the trochlear nerve (figs. 3-5, IV.) runs

forwards and outwards into the posterior side of the medial end

of the m. obliquus superior.

The naso-ciliary branch of the trigeminal nerve (figs. 3-5, V.)

passes out immediately lateral to the oculomotor, downwards

below the posterior side of the m. rectus superior, and forwards

above the optic stalk and along the anterior lower border of the

medial end of the m. obliquus superior to the snout.

Foetus of G.L. 17 and of 17-5 mm.

This stage consists of two pouch young, one measuring 17 mim.

and the other 175mm. The optic stalk has quite disappeared and

the eye muscles have practically attained their adult positions.

The distal end of the m. obliquus superior turns posteriorly

and comes to lie so close to the outer end of the m. rectus superior

that the two muscles are inserted almost at the same place.

The m. rectus internus has increased greatly in size and is

now a well-developed muscle ; it is, however, still conneeted with

the m. rectus inferior at its proximal end by a thin band of cells.

The m. obliquus inferior, as in our last stage, runs obliquely

backwards from the ali-nasal cartilage, its posterior end now

being inserted anterior to and on the lateral side of the m. rectus

externus.

The m. rectus externus and the m. retractor bulbi are now

separate muscles, The former is inserted on the posterior side

of the bulbus; at about its mid-region on the anterior side, an

indentation is present which in all probability represents the

first indication of the later division of the muscle into two portions.

The m. retractor bulbi has developed considerably; it extends

from the middle of the basisphenoid cartilage, to which it is

becoming attached, anteriorly and outwards, its distal end

surrounding the posterior side of the optic nerve in the shape of

a half moon, the pointed ends of which stretch out, the one above,

the other below the nerve. The m. retractor bulbi is innervated

by a small branch from the abducens nerve before the latter runs

into the m. rectus externus.

Fetus of G.L. 5-2 em. (Text-fig. 19.)

The m. retractor bulbi (text-fig. 19, 7.6.) has now attained its

adult position. It enters the orbit on the posterior side of the

optic nerve and, surrounding the latter, stretches outwards as a

circular sheet of muscle gradually diminishing in thickness and

increasing in circumference up to its insertion round the inner

side of the bulbus within the recti muscles. The two edges of

the sheet lie close together, especially at its outer end, but they

never completely fuse.

The proximal end of the m. rectus externus (text-fig. 19, r.é.)

is split up into two portions, already indicated at 17 mm., which

328 MISS E. A. FRASER ON THE DEVELOPMENT

unite distally to have a single insertion on the posterior side of

the bulbus.

Text-figure 19.

re.

xm MeL

rl.

Trichosurus vulpecula. G.L. 5°2 cm.

Longitudinal section (S23-1-5) passing through the optic nerve (II.) and the

surrounding eye muscles. The m. retractor bulbi (7.b.) forms a circular sheet

of muscle within the other eye muscles. The two portions of the m. rectus

externus (7.e.) are also seen. X 35.

l.p.s.=levator palpebre superioris. 0.s.=m. obliquus superior. VI.=abducens

nerve. 7.7.=m. rectus internus. 7.inf.=m. rectus inferior. 7.s.=m. rectus

superior,

Summary of Hvents in Trichosurus vulpecula.

In early stages a large premandibular cavity surrounded by

a single epithelial layer of cells is present on each side of the

fore-brain posterior to the optic vesicles.

In the 7 mm. embryo an extensive thickening occurs on the

lower two-thirds of the posterior wall, later (in the 6 min.

embryo) extending on to the ventral wall of the cavity. From

this thickening the m. obliquus inferior and the common

primordium of the m. rectus inferior and m. rectus internus

develop, the first to appear being the m. cbliquus inferior which

grows out laterally as a solid protuberance from the ventro-

lateral side; the m. rectus inferior arises a little later (in the

embryo of 8°5 mm.) as a forward extension from the thickened

wall immediately medial and slightly ventral to the m. obliquus

inferior,

OF THE EYE MUSCLES IN MARSUPIALS., 329

As development proceeds, the m. obliquus inferior moves

forwards below the eye, its distal end turning inwards in the

11-5 mm. embryo to become attached to the ali-nasal cartilage.

It becomes independent in the 13°5 mm. embryo and its origin-

ally proximal end moves outwards to reach its final insertion on

the antero-ventral side of the bulbus. This muscle thus under-

goes a noteworthy change in position. The m. rectus inferior

grows forwards and outwards, becomes independent and finally

extends downwards from below the inner end of the optic cup to

become inserted in the ventral side of the bulbus close to the

m. obliquus inferior.

From the dorso-lateral region of the cavity in the 6 mm.

embryo appears a bud-like outgrowth whose surrounding walls,

especially that of the posterior side, proliferate and give rise to

the m. rectus superior. The primordium of this omicele grows

antero-posteriorly over the eyeball, becomes independent and

very early attains its final position (11 to 12 mm.).

The remaining walls of the head-cavity undergo degeneration

and become obliterated in the surrounding mesenchyme.

The m. rectus internus is the last of the eye muscles to make

its appearance. It first develops in the 11 mm. embryo as an

outgrowth from near the proximal end of the m. rectus inferior

and grows dorso-laterally to become inserted on the anterior side

of the bulbus.

The abducens complex is first seen as a solid mass of cells

lying posterior to the premandibular cavity and consisting of

postero-dorsal and antero-ventral portions. The latter grows

out laterally as the m. rectus externus and the former grows

forwards and outwards, crossing the inner end of the m. rectus

externus, its distal end finally surrounding the optic nerve as the

m. retractor bulbi. The m. retractor bulbi is the last of the eye-

muscles to assume its final position.

The m. obliquus superior takes its origin from a mass of cells,

termed by me the intermediate mass, which can be traced down-

wards into continuity with the maxillo-mandibular mass of

mesenchyme. The anterior band-like part of the intermediate

mass separates off to form the primordium of the m. obliquus

superior. It grows inwards to form a muscular band, which

extends out from above the m. rectus internus to the anterior

side of the bulbus.

The more posterior portion of the intermediate mass, attached

to the maxillo-mandibular mesenchyme, is wider and composed

of more loosely arranged cells, and is also connected in our earlier

stages with the inven side of the primordium of the m. rectus

externus and for a. still shorter period with the postero-lateral

wall of the head-cavity. After the m. rectus externus and the

m. obliquus superior become independent the intermediate mass

disappears.

330 MISS E. A. FRASER ON THE DEVELOPMENT

Nores ON OTHER MARSUPIALS.

Phascolarctos cinereus. Embryo of G.L. 4 mm.

Our earliest stage of Phascolarctos cinereus measures 4 mm.

at its greatest length. The outer wall of the optic vesicle is

thickened and flattened and the optic stalk is in wide communi-

cation with the fore-brain.

Situated postero-medially to the optic vesicle on each side is a

large premandibular head-cavity, the long axis of which lies in

the sections in an obliquely antero-posterior direction. From

the posterior and postero-lateral walls active proliferation 1s

taking place by means of hollow buds which are growing out from

the cavity, a solid mass of cells being formed by the thickened

walls of one bud running into those of another. Next the fore-

brain the walls consist of a single epithelial layer, which in some

places, however, is not distinctly seen.

Unfortunately, this embryo is not well preserved and the

abducens muscle-mass is very difficult to distinguish from the

thickened posterior wall of the head-cavity.

Embryo of G.L. 7:5 mm. (Text-fig. 20.)

An older embryo, measuring 7°5 mm., shows the optic vesicle

much further invaginated and the primordium of the lens in the

form of a thickened hollow ingrowth from the ectoderm:

The head-cavity has increased considerably in size, having a

maximum diameter of *30 mm. x °26 mm. (text-fig. 20, 7./.c.), and

its antero-ventral portion has grown forwards above the optic

vesicle. The whole of the posterior wall is thickened; hollow

buds, many of which are long and tubular, run out into a mass

of closely packed cells, the entire region appearing as an irregular

mass pitted with small hollow spaces (text-fig. 20, p.). The wall

next the fore-brain, as in our first stage, is composed of a single

layer of cells.

The abducens muscle-mass, which is clearly recognisable, con-

sists of a narrow postero- dorsal portion lying along the medial

side of the vena capitis medialis; this continues into the larger

anterior part, which extends obliquely forwards and downwards

postero-laterally to the head-cavity to meet the intermediate

mass. The latter stretches forwards just above the optic vesicle

into a bulbous extension, the primordium of the m. obliquus

superior, and runs downwards into the maxillo-mandibular

mesenchyme; it is also connected with the postero-lateral wall

of the head-cavity.

The oculomotor nerve is well developed, its distal end termi-

nating some distance from the head-cavity.

THE EYE MUSCLES IN MARSUPIALS. 33

Text-figure 20.

Phascolarctos cinereus. G.L. 775 mm.

Horizontal section through the head (S 5-3-7), passing immediately above the

optic vesicle and just below the imiddle of the right head-cavity (7.h.c.) ‘The

posterior wall of the cavity is actively proliferating (p.). Size of cavity

*30 mm. X°26 mm. 110 and reduced by <.

a.c.a.=arteria cerebri anterior. £B.=Fore-brain. hp.=hypophysis.

Embryo of G.L.9 mm. (Text-fig. 21.)

At 9 mmm. the optic vesicle is closed except on the medial side

where the hollow optic stalk runs inwards to the fore-brain. The

lens has now the torm of a vesicle and is no longer connected

with the ectoderm.

The head-cavity is still larger (text-fig. 21, U.h.c.) and resembles

in shape that of 7richosurus, embryo (@) 725mm. A large bud

is present on the dorso-lateral side, the posterior and ventral walls

of which are thickened, whilst the whole of the postero-lateral

wall of the cavity is actively proliferating.

The primordium of the m. obliquus inferior runs out from the

postero-lateral side of the cavity behind the optic vesicle, and

medial to this muscle there is an indication of the future m. rectus

inferior.

The abducens muscle complex is well marked in this embryo,

and its connection with the intermediate mass and the m. obliquus

superior is clearer than in any of the embryos of Tvichosurus

examined (text-fig. 21). The abducens nerve runs into the

postero-dorsal portion which is situated in its usual position

medial to the vena capitis medialis; the anterior portion (text-

fig. 21, 7.e.) lies along the postero-lateral side of the cavity in

Proc. Zoou. Soc.-1915, No. XXIV. 24

332 MISS BE. A. FRASER ON THE DEVELOPMENT

front of the vena capitis medialis and stretches laterally into the

intermediate mass (text-fig. 21, iné.m.). The latter extends, on

the one hand, forwards and upwards to be connected by a narrow

band of cells with the m. obliquus superior (text-fiz. 21, 0.s.)

and on the other, downwards and backwards to join the maxillo-

mandibular mesenchyme.

Text-figure 21.

Phascolarctos cinereus. G.L.9 mm.

Horizontal section through the head (S 4-2-6) passing through the left head-

cavity (U.h.c.), about a quarter of the way down, and showing the m. obliquus

superior (0.s.) growing forwards from the intermediate mass (int.m.), this bemg

also united with the abducens muscle-mass (7.e.). In the next few sections

further ventrally, the intermediate mass is joined with the postero-lateral wall

of the head-cavity at approximately the region marked *. 110 and reduced

by Be

a.ca.=arteria cerebri anterior. #'B.=fore-brain. g.V.=Gasserian ganglion.

hp. =hypophysis. _2.c.V. = naso-ciliary branch of the trigeminal nerve.

v.c a.=vena cerebralis anterior. v.c.m.=vena capitis medialis. III.=oculo-

motor nerve.

The m. obliquus superior is a compact mass of cells lying

antero-laterally to the cavity above the optic cup. The inter-

mediate mass a few sections below the level of text-fig. 21 is

attached to the extreme postero-lateral wall of the cavity.

The oculomotor nerve runs down close to the postero-lateval

wall of the premandibular cavity (text-fig. 21, IIT.) and below the

level of the same splits up into many fibres near the point of

origin of the m. obliquus inferior.

OF THE EYE MUSCLES IN MARSUPIALS. Pars

Embryo of G.L. 11 mm.

In our next embryo, which measures 11 mm., all traces of the

head-cavity have disappeared, the m. obliquus inferior has begun

to move forwards below the eye and the m. rectus inferior is

further developed. Phascolarctos has now reached a stage in

development intermediate between that of the TZvrichosurus

embryos of 10 and 11 mm.

The later history of the eye muscles is similar to that of

Trichosurus and need not again be described.

PHASCOLOMYS MITCHELLI. (Text-fig. 22.)

The two youngest embryos of Phascolomys mitchelli measure at

their greatest length 9 mm.and 8°5 mm. respectively ; the former

is shghtly the younger of the two.

The optic vesicle is closed and the optic stalk is in wide com-

munication with the fore-brain; the lens is a hollow vesicle shut

off from the ectoderm.

Text-figure 22.

Phascolomys mite velli. G.L. 85 mm.

Longitudinal section (S 2-1-1) passing through the lateral side of the right head-

cavity (r.h.c.) and showing the proliferation from the dorso-lateral wall (d.2.)

and the primordium of the m. obliquus interior (0.inf.). The intermediate mass

(int.m.) situated between the cavity and the Gasserian ganglion (g./”.) is seen to

be connected with the maxillo-mandibular mesenchyime (maw.md.) by a band

of cells lying just on the medial side of the vena orbitalis inferior (v.0.1.).

110 and reduced by ¢.

a.c.a.=arteria cerebri anterior. op.c.=optic cup. v.c.a.=vena cerebralis anterior.

v.c.m. =vena capitis medialis. v.0.7.=vena orbitalis inferior. ILI.=oculomotor

nerve. es

24"

oor MISS E. A. FRASER ON DEVELOPMENT

In the 9 mm. embryo the head-cavity is large and triangular |

in shape, resembling that of the 9 mm. stage of Phascolarctos,

while in the 8°5 mm. specimen it is much reduced in size (text-

fig. 22, r.h.c.). In both embryos a dorso-lateral proliferating

muscle-bud is present (text-fig. 22, d./.), and the m. obliquus

inferior extends down from the ventro-lateral wall posterior and

ventral to the optic cup (text-fig. 22, o.inf.).

The anterior portion of the abducens muscle complex is joined

with the intermediate mass, and the connection of the latter with

the maxillo-mandibular mesenchyme by means of a wide band of

cells is well seen in the 8°5 mm. embryo (longitudinal series)

(text-fig. 22).

The m. obliquus superior has the same appearance as in the

9 min. stage of Phascolarctos; on its inner border it practically

surrounds the supra-orbital branch of the trigeminal nerve.

The oculomotor nerve is prolonged ventrally into the m. obliquus

inferior and the abducens nerve runs into the hinder end of the

postero-dorsal portion of the abducens muscle complex.

The examination of the two older stages, measuring 15°5 mm.

and 17-5 mm. respectively, shows that the further development of

the eye muscles proceeds as in Trichoswrus.

Macropus RUFICOLLIS, (Text-figs. 23 & 24.)

Only one embryo of Macropus rujicollis, with a maximum length

of 67 mm., came under observation. The optic vesicle is

Text: figure 23.

Macropus ruficollis. G.L. 67 mm.

Horizontal section through the head (S 4-2-1) showing the head-cavity on

each side (l.h.c. & v.h.c.). Size of cavities: left ="29 mm.X°14 mm.; right

= 30 mm.X'16 mm. X40 and reduced by {.

@.¢.a.=arteria cerebri anterior. £B.=fore-brain. hp.=hypophysis.

OF THE EYE MUSCLES IN MARSUPIALS. 335

flattened, its outer layer being much thickened and bulging into

the cavity, and the wide opening in the optic stalk is connected

with the fore-brain. The ectoderm opposite the vesicle is as yet

only slightly thickened on the ventro-lateral side of the latter.

This embryo possesses a large premandibular head-cavity on

each side of the fore-brain in the usual position medial and

posterior to the optic vesicle (text-figs. 23 & 24, Lh.c. & rh.c.).

The dorsal portion of the cavity is somewhat irregular in shape

with a small area marked off towards the median plane by a

Text-figure 24.

Macropus ruficollis, G.L. 67 mm.

Left head-cavity (/.2.c.) showing its lining of a single layer of epithelial cells.

Size of cavity="29 mm.X‘14 mm. 200.

constriction as in Tvrichosurus, embryo (a), 5 mm.; further

ventrally, it assumes a more oval form (text-figs. 23 & 24, Lh.c. &

r.h.c.), measuring on the left side -29 mm. x-°29 mm. x-14 mm.

and on the right *31 mm.x°30 mm.x-‘16 mm., the shortest

measurement being the transverse one. - It is lined throughout

by a single layer of epithelial cells.

The abducens muscle complex, the intermediate mass and

the primordium of the m. obliquus superior are united in the

usual way, but thei respective limits are diflicuit to determine

336 MISS E. A. FRASER ON THE DEVELOPMENT

as their component cells are loosely connected and rather spread

out, so that they are not easily distinguished from the surrounding

mesenchyme. The primordium of the m. obliquus superior is

more compact than the rest and forms a large rounded mass above

the optic vesicle, posterior to and on the outer side of the supra-

orbital branch of the trigeminal nerve.

PERAMELES spp. (Text-fig. 25.)

Our first stage of Perameles is an embryo of P. nasuta

measuring 5°7 mm. at its greatest length. The primary optic

vesicle is flattened and the outer layer thickened, and the

optie stalk contains a wide cavity communicating with the

fore-brain. The adjacent ectoderm is enlarged and _ slightly

invaginated to form the primordium of the lens.

Text-figure 25.

Perameles nasuta. G.L. 5'7 mm.

Horizontal section through the head (S 4-1-11), showing the left head-cavity

(l.h.c.) when at its largest. 110.

a.c.a. = arteria cerebri anterior. 2'B. = fore-brain. hp. = hypophysis.

op.v. = optic cup.

A small very reduced head-cavity is to be seen on each side

posterior and medial to the optic vesicle (text-fig. 25, .h.c.).

It is very irregular in form, is much broken up, and for the

most part is bordered simply by mesenchyme cells, a definite

OF THE EYE MUSCLES IN MARSUPIALS. 337

layer of epithelium having been observed only in the section

figured where the cavity is larger than it is elsewhere. There

appear to be proliferations of cells round parts of the cavity

but no definite muscle-masses can be distinguished.

The m. obliquus superior and the abducens muscle-mass are

connected by means of the intermediate mass and show the

same relations as described for other genera. A portion of

the intermediate mass is apparently joined with a side of the

head-cavity but the limits between the parts are again difficult

to make out.

In the next embryo, which measures 7 mm., the cavity of the

optic vesicle is almost closed and the lens forms a hollow vesicle

but the optic stalk is still widely open. Small head-cavities are

again found, the one on the left side being larger and better

developed than in the 5:7 mm. embryo, while that on the right

is more or less broken up. On both sides a thickening is present

on the postero-lateral side of the head-cavity and the primordium

of the m. obliqnus inferior extends down below the cavity ; on

the left a proliferating dorso-lateral bud is present.

In the 8°75 mm. embryo of P. obesula the cavities have almost

disappeared and may be compared with the traces of the head-

cavity still left in the 8°5 mm. embryo of Zrichosurus. The

muscles are developing as in the latter.

DIDELPHYS MARSUPIALIS. (Text-fig. 26.)

We had at our disposal only one embryo of Didelphys

measuring 8°5 mm. ‘The optic vesicle is completely invaginated

and its cavity obliterated; the optic stalk on the right side

contains a prominent cavity connecting it with the fore-brain

but on the left it is almost solid, only a few very small hollow

spaces remaining, and these are chiefly on the side next the brain.

On the left side the lens has just separated off from the ectoderm

but lies close to the latter and is of a remarkably small size, but

on the right no lens could be seen and had apparently not yet

begun to develop.

No head-eavities are present but in the usual position of the

latter, posterior and dorsal to the eye, lies a condensed mass of

mesenchyme cells from which the eye muscles are developing.

On careful examination, the postero-lateral side of this mass,

which extends obliquely forwards in front of the vena capitis

medialis, is ee without difficulty as the abducens muscle-

mass Ca fig. 26, r.e.); lying parallel to this, on the anterior

side of the whole group of cells, we see the future m. rectus

superior (text-fig. 26, 7.s.), between the medial side of which

and the abducens muscle-mass runs down the oculomotor nerve.

The lateral side of the abducens muscle-mass is joined with the

intermediate mass which, stretching forwards above the eye, is

continued into an outgrowth corresponding to the primordium

of the m. obliquus superior (text-fig. 26, 0.s. and it.m.). The

338 MISS E. A. FRASER ON THE DEVELOPMENT

intermediate mass is also united with the lateral side of the

primordium of the m. rectus superior, thus recalling the con-

nection of the intermediate mass with the postero-lateral wall of

the head-cavity in other marsupials. The central region of the

cell group is compact and as one passes further ventrally, below

the section figured, the primordium of the m. obliquus inferior

is seen to grow out downwards from this region, its approximate

position being marked in the diagram by a broken circle

(text-fig. 26). In still younger embryos it is possible that

vestiges of a cavity may be present, but if so they become

early obliterated.

Text-figure 26.

Didelphys marsupialis. G.L. 8°5 mm.

Horizontal section through the head (S 2-411), showing the mass of mesenchyme

dorsal to the optic cup, from which the eye muscles are developing. ‘he broken

circle indicates approximately the area from which, further ventraliy, the

primordium of m. rectus inferior grows out. > 100 and reduced by 4.

a.c.a. = arteria cerebri anterior. #F'B.=fore-brain. int.m. = intermediate mass-

0.s. =m. obliquus superior. +.e. = anterior portion of abducens muscle-mass.

r.s. =m. rectus superior. v.c.a. = vena cerebralis anterior. v.c.m. = vena

capitis medialis. v.0.i. = vena orbitalis inferior. ILI. = oculomotor nerve.

The conditions in Didelphys are interesting, as they may

possibly form a clue to the more accurate identification of the

group of mesenchyme cells from which the eye muscles arise in

higher mammals. Reuter (’97), as already mentioned, regards

the eye muscles in the pig as originating from a single mass

of mesenchyme cells. In the description of his earliest stage

(G.L. 10 mm.) he says (p. 384):—‘“Sie [die allererste Anlage

der Augenmuskeln| hat die Form einer gestielten Sichel und

umgreift mit den beiden nach vorn gerichteten Schenkeln den

Augenstiel, wiihrend der dritte hintere Schenkel vom N. abdu-

cens fortgesetzt wird. Die Spitze des oberen Schenkels bildet

OF THE EYE MUSCLES IN MARSUPIALS. 339

sich mit ihrem Nerven, dem N. trochlearis, am spiitesten aus.”

By the ‘second stage (G.L. 13 mm.) :—‘ Die Muskulaturanlage

wandert nach vorn gegen den N. opticus hin und verliert ihren

hinteren Schenkel, welcher von der Vena jugularis nach vorn

zusammengedrangt wird.” It will be remembered that in

Trichosurus (G.L. 7 mm.) the root of the vena cerebralis

anterior penetrates through the dorsal side of the larger anterior

portion of the abducens muscle-mass ; this does not occur in the

specimen of Didelphys which has perhaps passed this stage in

development.

DASYURUS VIVERRINUS.

A good series of embryos of Dasyurus viverrinus was examined

and no signs of any head-cavities were observed. The small size

of these embryos, however, makes it exceedingly difticult to

identify any small vestiges of cavities which might ‘possibly

persist.

We may conclude from the preceding facts that the occurrence

of well-developed premandibular head-cavities is characteristic of

the Diprotodontia, large cavities being found in TZ’richosurus,

Phascolarctos, Phascolomys and Macropus. In the Polyproto-

dontia, on the other hand, cavities may be present, as for

example in Perameles, but only in the form of comparatively

small irregular spaces, quite insignificant as compared with those

in the former group, or they are altogether absent, as in Dasyurus

and probably also in Didelphys, the muscles from ‘the first somite

then developing from a solid mesodermal mass as in higher

mammals. o.

Although there are variations in the mode of proliferation

from the walls of the premandibular cavity in the different

genera, the later development of the eye muscles presents much

sunilarity and Z'richosurus may be regarded as typical.

ConcLUDING REMARKS.

From the foregoing observations we have further evidence of

the unique position occupied by the marsupials in the Class

Mammalia. The possession of well-developed premandibular

head-cavities, as yet observed in no other mammals, recalls the

conditions existing in many reptiles. These cavities correspond

very closely with those found in the lizard (Corning, ‘99), in the

snake (Oppel, 90) and in the Chelonia (Filatoff, ‘07 & Johnson,

"13).

tee to lack of material of early stages it has not been

possible to determine the exact mode of origin of the cavities

at the anterior end of the head in marsupials, or even to establish

the existence of a median piece connecting the two across the

middle line. The connecting piece in reptiles very often forms

a wide cross-canal, which may persist until a comparatively late

340 MISS E. A. FRASER ON THE DEVELOPMENT

Stage or may disappear before the cavities have attained their

maximum size. This variation is mentioned by Corning (’00)

(p. 66):—“ Aus den Oppel’schen Figuren, wie aus den meinigen,

geht hervor, wie stark die Variationen sind, welche man in

Bezug auf die Ausbildung des mittleren Verbindungsstiickes

zwischen den beiden Kopfhéhlen antrifft. Nicht selten bleibt

die Verbindung noch in relativ spiter Zeit bestehen, in anderen

Fallen is sie schon zu einer Zeit verschwunden, wo die Héhlen-

bildung im lateralen Theile noch nicht auf ihrem Héhepunkt

angelangt ist, in noch anderen sehen wir im Stiel einzelne

kleinere Hohlenbildungen auftreten, die sich spaiter mehr oder

weniger vollstandig zu einer grossen Héhle vereinigen.”

In the 5 mm. embryo of 7richosuwrus and in the 6:7 mm.

Macropus, the small median portion partially constricted off from

the rest of the cavity (text-fig. 1, 2.) may possibly coincide with

the swollen part of the stalk, which runs from the somite to the

middle line in Anguis fragilis (called by Oppel the “ Hals”) and

which takes a part in the formation of the head-cavity.

We have seen that the walls of the premandibular cavity, as

in most other Vertebrates, give rise to the muscles innervated

by the oculomotor nerve. In Zrichoswrus and probably also in

Phascolomys, the primordium of the m. obliquus inferior and the

common primordium of the mm. rectus inferior and rectus

internus develop as solid outgrowths from the posterior and

ventral walls, whilst the primordium of the m. rectus superior

arises from the walls of a hollow evagination on the dorso-lateral

side, the latter mode of origin resembling that of all the oculo-

motor muscles in Lacerta (Corning, 99). In Phascolarctos,

elongated hollow outgrowths occur along the greater part of the

posterior wall of the cavity, an extensive budding here taking

place. The further development of these three muscles in the

marsupials agrees very closely with that of the same muscles in

Chelydra (Johnson, ’18).

The m. rectus internus and the m. rectus inferior develop from

a common primordium, the m. rectus internus first appearing as

an offshoot from near the proximal end of the m. rectus inferior ;

this offshoot grows directly upwards and outwards to the anterior

side of the bulbus. In Chelydra, according to Johnson (18), the

common primordium becomes transformed into ‘a solid elongate

mass. By the 11 mm. stage a constriction has appeared, slightly

beyond the middle of this mass, differentiating it into a proximal

M. rectus inferior and a distal M. rectus medialis. ......... The

proximal end, which at first is continuous with the distal end of

the M. rectus inferior, works up along the medial side of this

muscle so that the final separation of the two takes place at

their proximal ends, 7. ¢. their ends of origin.” No movement of

this kind has been observed in Jrichosurus, and if it occurs it

must take place in the short interval between the 10 mm. and

11 mm. stages.

OF THE EYE MUSCLES IN MARSUPIALS. 34]

The abducens complex and the m. obliquus superior are

difficult to identify in our earliest stages, but by comparison

with slightly older embryos it is possible to make out their

contours in the surrounding mesenchyme from which they are

not easily distinguished. In our first stages, up to 85 mm.,

the primordia of both these muscles are united with the maxillo-

mandibular mesenchyme by an intermediate mass of more loosely

connected cells, the position of which is well seen in Phascolomys

(text-fig. 21, p. 332). The m. obliquus superior arises from this

intermediate mass as an upgrowth which extends forwards above

the eye, the intermediate mass itself apparently degenerating.

In the rabbit, Edgeworth (03) regards the m. obliquus superior

and the m. rectus externus as ‘‘specialised portions of the man-

dibular and hyoid myotomes which separate, the former late, the

latter very early in development, from the upper ends of their

respective myotomes” (p. 82). In Chelydra, according to Johnson

(713), the m. obliquus superior (p. 159) ‘grows forward as a stream

of cells from the dorsal portion of the mesenchymal cell-mass

which results from the second head somite,” the ventral portion

of the latter at the 5 mm. stage being in close contact with the

mesoderm of the mandibular arch. If we compare these con-

ditions with those in Vrichosurus, Phascolarctos and Phascolomys,

we may conclude with some probability that the intermediate

mass answers to the second somite of the head whose cavity is

already obliterated, or in which a cavity has never developed,

and from whose dorsal region the m. obliquus superior takes its

origin. - In Chelydra also, at a certain stage in development, the

identification of the second somite is a matter of some difficulty.

In the embryo of 7 mm. Johnson says (p. 142) :—‘‘The second

head somite of the 7 mm. stage is of such indistinct and indefinite

form that it may easily escape notice. It reaches here the most

obscure phase of its development. The more or less conspicuous

cavities of earlier stages have collapsed and broken down, and

with their disappearance the cells of their walls are with difficulty

distinguished from the intruding and intermingling mesenchymal

elements.”

No cavity is seen in the abducens muscle-mass, which in our

earliest embryo is quite solid and shows a temporary attachment,

as above mentioned, to the intermediate mass, this connection

being probably a secondary phenomenon as in Chel ydra, where it

also oceurs. It is possible that the intermediate mass, 7. e. the

second somite, may contribute towards the formation of the

m. rectus externus as in some fishes (Dohrn, 04, Neal, 14), but

we have no direct evidence of this in 7richosurus. The m. rectus

externus and m. retractor bulbi develop exactly as in Chelydra ;

the origin of the m. retractor bulbi bearing no resemblance to

that of the pig where, according to Reuter (97), p. 376 :—

“« Dieser Muskel ensteht aus dem inneren Mantel des Augen-

muskelkelches durch Abspaltung von vorn nach hinten.”

342 MISS E. A. FRASER ON THE DEVELOPMENT

In Chelydra, however, the m. retractor bulbi separates off from

the m. rectus externus at an early stage before the second somite

has completely degenerated and when the m. rectus inferior and

internus first begin to differentiate from each other, whereas in

Trichosurus it only becomes an independent muscle at 17-5 mm.

after the other muscles of the eye have assumed their final

positions and at a time when the m. rectus inferior and internus

are almost completely separated. The m. retractor bulbi in

both animals is the last of the eye-muscles to reach its adult

position.

The cranial nerves III, [V and VI arise from the brain and

grow towards their respective muscles, their connection with the

mesodermal somites being a secondary one as shown by Neal (14).

No independent origins in the muscles themselves are present as

described by Filatoff (07) in Ymys lutaria. The oculomotor runs

at its distal end into the proliferation on the postero-lateral wall

of the head-cavity and breaks up into many fibres at the region

where the m. obliquus inferior and m. rectus inferior first grow

out, these two muscles being innervated at practically the same

time; later, at the earliest indication of the m. rectus internus,

fibres are seen to run into the root of the latter. The abducens

develops in the typical manner and runs into the posterior end

of the abducens muscle-mass ; when the m. retractor bulbi grows

forwards and separates off from the m. rectus externus, the nerve

branches into the two muscles at the point where the former

crosses the medial side of the latter. The trochlear, as is usually

the case, arises later than the other two nerves. In T'richosurus

it first appears at 8°5 mm. and does not reach the m. obliquus

superior until 11 mm., when it penetrates into the posterior side

of the medial end of this muscle.

’

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346

Fig. 1.

DEVELOPMENT OF EYE MUSCLES IN MARSUPIALS,

EXPLANATION OF THE PLATES.

Lettering.

= ali-nasal cartilage. | r.b. = m. retractor bulbi.

= ciliary ganglion. r.é. = Mm. rectus externus.

= fore-brain. | ri. = wm. rectus internus.

= eyeball. | v.inf. = m. rectus inferior.

= lens. | 7S. = Mm. rectus superior.

= naso-ciliary branch of the | III. = oculomotor nerve.

trigeminal nerve. | IV. = trochlear nerve.

= m. obliquus inferior. | V. = trigeminal nerve.

= optic cup. VI. = abducens nerve.

= optic stalk. | + = band of mesodermal cells.

== m. obliquus superior.

Pirate I.

Trichosurus vulpecula. G.L. 9S mm. (YV. 701).

Coloured drawing of wax-plate model, posterior view, showing the

anterior (7.e.), and the posterior (7.b.) portions of the abducens muscle-

mass, the primordium of the m. obliquus inferior (0.inf.) extending down

from the postero-lateral corner of the solid mass of mesoderm representing

the former head-cavity. The primordium of the m. rectus superior (7.s.)

stretches up above the optic cup from the dorso-lateral side of the sane

mass. Nerve III. (III.) runs down just in front of the abducens muscle

complex and into the hinder end of the latter runs Nerve VI.(VI.) A

band of mesodermal cells (+) is seen to extend out towards the ventral

border of the eyelid. % 800 and reduced by 4.

2. Anterior view, showing the primordium of the m. obliquus superior (os.

+} t=)

with its narrow prolongation lying anteriorly to the optic cup (op.c.), the

primordinm of the m. rectus inferior (7.inf.), extending forwards below the

optic stalk from the medial side of the m. obliquus inferior (0.inf.). The

primordium of the m. rectus superior (7.s.) is again seen. The naso-ciliary

branch (V.) of Nerve V. runs forwards below the m. obliquus superior.

Z=lens. The band of mesodermal cells (+) extending out to the ventral

border of the eyelid is again seen. XX 300 and reduced by 4,

lao Oe

Trichosurus vulpecula. G.L. 15 mm.

Fig. 3. Coloured drawing of wax-plate model, anterior view, showing the m. obliquus

superior (0.s.), the m. obliyuus inferior (0.inf.), and the m. rectus inferior

(x.inf.) from which branches out the m. rectus internus (7.7.). Nerve IV.

(IV.) runs into the postero-medial side of the m. obliquus superior (0.s.).

X 150 and reduced by approximately 2.

Posterior view, showing the m. rectus superior (7.s.), the m. retractor bulbi

(r.b.) still connected with the m. rectus externus (7.e.), and the m. obliquus

interior (0.inf.) in which runs Nerve III. Nerve VI. is seen at the point

of union of the m. rectus externus and the m. retractur bulbi. A faint

dotted line indicates approximately the line of junction between the

m. rectus externus and the m. retractor bulbi. XX 150 and reduced by

approximately 2.

Ventral view, showing the distal end of the m. retractor bulbi (7.6.) which is

growing forwards round the optic stalk, the m. obliquus inferior (0.inf.)

attached to the ali-nasal cartilage (al/m.), and the ciliary ganglion (cil.g.)

lying below the optic stalk. _ Nerve III. is seen to run out from the

m. rectus inferior (7.inf.) into the m. obliquus inferior. X 150 and reduced

by approximately 2.

P Z.S.1915. BROOM. PI. IIL.

R.B.del. Cambridge University Press.

ORGAN OF JACOBSON IN TALPA AND GENTETES.

R.B.del.

ORGAN

P Z.8.1915. BROOM. PI.IV

Cambridge University Prees.

OF JACOBSON IN CHRYSOCHLORIS.

ON THE ORGAN OF JACOBSON IN THE INSECTIVORA. 347

25. On the Organ of Jacobson and its Relations in the

“ Tnsectivora.’—Part II. Yalpa, Centetes, and Chryso-

chloris. By Lieut. R. Broom, M.D., D.Sc., C.M.Z.S.,

R.A.M.C.

[Received April 27, 1915: Read May 11, 1915.]

(Plates III. & IV.*)

INDEX.

Structure : Page

Melo, OSM OF VECOOEIN os consooseoanecorondenverasso Bas)

Centetes, Organ of Jacobson ............:..-...-....--. 349

Chrysochloris, Organ of Jacobson ..................... 851

When six weeks ago L undertook the examination of the

Organ of Jacobson and its relations in Zupaia and Gymnura,

I was merely anxious to see if T'wpaia agreed with Macroscelides,

and Gymnura with Hrinaceus, as an agreement would give

further evidence of the value of this region of the skull as a basis

of classification, and also strongly support the removal of Macro-

scelides, Twpaia, and allied genera from the Insectivora and the

placing of them in a distinct order—Menotyphla. As Ihave shown

in Part I. of this paper, Z’upaia agrees sufliciently closely with

Macroscelides—both having the Polyprotodont Marsupial type

of organ—to admit of their being placed in the same order;

while Gymnura was shown to have the higher HKutherian type of

organ met with in Hrinaceus, Felis, Ovis, and all higher forms in

which the organ is retained.

When I had finished the study of these two types, I thought

it would be worth while to look at the condition in the aberrant

African Golden Mole, Chrysochloris ; and as Prof. J. P. Hill had

two very fine specimens which I had given him some years ago—

a newly-born Chrysochloris hottentota and a slightly older Chryso-

chloris asiatica—he very kindly had the heads sectioned so that I

might examine them. The results were so interesting that it

-was necessary to go further with the research. Through the

kindness of Mr. Oldfield Thomas, of the British Museum, I

obtained the snout of a Centetes which I have examined, and also

adult specimens of the common mole and shrew. Prof. Hill

very kindly had sectioned for me the head of a very young

mole. ‘To these gentlemen, and also to Prof. Hill’s laboratory

assistant, Mr. F. Pittock, [ am deeply indebted.

%* For explanation of the Plates see p. 354.

Proc. Zoou. Soc.—1915, No. XXV,

348 LIEUT. R. BROOM ON THE

The Organ of Jacobson in Talpa, (PI. III. figs. 1-7.)

Prof. Kitchen Parker in 1885 figured a number of sections

of the nose of the young of Yalpa. While these show the general

structure and relations of the nasal cartilages, they do not clearly

show the structures in the neighbourhood of the naso-palatine

camall,

Theugh the young mole which I have examined has the

cartalage in the anterior part of Jacobson’s duct imperfectly

chondvified, the young animal reveals the structures more clearly

than does the adult in which the bones are so highly ossified and

anchylosed.

In the anterior part of the nose there is little worthy of special

note. Ome unusual feature is the thinning-out and perforation

of the cartilage of the septum a short distance behind the

plane of the nostril. As the sections of the snout have been cut

of uniform thickness, the number of the section figured or

deseribed will indicate the situation of the various features.

This anterior perforation of the septal cartilage is between

sections 19 and 22.

Immediately behind the perforation the upper part of the

septum begins to thicken. Fig. 1 represents section 36. The

thick alinasal cartilage is seen curving round and united with

the thick outer part of the anterior nasal-floor cartilage. Outside

the cartilage is seen the lacrimal duct, and inside, the duct of the

nasal gland.

From the plane of this last section to section 76, the nasal

cavity is completely surrounded by cartilage. The upper part of

the septal cartilage becomes gradually more thickened. From

section 60 a median groove passes upwards into the lower part of

the septum gradually dividing the cartilage into two. Fig, 2

represents section 74. Here the lower parts of the median

cartilage are almost separated from the upper septal cartilage to

form the recurrent cartilages. The anterior part of the pre-

maxilla is seen; the bones of the two sides are not clearly

separable.

Three or four sections beyond that shown in fig. 2 the

recurrent cartilage is free from the base of the septum and also

from the lower part of the alinasal. A similar condition occurs

for about twenty sections. In section 96 is seen the anterior

part of the naso- palatine canal.

Fig. 4 represents the condition at section 100. On the left

side the duct of Jacobson’s organ is seen opening into the canal.

On the right side, the duct of the organ is seen lying above and

slightly to the outer side of the canal. The anterior part of

Jacobson’s cartilage is seen lying above the duct. The recurrent

cartilage is well developed.

Fig. 3 represents section 103. The duct of Jacobson’s organ

is seen Immediately above the naso-palatine duct.

In fig. 5, which represents section 110, the naso-palatine duct

ORGAN OF JACOBSON IN THE INSECTIVORA. 349

is shown to be passing to the outer side of the Jacobson’s duct,

and Jacobson’s cartilage coming to lie on the inner side of the

duct. The palatine process of the premaxilla is seen on one side

free from the premaxilla.

A few sections further back, as shown in fig. 7, which re-

presents section 117, Jacobson’s ¢artilage is united with the

lower part of the recurrent cartilage, and the naso-palatine canal

is seen to have opened into the nasal cavity. In the next four

sections, veins are seen to pass through two fair-siged openings

in the cartilage about the level of the upper part of the palatine

process

In fig. 6 is represented section 126. Here the duct of

Jacobson’s organ is just about to open out into the organ proper.

The palatine processes form flattened splints on the inner sides

of the cartilages of Jacobson. A portion of the secondary palatal

plate of the maxilla is seen in the section.

The organ of Jacobson as distinct from the duct extends from

section 127 to section 178. The specimen examined is too young

to show clearly the arrangement of blood-vesséls in connection

with the organ. Beyond section 178 the organ is continued into

a glandular duct, which extends for ten more sections.

Talpa agrees with Gymnura in having feebly developed

glandular ridges; in having practically no inferior turbinal in

that part of the nose in which is situated the anterior part of

Jacobson’s organ ; in having the lower half of the nasal septum

converted into recurrent cartilages: and in Jacobson’s organ

having a long duct in front which opetts into the anterior end of

the naso-palatine canal. It differs in a number of details, such

as the mode of opening of Jacobson’s duct, the less forward

extension of Jacobson’s cartilage, in the arrangement of the

blood-vessels of the organ, and in the better development of

the recurrent cartilages.

There can, however, I think be little doubt but that Zalpa is

allied to Gymnura and may conveniently be left in the old

order Insectivora.

I have made a series of sections of the snout of an adult Sorew.

Unfortunately, the specimen was not in the best state of preser-

vation, and, further, the structures are so extremely ossified that

it is difficult to interpret some of the parts, and a young specimen

will require to be examined before one can speak with any con-

fidence on the affinities. In certain features Sorex resembles

Centetes even more than it does Valpa, and an examination of

the nose of a late fetus would probably reveal some interesting

facts.

The Organ of Jacobson in Centetes. (PI. IIT. figs. 8-14.)

The specimen examined was the snout of an adult Centetes

ecaudatus in fairly good presefvation. It was sectioned by

hand. ;

In the anterior part of the snout the median septum is well

25%

» 350 LIEUT. R. BROOM ON THE

developed and of about uniform width. On passing back, the

upper and middle part becomes thickened; then a little further

back the lower half widens out and becomes divided by a median

cleft into the two recurrent cartilages, as seen in fig. 8. Though

in this section the alinasal is seen detached from the re-

current cartilage, they are seen united a few sections further

forward. If this section be compared with a corresponding

section in Z’alpa, such as shown in fig. 2, it will be observed that

though there are many minor. differences there is considerable

agreement. Even the somewhat corresponding section which

I have figured in Gymnura will be seen to agree fairly well,

while the corresponding section in Zupaia will be seen to differ

very greatly.

On passing a little further back, the septal cartilage becomes

divided into an upper and a lower part. The upper part soon

becomes greatly reduced, and then completely lost, while the

lower part is continued backwards as a rounded cartilaginous rod.

In fig. 9 the section passes through the anterior part of the

papilla. The recurrent cartilages lie by the base of the septum.

‘The upper nasal cartilages are considerably reduced.

Fig. 10 represents a section further back. It shows the

anterior part of the naso-palatine canal. On the one side,

Jacobson’s duct is seen opening into it. At this plane the only

cartilaginous support of the ducts is the anterior extension of

the posterior nasal-floor cartilage. The recurrent cartilage is still

of large size.

Fig. 11 represents a section near the plane where the naso-

palatine canal opens into the nasal cavity. The recurrent

cartilage is greatly reduced. Jacobson’s duct is seen supported

by anterior processes of Jacobson’s cartilage.

Fig. 12 represents a section a short distance behind that shown

in fig. 11. A small process of the recurrent cartilage is still seen,

and quite free from Jacobson’s cartilage. Jacobson’s cartilage is

united at its outer lower side to the posterior nasal-floor cartilage.

This is a most unusual condition, and, so far as I am aware, is

only known elsewhere in Procavia.

In fig. 13 Jacobson’s organ and the cartilage are well developed.

There is a small nasal-floor cartilage still seen. The palatine

processes are large and partly roof over the organ. Above the

palatine process is seen the anterior end of the vomer.

Fig. 14 is considerably further back. The vomer nearly clasps

the nasal septum. Jacobson’s cartilage rests on the palatal plate

of the maxilla. The palatal processes, though thin, are still of

large size. In this same section, though not shown in the figure,

is seen a large inferior turbinal supported by a slender turbinal

bone, and further up on the outer nasal wall a very well developed

glandular ridge.

Oentetes presents a number of unusual characters. It agrees

with Gymnura and Talpa in that Jacobson’s organ has a long

. : : 5

narrow duct which opens into the anterior end of the naso-

ORGAN OF JACOBSON IN THE INSECTIVORA. 351

palatine canal, but differs from both in the anterior part of

Jacobson’s cartilage being less developed. It differs from both

in the greater development anteriorly of the inferior turbinal.

It resembles Gymnura in having a single large blood-vessel along

the outer side of the organ. In the loss of the upper part of the

septal cartilage, it agrees with Sorex. I know of no other form

besides Centetes in which the lower part of the septal cartilage is

completely surrounded by the vomer, as it is a little behind the

plane of Jacobson’s organ.

Whether Centetes should be placed with Gymnura and Talpa

in the order Insectivora, or placed in a separate order of the

Ccenorhinata, the evidence from the structures in the nose does

not conclusively show, and it will therefore be well at present to

leave Centetes and its allies in a very distinct family or suborder

of the Insectivora.

The Organ of Jacobson in Chrysochloris. (Plate IV.)

I have examined the organ and its relations in a newly-born

Chrysochloris hottentota, and a somewhat older Chrysochloris

asiatica. The former had a head-length of 12 mm., and the latter

a head-length of 16 mm.

Fig. 1 represents section 40 of the nose of the newly-born

Chrysochloris hottentota. The median septum is slender; the

alinasals are very wide. In the general arrangement of the

cartilages, and in the mode of opening of the naso-lacrimal

duct, the agreement with 7wpaia is fairly close.

Fig. 2 represents section 47. It will be unnecessary in the

present paper to discuss the morphology of the small cartilages

of the nasal valve, but this section shows a remarkable recurrent

process from the outer border of the alinasal. The only other

mammal I know of which possesses a similar structure is

Echidna.

In fig. 3, which represents section 55, the most anterior part

of the premaxilla is cut through. The alinasal is continuous

with the anterior nasal-floor cartilage, and a very sharp fold

passes into the inferior turbinal.

Fig. 4 represents section 64. Here the lower part of the

premaxilla is cut across as well as the lateral. The nasal-flooi

cartilage is no longer connected with the alinasal. The septal

cartilage shows the first indications of separating into upper and

lower parts.

In fig. 5, which represents section 69, we see the division of

the septal cartilage into upper and lower parts, and the com-

mencing division of the lower into the two recurrent cartilages.

Tn this and the previous sections, the great encroachment on the

nasal cavity by the folding of the mucous membrane is seen to

be in marked contrast to the condition seen in Valpa, Gymnura,

and Centetes.

' Fig. 6, which represents section 86, cuts through the first

352 LIEUT. R. BROOM ON THE

incisors. The premaxillaries are sending up processes which

form the bases of the palatine processes. The recurrent cartilages

resemble considerably those of the Diprotodont Marsupials,

Trichosurus and Pseudochirus,

In fig. 7, which represents section 114, is seen the anterior

part of the maxilla, Here the recurrent cartilage is even more

Diprotodont-like thai in the previous section figured. The

section is through the anterior part of the papilla.

Fig. 8, which represents section 120, is through the naso-

palatine canal, It will be observed that the eanal opens directly

up into the nose, as in the typical marsupials. Jacobson’s

cartilages have given off the outer bars.

Fig. 9 represents section 123. Here Jacobson’s organ is

seen opening directly into the lower part of the nasal cavity.

This section is strikingly like figures I have given of Jacobson’s

organ in Pseudochirws and T'pichosurus.

In fig. 10, which represents section 126, the organ is seen in

its fullest development.

A short distance behind this last section a small but well

chondrified posterior nasal-floor cartilage is developed, The

cartilage on passing backwards comes to lie underneath the outer

part of Jacobson’s cartilage.

Figs. 11-13 represent three sections through Jacobson’s organ

in the young Chrysochloris asiatica. Section 11 is through the

anterior end of the organ near where it opens into the lower

part of the nasal cavity : section 12 is through the middle of the

organ, and section 13 near its posterior end. One of the most

remarkable characters of the sections is the folding-down of the

inferior turbinal till it almost forms a secondary nasal floor.

The organ itself is remarkable in having no outer vascular

plexus nor, as is seen in most Marsupials and in Gymnura, Centetes,

and Tupaia, a large vessel along the outer side. Further, while

the organ in most mammals has a large gland or numerous glands

opening into it posteriorly, Chrysochloris has mucous glands

opening into tt all along its upper border.

It will be observed that Chrysochtoris differs in the structure

of this region of the nose entirely from Gymnura, Talpe, and

Centetes, and that while it agrees more with Pupara and

Macrogcelides, the affinity is not close,

Conclusions.

The research into the structure of the anterior nasal region of

the Insectivores, even though only a few of the more striking

types have been examined, shows I think conclusively that

the ‘‘ Insectivora ” is not a natural order. Hrinaceus, Gymnura,

Talpa, Sorex, and Centetes agree in each having an organ of

Jacobson which. ends in a long duct opening inte the naso-

ORGAN OF JACOBSON IN THE INSECTIVORA. 353

palatine canal near its anterior part. Jacobson’s cartilage in

each passes forwards with the duct, and the naso-palatine canal

is supported by an anterior process of the posterior nasal-floor

cartilage. Whatever subdivision may on other grounds be made

of these Insectivores they all belong to the Ccenorhinata, and

agree with most higher mammals, such as the Carnivores and

Ungulates.

Tupaia and Macroscelides agree in having a Polyprotodont

marsupial type of structure, and cannot have any near affinity

with the typical Insectivores, and the order Menotyphla to which

they belong must be removed far from the Insectivora and

placed in quite a different phylum and not far from the early

Marsupials.

Chrysochloris, which has generally been regarded as allied to

Centetes, is seen to be in no way closely related to it. Like

Lupaia, Chrysochloris belongs to the Archxorhinata, but it

cannot belong to the order Menotyphla. The resemblance of its

nasal structures to those of the Diprotodont marsupials is

puzzling, and until further work is done on the development

of Chrysochloris, one must hesitate in discussing its aflinities

further.

In 1883 Dobson *, in discussing the affinities of Chrysochloris,

wrote as follows :—“‘ The natural relations of the species of the

family are with the Centetide, which they resemble in the form

of their molar teeth, in the absence of an interorbital constriction,

in the position of the testes, and in some important myological

characters ; but they also present many most important differ-

ences .... such as the presence and peculiar development of the

zygomatic arches, the presence of tympanic bulle, and the very

different form of the male generative organs, characters which,

while indicating separation at a very remote period from the

ancestral Centetidz, do not ally them with any other family of

recent Insectivora.”

The peculiar structure of the molars has been the main

character which has suggested an affinity with Centetes. But

we find-a similar type of teeth in the marsupial Votoryctes, and

the similarity in the three genera is probably due to convergence,

and does not indicate any attnity.

I hope shortly to undertake the study of the structure and

development of the skull, and may be able to throw some further

light on the affinities of Chrysochloris.

In the meantime I have no hesitation in removing it from the

Insectivora, and placing it in a distinct order, for which the

name Chrysochloridea, proposed by Dobson, may be retained.

* A Monograph of the Insectivora, Systematic and Anatomical, Part I1., 1883,

p- 108.

304

ON THE ORGAN OF JACOBSON IN THE INSECTIVORA.

EXPLANATION OF THE PLATES.

Lettering.

a.J.c., anterior process of Jacobson’s cartilage; b.v., blood-vessel ; d.J.0., duct of

Jacobson’s organ; g.d.J.o., duet of gland leading into Jacobson’s organ ; J.c., Jacob-

son’s cartilage; J.o., Jacobson’s organ; J.d., lacrimal duct; Mx., maxilla; Na.,

nasal; 2,f.c., nasal-floor cartilage; 1.9.d., nasal-gland duct; .p.c., naso-palatine

canal; 0.6.J.c., outer bar of Jacobson’s cartilage; Pma., premaxilla; p.Pme., pala-

tine process of premaxilla; 7.a.., recurrent process of alinasal; r.c., recurrent

cartilage; Vo., vomer.

PuateE III.

Figs. 1-4. Sections through the snout of Zalpa ewropea. Young. X 15,

5-7. Sections through the snout of Zalpa europea. Young. Showing

Jacobson’s duct. X 34.

8-9. Sections through the snout of Centetes ecaudatus. 5.

10-14. Sections through the snout of Centetes ecaudatus. Showing Jacobson’s

organ and its duct, and their relations. > 10.

PratvE LV.

Figs. 1-7. Sections through the snout of a newly-born Chrysochloris hottentota

(greatest length 28 mm.). Figs. 14, X 15. Figs. 5-7, X 20.

8-10. Sections through the papillary region of newly-born Chrysochloris

hottentota, showing the mode of opening of Jacobson’s organ. X 30.

11-13. Sections through the snout of young Chrysochloris asiatica. (About

half-grown.) X 17.

ON ANOMODONT REPTILES. 355

26. On the Anomodont Genera, Pristerodon and ee

stoma. By Lieut. R. SEOUL, MADD Sc CML ZiS

R.A.M.C.

[ Received April 27, 1915 : Read May 11, 1915.]

(Text-figures 1-4.)

InDEX.

Structure ; Systematic: Page

IEVUSHAROGIOD TOCIGIOPD, ono aceon eco c0on6000G ROR onDeND EGO CDS)

IPE GIVICE DSI Rea Psy RoR ROE, OO

d EARS IOUS mocranmedn cio deans a coo nore do Rca on oNon ec roneN sas)

IPs WIRECRGKIVDS dooce oo s4c1000009 960000980. 208800002000000 ITS)

Tropidostoma microtrema ...........00ee eee eevee 808

PRISTERODON MCKAYI Huxley. (Text-fig. 1.)

In 1868 Huxley described, under the name Pristerodon mckayi,

a fairly well-preserved skull of a small Anomodont reptile.

Almost the whole of the right side is shown in good condition,

and a considerable part of the leftside. There are also associated

with the specimen, besides the jaws belonging to the skull, a few

other :mandibular remains. Huxley apparently believed the

animal to be a lizard, as he refers to the specimen as ‘“‘ a shattered

lacertilian skull, having very much the general shape of that of

Rhynchosaurus, beg very broad posteriorly owing to the large

size of the supratemporal fossa, and tapering anteriorly.”

The skull when complete probably measured 87 mm. from

the snout to the transverse plane of the squamosals, and the

greatest breadth is about 70mm. The interorbital measurement

is 17 mm., and the intertemporal measurement 18°5 mm. The

antero-posterior measurement of the orbit is 22mm. In the

type-skull there is no tusk, the animal having been a female, but

a specimen in the South ‘African Museum shows a Dicynodon-

like tusk. The maxilla and the mandible have each, as shown

by Huxley, a series of small teeth which have high crowns

remarkable for being smooth in front and having about 8 or 9

relatively strong denticulations on the posterior side. The exact

number of teeth appears to vary with age, but in the type there

appear to be as many as 12 arranged somewhat irregularly and

one or two being probably replacing teeth.

In structure the skull is typically Dicynodont. The frontals

are large and pass backwards between the postfrontals and

preparietal. The postfrontal is well developed, but in the

specimen it is difficult to be quite sure of the sutures of its

anterior and outer end, but. they are probably as I have figured

them, in dotted line. "The postorbital is a larger bone which

forms most of the postorbital arch and the whole of the

inner border of the temporal fossa, ‘The preparietal is long and

356 LIEUT. R. BROOM ON

narrow, and extends back nearly to the pineal foramen.

however, doubtful if it reaches the foramen, the appearances

being rather in favour of the parietals just meeting in front of

the foramen and shutting out the preparietal.

fairly large flat bone. The interparietal is of moderate size and

forms the upper part of the occiput. The squamosal is large and

typically Dicynodont in character. The specimen shows part of

the quadrato-jugal in front of the lower part of the squamosal.

Text-figure 1.

wor

e .

crm em @peen em

Skull of Pristerodon mckayi Huxley. Nat. size.

Left side and snout restored. B.M., R 1810.

Fy., frontal; I.P., mterparietal; Jw.,jugal; D., lacrimal; Ma., maxilla; Wa., nasal;

Pa., parietal; Pal., palatine; Pm., premaxilla; Po.F., postfrontal; Po.O.,

postorbital; P.P., preparietal; P7r.F., prefrontal; Pt., pterygoid; P.Vo.,

prevomer; 7'.P., transpalatine.

The lower jaw is not well preserved. As is seen in one of

Huxley’s figures, the angular has, as in Dicynodon, a fan-like

expansion passing inwards and downwards. In front of the

articulation is a plate of bone passing outwards and forwards,

apparently part of the surangular.

It is,

The parietal is a

ANOMODONT REPTILES. 357

PRISTERODON RANICEPS (Owen) *. (Text-fig. 2.)

In 1876 Owen briefly described, under the name Ouxudenodon

raniceps, a small skull and jaws from the same locality, Kast

London, as Huxley’s type. Owen gives no figure of the specimens,

but merely describes the skull as follows :—“ The occipital condyle

is low and broad; the lateral portions contributed by the

exoccipitals simulate the Batrachian double condyle. ‘The supra-

occipital surface is much depressed. The temporal fosse are large,

with the long diameter lengthwise. The orbits appear to have

been small.”

Text-figure 2.

Frontal and parietal regions in Pristerodon waniceps (Owen). Nat. size.

B.M., R 1650.

Lydekker, in his Catalogue of 1890, refers the specimens

doubtfully to Udenodon baini Ow., believing that they were of

a young individual.

It is rather remarkable that both Owen and Lydekker should

have referred the specimens to Oudenodon, seeing that a consider-

able number of teeth are well exposed to view in the lower jaw.

The teeth agree closely with those of Pristerodon mckayi, but

the denticulations are relatively rather coarser,

The skull is much crushed, but probably measured about

92 mm. from the snout to the plane across the posterior borders of

the squamosals, and the greatest breadth is about 80 mm. The

interorbital measurement is 16 mm.and the intertemporal 28 mm.

The antorbital portion of the skull appears to be relatively shorter

than in Pristerodon mckayi. Unfortunately no sutures can be

clearly made out in the present condition of the specimen, but,

as will be seen by the figure I give of the parietal region, the

* [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. |

358 LIEUT. R. BROOM ON

relative proportions of the parts differ very considerably from

those of the type species.

In the lower jaw there appears to be a series of 6 teeth with

some replacing ones.

Seeing that Owen’s specimen came from the same locality as

Huxley’s I was inclined to regard thein as belonging to the same

species, but the differences in proportions appear to be sufficiently

great to warrant us at least provisionally in regarding them as

distinct.

PRISTERODON AGILIS (Broom).

In 1904 I described, in the Records of the Albany Museum,

the skull of a small Anomodont found by me at Pearston. At

that time it was not known that Oudenodon is the female of

Diceynodon, and as the Capetown specimen of Pristerodon mckayr

is tusked, and the new allied form which I discovered being

tuskless, I placed it in a new genus and called it Opisthoctenodon

agilis. As, however, the tusk is now known to be a sexual

character, this species must be placed in Huxley’s genus

Pristerodon.

The skull is much smaller than in P. mckayi and narrower

relatively, but in most characters it agrees closely, and the molars

are very similar,

Pristerodon mckayi and P. raniceps both come from the

Pareiasaurus zone, but P. agilis is from the Hndothiodon zone.

PRISTERODON BRACHYOPS (Broom).

This species when described was also placed in the genus

Opisthoctenodon. As the crowns of the molars ave unknown it is

impossible to be certain that it belongs to LPristerodon, but it

agrees sufficiently to admit of its being placed here, at least

provisionally, It is probably from the Cistecephalus zone.

TROPIDOSTOMA MICROTREMA (Seeley). (Text-figs. 3 & 4.)

In 1889 Seeley described an Anomodont occiput under

the name Dicynodon microtrema. The occiput formed part

of Mr. T. Bain’s collection and was obtained, according to

Mr. Watson, on the farm Tafelberg, in the Beaufort West

district. In the British Museum there is a good series of

Dicynodont skull-remains from the same locality, and the

majority belong, I think there is little doubt, to the one species.

Fortunately there are a number of snouts and a few mandibles.

Both tusked males and tuskless females are represented, and

most specimens show evidence of small molars behind the tusk,

or caniniform process. ‘The specimens differ considerably in size,

doubtless due to difference in age, and the differences in dentition

are probably due to the same factor. There appear to be

normally four molars, but as age advances they become reduced

to three, two, or one, and in old age get completely lost.

ANOMODONT REPTILES. 359

A small mandible shows the crowns of two teeth in fairly good

preservation. They are seen to agree pretty closely with those of

Pristerodon. Though the teeth are rather more robust, they are

really relatively very much smaller. One might incline on the

evidence of the teeth to place these large specimens in the

genus Pristerodon, but the structure of the parietal region differs

so considerably that it is necessary to place them in a different

genus. The snout agrees so closely with Dicynodon except for

the presence of the molars, that one has to consider whether it

might not be possible that all species of Dicynodon had molars

when young which they lost later. This, however, is hardly

possible. In no species of Dicynodon have molars ever been

detected—even where the skulls are manifestly of young animals.

Of Dicynodon platyceps we know over a dozen skulls, from small

ones about 3 inches to others over a foot in length. But in none

is there any trace of molars. Further, the peculiar condition of

the intertemporal region found in the present species is unknown

Text-figure 3.

3ones of the frontal and parietal regions of the skull of Tropidostoma

microtrema (Seeley). About + nat. size.

For lettering see text-fig. 1, p. 356.

in any species of Dicynodon. Wemay, therefore, safely conclude

that Dicynodon microtrema Seeley must be placed in a distinct

genus. Another specimen, figured by Seeley in 1889, was an

occiput which he named Dicynodon (Lropidostoma) dunni.

Lydekker regarded this specimen—quite rightly, I think—as

belonging to the same species as that named D. microtrema. And

if this be so we must accept the generic name Tropidostoma

for the type.

Though there is no complete skull in the collection, one is

fairly complete, and there are so many snouts, occiputs, and

other portions, that practically every detail of the structure can

be made out.

The skull, in what appears to be an adult male, measures from

the snout to the plane cutting the posterior borders of the

squamosals 266 mm., and the greatest width across the squamosals

is about 220 mm,

‘

360 LIEUT. R. BROOM ON

The following are the other principal measurements :—

Measurement between the canines, 40 mm. to 48 mm.

Interorbital width, 25 mm.

Measurement across the nasal bosses, 44 mm. to 52 mm.

Interorbital measurement, 36 mm. to 38 mm.

Intertemporal measurement, 28 mm. to 36 mm.

The premaxilla is relatively narrow in front. It has the

usual pair of palatal ridges seen in most Anomodonts. It forms

much the larger part of the hard palate, and meeting the palatine

behind shuts owt the maxilla from the margin of the posterior

nares. In the middle line the premaxilla meets the prevomer,

as shown in the figure.

Yext-figure 4.

Anterict palatal region of Tropidostoma microti ema (Seeley). 4 nat. size.

Showing the relations of the palatal elements and tooth-sockets behind the canines.

B.M., R 860.

For lettering see text-tig. 1, p. 356.

The maxilla is very similar to that of Dicynodon except for the

presence of molarteeth. With the premaxilla it forms the lower

border of the nostril, meeting the septomaxilla. Its upper

border articulates with the lacrimal, which completely separates

it from the nasal.

In nearly all the specimens of Tropidostoma microtrema there

are seen some molar teeth or tooth-sockets. In young specimens,

which are about half the adult size, there are four molars. of

which the first is the largest, and they decrease in size to the

fourth. The four molars measure 10mm. Only one specimen

shows the crowns. These resemble closely the molars of

Pristerodon, but those of Tropidostoma are rather stouter and

the denticulation slightly coarser. In the Ist molar there are

ANOMODONT REPTILES. 361

probably seven denticulations, and in the 3rd, five. As Tropido-

stoma approaches the adult condition the molars seem to be

gradually lost. Some specimens show only three, others but two

or one, and in old age all trace of even the sockets disappears.

In those specimens which are regarded as males there is a

pair of powerful tusks. In the supposed females, in place of the

tusks are, as in ‘‘ Oudenodon,” caniniform processes. One

specimen (R871) is remarkable for having on the right side a

well-developed tusk, and on the left side no trace whatever of a

tusk. This specimen is probably the oldest animal of the series,

and I regard it asa female which has developed a tusk on the

one side.

The septomaxilla lies within the nostril forming its lower wall.

The lacrimal extends forwards between the nasal and the

maxilla, and meets the septomaxilla.

The relationships of the bones of the top of the skull will

best be understood from the diagram I have given. The greater

part of the figure is drawn from specimen R 860, but the anterior

and posterior portions are from two other specimens. Nearly

every suture is confirmed by more than one specimen.

The most noteworthy features of the upper side of the skull

are the presence of thickened bosses on the nasals, and the

development of the postorbitais and parietals intoa pair of lateral

crests with a deep groove between them.

The palate has already been figured by Lydekker; but as I

think he is in error in a number of points, and as the whole

structure of the anterior palatal region is exceedingly well shown

in specimen R 860, I think it worthy of being figured again.

The great size of the palatal portion of the premaxilla is seen and

the relations of the palatine, transpalatine, and pterygoid bones

to each other. The prevomer has its lower border developed as

a pair of plates. In this Tvropidostoma differs from Dicynodon,

and agrees with Hndothiodon and Emydorhynchus.

The occiput has been figured and described by Seeley.

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ON TRIASSIC SLEGOCEPHALIANS, 363

27. On the Triassic Stegocephalians, Brachyops, Bothriceps,

and Lydekkerina, gen. nov. By Lieut. R. Broom, M.D.,

DiSe., C-M:Z.S., R-A.M.C.

[Received May 26, 1915; Read June 8, 1915. |

(Text-figures 1-3.)

INDEX.

Page

Systematic :

. 366

Lydekkerina, gen. nov.

Structure ;

Brachiyops Vaticeps \..c.1ccecee-ee eee ses eee eeneseene vss BOD

Bothriceps australis a6 . 3864

Lydekkerina hualeyt ....200cccccceeveeeenee veeeee eee 366

BRACHYOPS LATICEPS Owen. (Text-fig. 1.)

In 1854 Owen described, under the name Brachyops laticeps,

the skull of a small Stegocephalian from Mangali, Central India.

Though the skull is fairly complete it is very badly weathered,

Text-figure 1.

Skull of Brachyops laticeps, about 3 nat. size.

Va., Nasal; Pa., Parietal ;

Pmzx., Premaxilla; PoF., Postfrontal ; PoO., Postorbital; Pr.F., Prefrontal ;

P.Pa., Postparietal; Q.J.. Quadratojugal; Sq., Squamosal; S.Sq., Supra-

Fr., Frontal; Ju., Jugal; D., Lacrimal; Mex., Maxilla;

squamosal (Supratemporal) ; 7., Tabular.

Proc. Zoou. Soc.— 1915, No. XX VI. 26

364 LIEUT. R. BROOM ON .

and very little can be made out beyond the impression of the

inner sides of the upper cranial bones. Very little remains of

the elements in front of the orbit and practically nothing of the

tabular and postparietal regions.

The parietals, which are clearly paired, are unusually large and

have between them a large pineal foramen. The anterior end of

the parietal is in front of the transverse plane through the

postorbital margin.

The frontal is relatively small. The postfrontal and post-

orbital are of about equal size—each a little larger than the orbit.

The jugal has a very well-developed portion lying between the

orbit and the quadratojugal and squamosal, and here there has

evidently been a moderate-sized boss.

The squamosal is large and the suprasquamosal only a little

smaller. The tabular has evidently been small and the post-

parietals short antero-posteriorly.

The occipital region slopes back from the postparietals, and

the exoccipital condyles are considerably behind the plane of the

quadrates.

In the figure I have given, the sutures in line are those seen on

the specimen ; those in broken line are from those of the opposite

side ; those in dot are hypothetical.

BotHRIcEPs AUSTRALIS Huxley. (Text-fig. 2.)

In 1859 Huxley described a small Stegocephalian from

Australia, under the name Bothriceps australis. ‘Though smaller

than Brachyops laticeps it is fairly closely allied to it. Huxley

gives reasons sufficient to show that it is at least specifically

distinet. The few reasons he gives for regarding it as generic-

ally distinct are less conclusive. Brachyops has a broader skull,

and the eye is placed further forward and more laterally than in

Bothriceps, but in the imperfect state of the specimens no

characters of generic importance can be seen to separate the two

forms. ‘There may, however, be in association with the anterior

portion of the orbit in Brachyops, some differences in relations of

the lacrimal, or septo-maxillary, or of the sensory grooves, or

in the structure of the palate, which may be sufficient to separate

the types into distinct genera, and in the meantime Bothriceps

may be retained as possibly distinct.

The figure given by Huxley shows the sutures as preserved in

the specimen. The new figure I give represents a restoration

of the skull. As, however, every element of the upper side of

the skull is shown on one side or the other, there is no element

in any doubt, and the only sutures concerning which there is any

doubt are indicated in dotted line.

The following are some of the more interesting points in the.

structure of the skull. The lacrimal is small, and nearly extends

from the orbit to the nostril, but does not reach the border of

TRIASSIC STEGOCEPHALIANS. 365

either. The postfrontal is more than twice as long as broad.

The jugal forms the lower margin of the orbit, but much the

larger part of the bone is situated behind the orbit. ‘The supra-

squamosal is only about half the size of the squamosal. |

The occiput slopes away from the postparietals as in Brachyops,

but being much better preserved we can make out something of

the structure in Bothriceps. The postparietal is well developed,

and has a considerable part on the occipital aspect which articu-

lates with the exoccipital. The tabular is smaller than the

Text-figure 2.

Z Y a a y :

M1 3 / Mi) .

oy,

‘

Skull of Bothriceps australis, about % nat. size.

For lettering see text-fig. 1.

postparietal, and if only the upper surface were seen might be

regarded as forming a small posterior horn, but the exoccipitals

extend much behind it, and inferiorly and internally it articu-

lates with what is probably the paroccipital.

The parasphenoid is large and there are no teeth on it.

Brachyops, Bothriceps, and Batrachosuchus are allied genera

forming a very distinct family which may be called the

26*

366 LIEUT. R. BROOM ON

Brachyopide. It is interesting that one should be known only

from India, one from Australia, and the third from South

Africa,

LYDEKKERINA HUXLEYI (Lydekker), gen. nov. (Text-tig. 3.)

In 1890 Lydekker described a small Stegocephalian from the

Orange Free State under the name Bothriceps huwleyi, believmg

the form to be allied to Huxley’s Bothriceps australis. No later

worker, so far as I am aware, has doubted the correctness of

Lydekker’s placing it in Huxley’s genus. But as I hope to show

that the South-African form differs very markedly from Bothri-

ceps, and possibly even belongs to a different family, I propose

to establish a new genus for it, which I have much pleasure in

calling Lydekkerina, after Mr. R. Lydekker, who has so recently

passed away, and whose work on South African fossil reptiles

was extremely good.

The only known specimens of Lydekkerina hualeyi are four

skulls, with some portions of the rest of the skeleton, in the

British Museum, and a number of fairly good specimens in the

Bloemfontein Museum ; and all were obtained, I believe, from a

locality near Edenburg, O.F.S. -

The skull is about a half longer than broad. The orbits are

moderately round and placed near the middle of the skull. The

nostrils are large; and there is a very distinct otic notch

bounded internally by a well-developed tabular.

The premaxillaries are fairly well developed and the maxillaries

are long but very slender. The septo-maxillary, if developed as

is probable, is entirely inside the nostril.

The nostril is large, and is separated from the maxilla by the

lacrimal.

The lacrimal, is very well developed, extending backwards from

the nostril to near the orbit, and having on its inner side the

nasal and prefrontal, and on its outer the maxilla and jugal.

The prefrontal is slightly larger than the lacrimal, and forms

most of the anterior margin of the orbit.

The frontal is long and narrow. The prefrontal is also narrow,

and of about the same length as the frontal. The postorbital is

broad and slightly smaller than the prefrontal.

The jugal is long, and in the orbital region fairly wide. It

lies above the maxilla, and along its upper border are the

lacrimal, the prefrontal, the orbit, the postorbital, and the

squamosal. Nearly as much of the jugal lies in front of the orbit

as behind it.

The parietal is about as long as the frontal, but slightly

broader. There is a small pineal foramen situated between the

bones, and nearer to the anterior than to the posterior borders.

The suprasquamosal (supratemporal) lies on the outer side of

the parietal. It is about the same size as the postorbital.

TRIASSIC SYEGOCEPUALIANS. 367

The squamosal is fairly large. It lies between the supra-

squamosal and the quadratojugal, and passes downwards on the

anterior wall of the otic notch to meet the pterygoid.

The quadratojugal is rather smaller than the squamosal.

The postparietal is a small quadrangular bone. On its outer

side lies the tabular, which is produced backwards into what,

when viewed from above, looks like a little posterior horn.

Text-figure 3.

ten sae eet

‘

Ci et . }

Ree ~-

Skull of Lydekkerina huevleyi, nat. size.

For lettering see text-fig. 1.

The occiput is not sufficiently well preserved in any of the

Specimens to show the sutures, but the general structure can be

satisfactorily made out, It differs from that of both Brachyops

and Bothriceps in having the condyles relatively small, and in

their not extending back behind the plane of the upper part of

the postparietals, so that they are not seen when the skull is

viewed from above. ‘There is a relatively large opening between

the exoccipital and postparietal on the one hand, and the par-

occipital and tabular on the other.

The palate differs from that of Bothriceps in having on the

368 ON TRIASSIC STEGOCEPHALIANS.

parasphenoid, the prevomers, and the pterygoids, innumerable

very minute teeth. In this it agrees with the large South

African form hinesuchus.

The mandible, so far as can be seen in the specimens, agrees

closely with that of Zrimerorhachis. On the lower and outer

‘sides can be seen a small splenial in front, with behind it a rather

larger preangular, and behind this latter a large angular. The

preangular has on its inner side relations to the prearticular

and to what is probably the intercoronoid similar to those of

Trimerorhachis. ‘The structure of the jaw is best seen in the

British Museum specimen R 506.

ON THE SNAKES OF MADAGASCAR. 369

28. A List of the Snakes of Madagascar, Comoro, Mascarenes,

and Seychelles. By G. A. Boutencer, F.R.S., F.Z.S.*

[Received May 12, 1915: Read May 25, 1915. |

INDEX.

GEOGRAPHICAL: Page

Madagascar etc., List of the Snakes.......00..0.....0cc = 869

SYSTEMATIC:

Ophidia. Keys to the identification of the Snakes of Mada-

PAS CAT CLC HUN Rae NeR ae! Hie A on is te Godt AMAR E spite, OO

This list has been prepared on the same lines as the one

recently published in these ‘ Proceedings’ 7, and likewise with the

help of Mr. F. Kingsbury and M. G. de Witte. The reader is

referred to the previous paper for an explanation of the terms

used in the keys to the genera and species.

The fauna of the islands here dealt with is remarkable, not

only for its very striking difference from that of the African

continent, but also for the absence of Snakes dangerously

poisonous to man (Elapine Colubrids, Viperids), with the ex-

ception of the two Sea-snakes which are known from the western

parts of the Indian Ocean.

Synopsis of the Families.

I. Worm-like, with small inferior mouth ; eyes hidden or visible

under the head-shields; body covered with uniform imbri-

cate scales above and beneath .............-......:.c0000eeeeeee LYPHLOPIDE.

II. Mouth large; eyes exposed; body with enlarged shields beneath (except im

Sea-snakes, with strongly compressed tail).

Frontal and parietal-shields absent, or broken up into small

eT RICINGIG), | asl cae sane ea anenser ca arene tac bata ttprantemensnescepeNetrans at! eXORSID)E2

Frontal and parietal shields present..........0...0..::::0c0 eee, COLUBRIDZ.

Family TyPHLOPIDA,

A single genus.

. 1. TYPHLops.

Schneid. Hist. Amph. it. p. 339; Bouleng. Cat. Sn. 1. p. 7.

Synopsis of the Species.

I. Prieocular present.

A. Eyes distinct.

a. Snout rounded; prxocular not much narrower than ocular; diameter of

body 35 to 55 times in total length.

Nasal completely divided, the anterior portion extending to

the upper suriace of the head; 20 scales round middle

GIP ]NGION) ee ceeeresascendace escabosne coadacadeenasts senceedersocoomtnn fe Gaels HOU LOU

Nasal not completely divided ; 20 scales round middle of

body; preocular as broad as ocular ........................ Z. eomorensis.

* Published by permission of the Trustees of the British Museum. :

+ “A List of the Snakes of the Belgian and Portuguese Congo, Northern Rhodesia,

and Angola,” P. Z. S. 1915, p, 193. ovaries

370 MR. G. A. BOULENGER ON THE

Nasal not completely divided; 20 scales round middle of

body; preocnlar narrower than ocular .................. T. microcephalus,

Nasal not completely divided; 24 scales round middle of

IOCKY aeoosenea cesses... LT. mucronatus.

6. Snout seh cobiusely janes Houontal edge; nasal semi-divided ; prse-

ocular much narrower than ocular.

Diameter of body 44-50 times in total length ; 20-22 scales

round middle of body ..... sonancsadnosaosdes » lb DOARIGCIU

Diameter of body 39 times in total length; 26 scales round

middle of body . ete Scots ‘abesdesu ose 2 2Es Glaconaads

c. Snout ohh harp horizontal eile nasal com-

pletely divided ; diameter of body 52-68 times

in total length ; 20 scales round middle of body. TT. arenarius.

B. Eyes hidden ; nasal incompletely divided ; diameter

of body 46 times in total length ; 24 scales round

middle of body .. See eee eee eee ieee CL madagascanelisis.

Tl. Preocular Anta eyes tien snout with sharp

horizontal edge: nasal completely divided; dia-

meter of body 71-78 times in total length ; 20

scales round middle of body ........................ UT. grandidiert.

1. TYPHLOPS BRAMINUS.

Eryx braminus Daud. Hist. Rept. vil. p. 279.

Typhlops braminus Bouleng. Cat. Sn. 1. p. 16.

Southern Asia; islands of the Indian Ocean; South Africa ;

Mexico (probably transported by human agency).

2. TypuLops comorENsIs Bouleng. Ann. & Mag. N. H. (6) iv.

IESE jos BOM. camel te, © jos Alle

Comoro Islands.

3. TYPHLOPS MICROCEPHALUS Werner, Jahresh. Ver. Nat.

Wirttemb, Ixv, 1909, p. 60.

Madagascar,

4, TyPHLOPS MUCRONATUS.

Typhlops (Ophthalmidion) mucronatus ect! Zool. Anz. 1880,

p. 279, and Abh. Senckenb. Ges. xii. 1881, p. 438, jollgeis ake, Il

Typhlops mucronatus Bouleng. t. ¢. p. 37.

Madagascar (Nossi Be).

5. TYPHLOPS BOETTGERI Bouleng. t. ¢. p. 39, pl. 1. fig. 6.

Madagascar.

6. TyeHLops DEcoRsET Mocquard, Bull. Mus. Paris, 1901,

p. 255.

Madagascar.

7. TYPHLOPS ARENARIUS.

Onychocephalus arenarius Grandid. Ann. Se. Nat. (5) xv. 1879,

art. 20, p. 9.

Typhlops arenarius Bouleng. t. ce. p. 49.

Madagascar.

SNAKES OF MADAGASCAR. Bl

8. TYPHLOPS MADAGASCARIENSIS Boettg. Abh. Senckenb. Ges.

xg IBiieope depletion! 3 Boulemg nbs Coup. /25.

Madagascar (Nossi Be).

9. TYPHLOPS GRANDIDIERI Mocquard, Bull. Mus. Paris, 1905,

p- 287.

Madagascar.

Family Boip #.

Synopsis of the Genera.

I. Head distinct from neck; scales smooth or unicarinate.

Scales smooth; labials with deep pits ..... . Corallus.

Scales smooth; nasals separated iM small. scales ; ‘tail short (20-41

subcaudals) eee . Boa.

Scales keeled ; nasals- separ ated by # a pair -of internasals ; 5 “tail long (120-

133 subcandals) eee nee AA RU Aen MOE ee sean eee ome OSU REUS

If. Head not distinct from neck; scales tri- or quinquecarinate ...... Bolieria.

1. CorRALLus.

Daud. Hist, Rept. v. p. 106; Bouleng. Cat. Sn. 1. p. 99.

1. CORALLUS MADAGASCARIENSIS.

NXiphosoma madagascariense Dum. & Bibr. Erp. Gén. vi. p. 549.

Corallus madagascariensis Bouleng. t. ¢. p. 103.

Madagascar.

; Boa.

Linn. Syst. Nat. i. p. 373 ; Bouleng. Cat. Sn. 1. p. 116.

Two species :—

11-16 scales round the eye; scales in 59-65 rows......... B. dumerilii.

6-9 scales round the eye; scales in 69-77 rows............. B. madagascariensis.

1. BoA DUMERILIL.

Acrantophis dumerili Jan, Icon. Gen. p. 83, 1. 1, pl. il.

Boa dumerilvi Bouleng. t. ¢. p. 120.

Madagascar.

29, BoA MADAGASCARIENSIS.

Pelophilus madagascariensis Dum. & Bibr. Erp. Gén. vi. p. 524.

Boa madagascariensis Bouleng. t. ¢. p. 120.

Madagascar.

3. CASAREA,

Gray, Zool. Miscell. p. 43; Bouleng. Cat. Sn. i. p. 121.

1. CASAREA DUSSUMIERI.

Boa dusswmieri Schleg. Phys. Serp. 11. p. 396.

Casarea dussumiert Bouleng. t. c. p. 121.

Round Island, near Mauritius.

372 MR. G. A. BOULENGER ON THE

4, Bourertia.

Gray, Zool. Miscell. p. 46; Bouleng. Cat. Sn. i. p. 121.

1. BoLiERIA MULTICARINATA.

Eryx multocarinata Boie, Isis, 1827, p. 513.

Bolieria multicarinata Bouleng. t. c. p. 122

Round Island, near Mauritius.

Family CoLUBRID &.

Aglypha.

Synopsis of the Genera.

I. Pupil round; scales smooth (Madagascar and Comoro Islands).

A. Body cylindrical; scales not oblique.

1. Snout rounded; anal divided.

a. Kye moderate or rather large; nostril between two nasals; subcaudals

65-160; scales in 17 or 19 rows.

Head short, not or but scarcely distinct from necks frontal

not narrower than supraocular ......... secesessseeese Polyodontophis.

Head moderately elongate, distinct from | neck ; ‘frontal not

narrower than supraocular, usually shorter than parietals . Tropidonotus.

Head elongate, distinct from neck; frontal, in the middle,

narrower than ecru, its length not less than that of

parietals .. Bete .. Dromicodryas.

b. Eye naib Tena: ea Singles 5 series “0 191 rows ;

subcandals 410... eee, Compsopiis.

c. Eye small; scales in 17 to 25 rows ; ~ gelbeauslals 35 to 64,

a. Loreal present.

Nostril between the nasals and the internasal; frontal pio

twice as long as broad ......... .. Liophidium.

Nostril between ‘the nasals; ‘frontal not more ham ¢ once > and a

hnalijasilong jas bxdad orien setae eee Lee ee tne Psendonyrhopuss

(3. Loreal absent.

Nostril between the nasals and the internasal; frontal a little

longer than broad; ventrals 221-255 ...... . Idiophis.

Nostril between the nasals ; frontal a little broader than ‘long;

WEIN HCMIS G3} ooeon senses cor .. Pararhadinea.

2. Snout pointed, strongly mEnecoine ocuals very 7 tra,

trihedral ; scales in 21 or 23 rows; anal entire ...... Lioheterodon.

3. Snout depressed, with horizontal edge; scales in 17

rows; anal divided ............... F ...... Heteroliodon.

B. Body slightly pee ccalesh narrow, » obliane, in

PEARS aster aa soseeeoeosonAseuerocansnoucsccoosn WMeEROpISbOdon.

Il. Pupil vertically ilies ; realy opieanen (Sey pteies and Mascarene Islands).

Scales keeled, in 17 rows; no loreal .....0.......0.00..00cccccc00000... Lyycognathophis.

Scales moon Tim PAIL WP BB} TONGS — soscaucascosctoscacoocotspevnvernoocce IEDM AD.

Sealesismoo uly mmul// MOWSi erin. .eusse- Use seereeae cree nee neOmo7iE

1. PoLyopontopuis.

Bouleng. Faun. Ind., Rept. p. 301, and Cat. Sn. i. p. 181.

Three species :—

I. Scales in 17 rows.

Loreal deeper than long; frontal much shorter than parietals... P. rhodogaster.

Loreal as deep as long or longer than ey frontal but slightly

shorter than parietals ssbecnssoenasacnasaspbsdsboecdcesenoboacnesdos0a | eo UOLGUMTEBe

JOG, SyemMlesss a IUD OWS cocoon cazuasnnesno poate sve wsanundnortnnaaapananoar 2s MOGOHCRBES:

SNAKES OF MADAGASCAR. aes

1. PoLYODONTOPHIS RHODOGASTER.

Herpetodryas rhodogaster Schleg. Phys. Serp. 1. p. 193.

Polyodontophis rhodogaster Bouleng. Cat. Sn. i. p. 182.

Madagascar.

POLYODONTOPHIS TORQUATUS.

Coronella torquata Bouleng. Ann. & Mag. N. H. (6) 1888,

p. 103, pl. v. fig. 3.

Polyodontophis torquatus Bouleng. Cat. Sn. 1. p. 183.

Madagascar.

. PoLYODONTOPHIS MAYOTTENSIS.

ae rhodogaster, var. mayottensis Peters, Mon. Berl. Ac.

1873, p. 793.

Polyodontophis mayottensis Bouleng. Cat. Sn. 1. p. 185.

Mayotta, Comoro Islands.

2. 'TROPIDONOTUS.

Kuhl, Bull. Se. Nat. ii. 1824, p. 81; Bouleng. Cat. Sn. 1.

je UM

Liopholidophis Moequard, Bull. Mus. Paris, 1904, p. 302.

Synopsis of the Species.

TI. Scales in 17 rows.

oe nearly as broad as deep; eye rather small; subcaudals

2-152...... T. sexlineatus.

Rostra much broader than deep eye ‘rather lar we; subcaudals

97-160.. Sy vesstvereese.. 2 dolichocercus.

Rostral twice as broad as s deep ; : ‘subcaudals 221......cccccec. grandidieri.

II. Scales in 19 rows; subcaudals 68-104.

Frontal once and a half as long as broad.. § Pe eRSOCETUp i Uus

Frontal once and two-thirds to twice as long # as broad ............. 7. lateralis.

1. TROPIDONOTUS SEXLINEATUS.

Dronicus sexlineatus Giinth. Ann. & Mag. N. H. (5) ix. 1882,

p. 264, fig.

Tropidonotus sexlineatus Bouleng. t. c. p. 246.

Madagascar.

2. TROPIDONOTUS DOLICHOCERCUS.

Dromicus dolicocercus Peracea, Boll. Mus. Torin. vil. 1892,

mos WZ ple ty eet.

Tropidonotus dolichocercus Bouleng. t. c. p. 246.

Madagascar.

3. TROPIDONOTUS GRANDIDIERI.

Liopholidophis grandidiert Mocquard, Bull. Mus. Paris, 1904,

p. 304.

Madagascar.

374 MR. G. A. BOULENGER ON THE

4, TROPIDONOTUS STUMPFFIL.

Dromicus stumpfii Boettg. Zool. Anz. 1881, p. 358.

Tropidonotus stumpfii Bouleng. t. c. p. 247.

2? Liophidiwm gracile Mocquard, Bull. Mus. Paris, xiv. 1908,

p. 261.

Madagascar.

5. TROPIDONOTUS LATERALIS.

Leptophis lateralis, part., Dum. & Bibr. Erp. Gén. vi. p. 544.

Tropidonotus lateralis Bouleng. t. c. p. 248.

Madagascar.

3. DROMICODRYAS.

Bouleng. Cat. Sn. 1. p. 189.

Two species :—

Subcaudals 81-117 ; usually two light dorsal stripes ...... D. bernieri.

Subcaudals 108-122; four black dorsal stripes ............ D. quadrilineatus.

1. DRomIcoDRYAS BERNIERI.

Herpetodryas berniertti Dum. & Bibr. Erp. Gén. vi. p. 211,

pl. Ixvi.

Dromicodryas bernieri Bouleng. t. ce. p. 189.

Madagascar.

2. DRoOMICODRYAS QUADRILINEATUS.

Herpetodryas quadrilineatus Dum. & Bibr. t. e. p. 212.

Dromicodryas quadrilineatus Bouleng. t. c. p. 190.

Madagascar.

4. CoMPSOPHIS.

Mocquard, CR. Soc. Philom. 1894, no. 17, p. 8.

1. CompsoPHis ALBIVENTRIS Mocquard, 1. c.

Madagascar.

5, LiopHipium.

Bouleng. Cat. Sn. 11. p. 598.

1. LiopHipruM TRILINEATUM Bouleng. t. c. p. 599.

Madagascar.

6, PsEUDOXYRHOPUS.

Ginth. Ann. & Mag. N. H. (5) vii. 1881, p. 359; Bouleng.

Cat. Sn. i. p. 314.

Rhabdotophis Werner, Jahresh, Nat. Ver. Wiirttemb. Ixy.

1909, p. 58.

SNAKES OF MADAGASCAR, 375

Seven species :—

I. Seales in 25 rows; loreal twice as long as deep; frontal

meamhy AS long! MS WOME — coooscasosssacanvousscoockbosscuonarnes Leb ULUCIROOSs

II. Scales in 21 rows.

8 upper labials, 4th and Sth entering eye; rostral just visible

froma Overereeaneneetcec sects cc sent csacace us sseoanenecesn, | ee OCET ures:

8 upper labials, 4th and 5th entering eye; portion of rostral

visible from above at least half as long as its distance

trom the frontal atee Ree eRe ee tees ede ene en ) ea Qeenguelanentus.

7 upper labials, 3rd and 4th entering eye........................... DP. ambreensis.

TOC SCENES sh) UD) TRONS co cooboo soo snolboo nnobne ubeibbeDspodonéonacnbason. .. Leo PHUGLFUKcas

IV. Scales in 17 rows.

Rostral broader than deep; ventrals 183; subcaudals 64 ...... P. occipitalis.

Rostral deeper than broad; ventrals 167; subcaudals 37 ...... P. subcaudalis.

1. PsEUDoxyruHopus Microps Giinth. Ann. & Mag. N. H. (5)

vii. 1881, p. 359, fig.; Bouleng. t. c. p. 315, & i11. p. 613.

Pseudoxyrhopus dubius Mocquard, Bull. Mus, Paris, 1904,

no. 6, p. 305.

Madagascar.

2. PSEUDOXYRHOPUS HETERURUS.

Homalocephalus heterurus Jan, Arch. Zool. Anat. Phys. ii.

1863, p.. 286.

Pseudoxyrhopus heterurus Bouleng. Cat. Sn. i. p. 315.

Madagascar.

3. PsEUDOXYRHOPUS QUINQUELINEATUS.

Liophis quinquelineatus Giinth. Ann. & Mag. N. H. (5) vu.

1881, p. 359, fig.

Pseudoxyrhopus quinquelineatus Bouleng. t. c. p. 315.

Madagascar.

4. PsSEUDOXYRHOPUS AMBREENSIS Mocquard, CR. Soc. Philom.

1894, no. 9, p. 4; Bouleng. Cat. Sn. ii. p. 613.

Madagascar.

5, PsEUDOXYRHOPUS IMERINA.

Liophis imerine Giinth. Ann. & Mag. N. H. (6) v. 1890, p. 71.

Pseudoxyrhopus imerine Bouleng. Cat. Sn. 1. p. 316.

Madagascar.

6. PsEUDOXYRHOPUS occIPITALIS Bouleng. Cat. Sn. ii. p. 613.

Madagascar.

7. PSEUDOXYRHOPUS SUBCAUDALIS.

Rhabdotophis subcaudalis Werner, Jahresh. Nat. Ver. Wiirt-

temb. Ixv. 1909, p. 58.

Madagascar.

376 MR. G. A. BOULENGER ON THE

7. [pIoPHis.

Mocquard, Bull. Mus. Paris, 1901, p. 252.

1. Ipiopmis yvariuanti Mocquard, |. c. and Bull. Soc. Philom. (9)

Ti WOM. jos AS alle tee 3}.

Madagascar.

8. PARARHADINAEA.

Boettg. in Voeltzk. Reise Ostafr. 111. p. 325.

1. PARARHADINHA MELANOGASTER Boettg. t. c. p. 326.

Madagascar (Nossi Be).

9, HETEROLIODON.

Boettg. in Voeltzk. Reise Ostafr. 111. p. 313.

1. HETEROLIODON ToRQUATUS Boettg. 1]. c. pl. xxvi. fig. 4.

Madagascar.

10. LioHETERODON.

Dum. & Bibr. Erp. Gén., Atlas, pl. Ixix.; Bouleng. Cat. Sn. i.

p. 268.

Three species :—

Scales in 23 rows; ventrals 209-216 ; praefrontals separating

internasals from frontal.....0.....0.0...c cece Le. madagascariensis.

Scales in 23 rows; ventrals 191-194; internasals in contact

mibhinrontallle Aue Naoeieo ee senn vache nce sat teases cen ean ae el cer ee LL. geayi.

Scales in 21 rows; ventrals 159-170; internasals in contact

wath tirontall fe. slfcch. ctusnte tee teeen ts tes eee En LL. modestus.

1. LIOHETERODON MADAGASCARIENSIS.

Heterodon madagascariensis Dum. & Bibr. op. cit. vii. p. 776,

ol, Ikabe

Lioheterodon madagascariensis Bouleng. t. c. p. 269.

Madagascar.

2. LioHETERODON GEAYI Mocquard, Bull. Mus. Paris, 1905,

[Os es 7

Lioheterodon voeltzkovii Boettg. in Voeltzk. Reise Ostafr. iii.

p. 313, pl. xxvin.

Madagascar.

3. LIOHETERODON MODESTUS.

Heterodon modestus Giinth. Ann. & Mag. N. H. (3) xii. 1863,

p. 396.

Lioheterodon modestus Bouleng. t. c. p. 269.

Madagascar.

“SNAKES OF MADAGASCAR. Ba

11. MrcropisrHopon.

Mocquard, CR. Soc. Philom. 1894, no. 17, p. 7.

1. MrcropistHopon ocHRACEUS Mocquard, |. ¢. p. 8.

Madagascar (Nossi Be).

12. LycoGNATHOPHIs.

Bouleng. Cat. Sn. i. p. 317.

1. LycoGNaATHOPHIS SECHELLENSIS.

Psammophis seychellensis Schleg. Phys. Serp. 11.

Lycognathophis sechellensis Bouleng. t. c. p. 317.

Seychelles.

[ao

13. Boopon.

Dum. & Bibr. Mém. Ac. Se. xxiii. 1853, p. 460; Bouleng. Cat.

Sls 6 TOs Ball

1. Boopon GEOMETRICUS.

Lycodon geometricus Schleg. Phys. Serp. i. p. 111.

Boodon geometricus Bouleng. t. c. p. 329.

Seychelles.

14, Lycopon.

Boie, Isis, 1827, p. 521; Bouleng. Cat. Sn. i. p. 348.

1. LycoDON AULICUS.

. Coluber aulicus Linn. Mus. Ad. Frid. i. p. 29, pl. xii. fig. 2.

Lycodon aulicus Bouleng. t. ec. p. 352.

Tutroduced in the Mascarene Islands.

Opisthoglypha.

Synopsis of the Genera.

J. Prefrontal not in contact with the upper labials; a loreal separating the nasal

from the preocular.

A. Pupil round.

Scales in 19 rows; tail moderate or short (subcaudals 31-76) ; anal

EMLINES: veiepsantea ae Saaee alse ee erak orem meee aval eeemnununtevneacsccry COULDSASs

Scales in 21 rows; tail long (subcaudals 121-175) ; anal divided ;

loreal once and a half to thrice as long as deep..................... Ithyeyphus.

B. Pupil vertically elliptic.

1. Body cylindrical.

Scales keeled, in 19 rows; snout ending in a long appendage ...... Langaha.

Scales keeled, in 25 rows; subcaudals single ..............0........ <Alluaudina.

378 MR. G. A. BOULENGER ON THE

Scales smooth, in 25 to 29 rows; eye separated from the labials by

suboculars Eteirodipsas.

2. Body compressed; scales smooth, in 17 to 25 rows.

Stroubmounded caswensscaceecnmeceeee sss ee eee ae nee eee recast auaerscceein MO CCOMIUISS

Snout much depressed, broad, truncate .........0....:ececee eee eeeeeeeee se =Lycodryas.

Il. Prefrontal in contact with upper labials, separating the

loreal from the preocular; nasal entire or semidivided ;

RENE MY NZ ASKOKSldegacdssacen shc05dca9 000 360 ono davadsaee sodeadespodasdence | JURA IGS

1. GEopIPsAs.

Bouleng. Cat. Sn. 111. p. 32.

Two species :—

A single loreal; ventrals 172-189; subcaudals 55-76............... G. infralineata.

Two superposed loreals; ventrals 187; subcaudals 31 ............ G@. bowlengeri.

1. GEODIPSAS INFRALINEATA.

Tachymenis infralineatus Giinth. Ann. & Mag. N. H. (5) ix.

1882, p. 265.

Geodipsas infralineata Bouleng. 1. c. pl. iii. fig. 1.

Madagascar.

2. GEODIPSAS BOULENGERI.

Tachymenis boulengeri Peracea, Boll. Mus. Torin. vu. 1892,

TO, IEPA 05 Bs

: g e.

Geodipsas boulengeri Bouleng. |. c.

Madagascar.

2. ITHYCYPHUS.

Giinth. Ann. & Mag. N. H. (4) xi. 1873, p. 374; Bouleng. Cat.

Sn. il. p. 34.

Two species :—

Preocular in contact with frontal; loreal twice to thrice as long as

dcep)smtihreepostocularsi. .9--se eee: a eeeee reee rene eesseeee: ese meen OuOud.

Preocular not reaching frontal; loreal once and a half to twice as

long as deep ; two (rarely three) postoculars.......................... I. miniatus.

1. IrHYCYPHUS GOUDOTI.

Herpetodryas goudoti Schleg. Phys. Serp. i. p. 187.

Ithycyphus goudott Bouleng. |. c.

Madagascar.

2. ITHYCYPHUS MINIATUS.

Coluber miniatus Schleg. t. c. p. 148.

Tthycyphus miniatus Bouleng. t. ¢. p. 35-

Madagascar ; Comoro Islands.

> >

SNAKES OF MADAGASCAR. 379

3. LANGABA.

Bruguiére, Journ. de Phys. xxiv. 1784, p. 132; Bouleng. Cat,

Sn. ili. p. 35.

Synopsis of the Species.

I. A single supraocular; rostral appendage pointed.

Rostral appendage at least twice as long as snout, ensiform, not

(STREP MIG le eens Sco cho nbboc Suobas SpeneS coq Onenes oae Roe eee ee ence aconoe Pl RaU eCUHGR

Rostral appendage once and a half to once and two-thirds as long

as snout, tapering to a sharp point, and serrated above at the

ETC ees esc ae eres MRE Aion Rise ea eee Pamabermed eae

Rostral appendage not more than once anda half as long as snout,

Sermtedsabove andubeneat bere erss- pee. aeee-e en re ceeemeeeeseese- anne lan CRISLORG CULE.

II. Three supraoculars, in addition to an erect horn-like scale

above the eye; rostral appendage not much longer than

snout, very obtuse, grooved beneath ...... BRP ast secrete Be TL. alluandi.

1. LaAncAHA- NAsuTA Shaw, Nat. Misc. xxii. pl. cmlxviii.;

Bouleng. t.c. p. 36.

Madagascar.

2. LANGAHA INTERMEDIA Bouleng. Ann. & Mag. N. H. (6) i.

IS88ip; LOS; pl v. fig. 6 ;"and t. ep. 37.

Madagascar.

3. LANGAHA CRISTA-GALLI Dum. & Bibr. Erp. Gén. vii. p. 806,

pl. lxxi.; Bouleng. 1. c.

Madagascar.

4, LANGAHA ALLUAUDI Mocquard, Bull. Mus. Paris, 1901,

p. 293.

Madagascar.

4, ALLUAUDINA.

Mocquard, CR. Soc. Philom, 1894, no. 17, p. 9; Bouleng. Cat.

Sn. 11. p. 38.

1. ALLUAUDINA BELLYI Mocquard, |. c.; Bouleng. 1. ¢.

Madagascar.

5. ETETRODIPSAS.

Jan, Hlenco sist. Ofid. p. 105; Bouleng. Cat. Sn. iii. p. 38.

1. ErErRODIPSAS COLUBRINA.

Dipsas colubrina Schleg. Phys. Serp. ii. p. 273.

Lteirodipsas colubrina Bouleng. t. c. p. 39.

Madagascar.

Proc. Zoor. Soc.—1915, No. XX VII. 27

380 MR. G. A. BOULENGER ON THE

6. STENOPHIS.

Bouleng. Cat. Sn. ii. p. 39.

Synopsis of the Species.

I. Scales in 17 or 19 rows; internasals much shorter than the prefrontals.

A. Scales in 17 rows; ventrals 187; subcaudals 67 pairs... S. guenthert.

B. Scales in 17 rows; ventrals 228-276.

Posterior chin-shields a little shorter than the anterior and in

contact with each other; ventrals 229-248; subcaudals

103-125, all or greater part In pairs .. . 8S. granuliceps.

Posterior Ghing shields longer than the anterior and in contact

with each other ; ventrals 228; subcaudals 110, greater

part in pairs ....... . SS. inornatus.

Posterior chin-shields shorter than the ‘anterior and ‘separated

by scales; ventrals 255-276; subcaudals 93-116, all or

GAEAIGIP FORTRY SMEG Goocagupsccansocen savonoas.scenpanavoodceoonaecaaua So GAMMA

C. Scales in 19 rows; ventrals 243; subcandals 126, single. S. maculatus.

II. Seales in 21 to 25 rows.

A. Internasals much shorter than the prefrontals; scales

in 21 rows; ventrals 173; subcaudals 157 pairs......... S. longicauda.

B. Internasals nearly as long as, or a little longer than the prefrontals.

1. Loreal separated from the eye by the preocular; subcaudals 152-159,

single.

Posterior chin-shields large and in contact with each other;

scales in 21 or 23 rows; ventrals 225-2386 ...........:......... S. arctifasciatus.

Posterior chin-shields very "small or absent ; ; scales in 23 or 25

TRONS WelmneAls CNP) as coacboosncnscnsboocbenbe sonebSenousueences [Sb GUMRACHDOIIS.

2. Loreal entering the eye; scales in 23 rows; ventrals

226 ; subcaudals 106 pairs ........... cece, S. betsileanus.

1. STENOPHIS GUENTHERI Bouleng. t. c. p. 40, pl. iv. fig

Madagascar.

2. STENOPHIS GRANULICEPS.

Dipsas (Heterurus) gaimardi, var. granuliceps Boettg. Abh.

Senck. Ges. xi. 1877, p. 14, pl. i. fig. 3.

Stenophis granuliceps Bouleng. t. ¢. p. 41.

Madagascar.

3. STENOPHIS INORNATUS Bouleng. t. c. p. 42.

Madagascar.

A, SrENOPHIs GAIMARDIL.

Dipsas guimardit Schleg. Phys. Serp. ii. p. 293.

Stenophis gaimardw Bouleng. t. c. p. 42.

Madagascar; Comoro Islands.

5. STENOPHIS MACULATUS.

Dipsadoboa maculata Giinth. Cat. Col. Sn. p. 183.

Stenophis maculatus Bouleng. t. c. p. 43, pl. iv. fig. 2.

Madagascar (2).

SNAKES OF MADAGASCAR. S81

6. STENOPHIS LONGICAUDA Boettg. in Voeltzk. Reise Qstafr. iii.

p..o15) pk sev tes 7:

Madagascar.

7. STENOPHIS ARCTIFASCIATUS.

Heterurus arctifasciatus Dum. & Bibr. Erp. Gen. vii. p. 1176.

Stenophis arctifasciatus Bouleng. t. c. p. 43.

Madagascar.

§. SrENOPHIS VARIABILIS Bouleng, t. c. p. 48, pl. iv. fig. 3.

Madagascar.

9. SPENOPHIS BETSILEANUS.

Dipsas betsileana Giinth. Ann. & Mag. N. H. (5) vi. 1880,

p. 238.

Stenophis betsileanus Bouleng., t. ¢. p. 44, pl. iv. fig. 4.

Madagascar.

7. LiycopRYAS.

Giinth. Ann. & Mag. N. H. (5) iii. 1879, p. 48; Bouleng. Cat.

Sn. 11. p. 44.

1. LycopRYAS SANCTI-JOHANNIS Giinth. 1. e.; Bouleng. t. c.

p. 45, pl. iii. fig. 2.

Comoro Islands.

8. Mrmopuis.

Giinth. Ann. & Mag. N. H., (4) i. 1868, p. 421; Bouleng. Cat.

Sn. il. p. 171.

1. MimorHiIs MAHFALENSIS.

Psammophis mahfalensis Grandid. Rey. et Mag. Zool. xix.

1867, p. 234,

Mimophis makfalensis Bouleng, 1. e.

Madagascar.

Proteroglypha.

(Sea-Snakes, with oar-shaped, compressed tail.)

Two genera :—

No distinct ventrals ; frontal at least as long as snout ...............-:- HAydrus.

Ventrals distinct, but very small; frontal shorter than its distance

ROWER OF SHUG | Sy assosoush son codoutosd sdb oorprcoce dos seocebe-easarmnnanaes | 2 ODOUR

Die

2 ON THE SNAKES OF MADAGASCAR,

1. Hyprus.

Schneid. Hist. Amph. i. p. 233; Bouleng. Cat. Sn. iil. p. 266.

1. Hyprus PLATURUS.

Anguis platura Linn. 8. N. i. p. 391.

Hydrus platurus Bouleng. t. c. p. 267.

Indian and Pacific Oceans.

2. KNHYDRINA.

Gray, Cat. Sn. p. 47; Bouleng. Cat. Sn. i, p. 302.

1. ENHYDRINA VALAKADIEN.

Hydrus valakadyn Boie, Isis, 1827, p- 904,

Enhydrina valakadien Bouleng. 1. c.

Tndian Ocean, eastwards to Papuasia.

ON A PARASITIC GAMASID MITE. 383

29. On a Blood-sucking Gamasid Mite (Jchoronyssus ser-

pentium, sp. n.?), parasitic on Couper’s Snake. By

Sranuey Hirst, F.Z.S8.*

[Received April 30, 1915: Read May 25, 1915.]

(Text-figures 1 & 2.)

It is quite probable that the species dealt with below is identical

with that to which Gervais gave the name Dermanyssus natricis

in the year 1844, but as there is some doubt on this point, I am

describing it under a new name. Gervais says very little about

the structure of his species, and the only account of it which is

at all complete is that written by Paul Mégnin in 1884, who

created a new genus (Ophionyssus) for it. References to several

earlier papers and works in which this parasitic mite is mentioned

are given in Mégnin’s paper. Unfortunately he makes a serious

mistake in his description, stating that there is no ventral plate

(plastron) on the lower surface of the body, and that the

‘‘ oviducte” has the form of a longitudinal sht. It is quite clear

that he mistook the narrow genito-ventral plate for the genital

opening itself. In reality, the genital aperture is quite normal

in appearance, being transverse and protected by the usual deli-

cate chitinous flap or operculum, which is joined posteriorly to

the genito-ventral plate. Ophionyssus natricis is mentioned by

G. Canestrini in his ‘ Prospetto dell’ Acarofauna Italiana,’ and

also by Prof. Antonio Berlese in his well-known work on the

Acari, etc. of Italy; but these authors did not have the oppor-

tunity of examining specimens, and the latter expresses doubt as

to the validity of the genus, rightly considering it as probably

identical with Leiognathus Can. | =Ichoronyssus Kolenati].

It may be of interest to note that, in having two dorsal shields

and also minute intermediate platelets, the adult females described

below present a strong resemblance to the protonymph stage

of certain other species of Jchoronyssus (for instance, to that of

I, bacoti mihi).

Genus IcHoronyssus Kolenati.

Dermanyssus (ad part.) Gervais, in Ins. Apt. ili. p. 223 (1844) ;

Ichoronyssus Kolenati, Wien. ent. Monatschr. 11. p. 5 (1858); id.

Sitz. K. Akad. Wiss. Wien, xxxv. p. 173 (1859); Ophionyssus

Mégnin, Bull. Soc. Zool. France, p. 109 (1884); Ophionyssus+

Leiognathus G. Canestrini, in Prospet. Acarof. Ital. part 1. p. 121

(1885) ; Liponyssus of some recent authors, but probably not

that of Kolenati.

* Published by permission of the Trustees of the British Museum.

384 MR. STANLEY HIRST ON A

IcHORONYSSUS SERPENTIUM, sp. n. ?

©. Body long oval in shape. Dorsal surface furnished with

numerous hairs, which are not very long and are distinctly

curved; apparently the fine little offshoot or hair present near

the end of the hairs of certain other species of Jchoronyssus (for

instance, on those of J. bacotz) does not occur on the hairs of this

Text-figure |.

Fehoronyssus serpentium, &. Ventral view.

species. There are two dorsal shields, the anterior one being

comparatively large (length -3 mm., width -27 mm.), but the

posterior one, which is situated far back near the hinder end of

the dorsal surface, is very small and inconspicuous. Outline of

anterior shield shaped almost like that of a lemon; twenty hairs

PARASITIC GAMASID MITE. 385

are present on its surface, six pairs practically forming a longi-

tudinal series running down the middle and four other hairs

being placed on each side, three of these lateral hairs being

marginal, The minute posterior shield is not provided with

any hairs, but one or more pairs of minute punctations, which

may possibly represent the sockets of hairs, occur on it.

On each side of the dorsal surface, a little behind the anterior

shield, there are two distinct but very minute platelets, and

they are followed posteriorly by about four very inconspicuous

Text-figure 2.

Ichoronyssus serpentium, 2. Dorsal view.

(obsolete) linear platelets, arranged in a longitudinal series.

Numerous hairs are present on the posterior part of the

ventral surface, but they are shorter than those on the dorsal

surface. Sternal plate trapezoidal in shape, being much wider

than long and furnished with only two pairs of hairs. Genital

aperture normal in situation and appearance ; it is protected by

the usual thin membranous operculum. Genito-ventral plate long

and very narrow, the hinder end being sharply pointed. Anal

386 ON A PARASITIC GAMASID MITE.

plate pear-shaped, and it has the three usual hairs on its surface.

Peritreme slender and rather short; it reaches a little further

forwards than the coxa of the third leg. Fingers of chelicera

short, and apparently without any trace of teeth. Legs. First

leg the longest, and the fourth leg also long, the legs of the

second and third pairs being shorter. Ventral surface of coxee

without any strong spurs, only fine sete or hairs being present.

There is, however, a very slight projecting spinule at the distal

end of the inner surface of the coxa of the first leg.

Length *9 ram.

Colour (an spirit) reddish brown.

Material. Five female examples found on a Couper’s Snake in

the London Zoological Gardens (May 5th, 1909).

ON EXTERNAL CHARACTERS OF THE PARADOXURINA. 387

30. On the Feet and Glands and other External Characters

of the Paradoxurine Genera Paradoxurus, Arctictis,

Arctogalidia, and Nandinia. By R. 1. Pococg, F.R.S.,

F.1L.S., F.Z.8., Curator of Mammals.

[Received May 11, 1915: Read June 8, 1915.]

(Text-figures 1-10.)

INDEX. Page

ISG Oe JEGIAMEOTENIPOS cccanh cocacd sbsanosechodon one cobscn senkise seams » GIT

Bare AT CLUCHUS Mn eenned ee Sena ee cence ae cascear ea. OOO

Bi ARCLOG EL CO LC meet eerste eeacsa atens Sorte OOO,

5 NO DOH OUR concanee 392

Comparison between the Pinadlosumine cml Winwadina fe pes

Ok fectv=n eyes: SERED GUC OREOR ESTEE OR GL |

Rhinarium and Fibre at Bp yredonenines RAR Re eo Oy

Ear of Paradoxurines and Viverrinues . sae anos.) Bes

Perfume-gland and external genitalia oe Paradenwe WE so, 4houl

3 = 5 bp AUACGUCHES sotecaceo 4.06

= “s ts F Arctogalidia ... 407

% 6 5p INandinig =... ... 409

General conclusions with regard to the perfume-glands ... 411

This paper, the result of researches carried on at intervals for

several years in the Society’s Prosectorial Department, may be

regarded as a continuation of the one dealing with the genera

of Viverrine which was published in the ‘ Proceedings’ for

March 1915 (pp. 131-149). Its subject-matter is treated on the

same general linesas those therein adopted, and its main purpose

is to show first that the genera discussed, hitherto diagnosed

mainly by cranial and dental characters, may be equally well,

perhaps better, distinguished by the cutaneous features examined ;

and secondly, that these features fully justify the conclusion,

hinted at but not adopted by Mivart, that the Viverrine and

Paradoxurine genera should be relegated to distinct subfamilies

of the Viverride.

The genus Paradoxurus, including the species referred to

Paguma and Macrogalidia, ranges from India and Ceylon to the

Philippine Islands and Celebes. <Arctictis extends from the

eastern Himalayas to Borneo, and Arctogalidia, with the same

eastern limit, reaches Assam. Wandinia, on the contrary, i

restricted to the forests of tropical Africa.

The Feet of Paradoxurus.

Of this genus I have examined examples of the three species

P. larvatus trom Szechuen, P. niger from India, and P. hermaphro-

ditus from Singapore. According to Gray’s nomenclature, which

there is a tendency at the present time to revive, larvatus should

be referred to the genus Paguma; but in this paper I propose,

388 MR. R. I. POCOCK ON EXTERNAL

without prejudice, to follow Blanford and most modern authors

in regarding Paguwma as a synonym of Paradoxurus.

The fore foot of P. larvatus is broad; the digits are capable of

considerable distension and are webbed up to the proximal ends

of the digital pads; the underside of the webs is smooth, save for

the presence of four patches of short hair near the distal margin

of each. The pollex is well developed and lies close to the second

digit. The plantar pad is large and wide owing to the size of its

pollical lobe, which approximately equals in dimensions any one of

the three main lobes corresponding to the intervals between the

Text-figure 1.

Paradoxurus larvatus.

A. Left fore foot, digits fully stretched.

B. Left hind foot, _,, *

four principal digits. The lobes are well defined by grooves.

The double carpal pad, separated by a deep naked crease from

the plantar pad, is at least as wide and long as the latter in-

cluding its pollical lobe. The external lobe of the carpal pad is

about twice as large as the internal and is defined from it by a

deep groove. These carpal pads occupy practically the whole

width of the paw behind the plantar pad and, like the latter and

the digital pads, are covered with coarsely granular or scale-like

integument.

The claws are protected by rudimentary skin-lobes and are

CHARACTERS OF THE PARADOXURINA, 389

retractile in the sense that the terminal phalanx can be drawn

back so as to lie along the outer side of the penultimate phalanx

as in most of the Viverride and the Felide.

The hind foot resembles the fore foot in its main features, but

is narrower and the digital pads of the third and fourth digits

are united at the base; behind their point of junction there is a

small triangular patch of short hair. Similar patches of hair are

present on the underside of the proximal half of the webs, as in

the fore foot. The hallux is shorter than the pollex, but the

hallueal lobe is large and adds considerably to the width of the

plantar pad, the elements of which are well defined. Behind the

plantar pad the greater part of the underside of the metatarsus

is naked; the naked area, narrowing posteriorly or proximally,

reaches to within about half an inch of the heel, which is

covered with hair, This naked area exhibits a median depression

which is narrower distally close to the plantar pad than

proximally towards the heel, and is bounded at the sides nearly

throughout its length by two thick ridges of integument, the

metatarsal pads. The external of these, narrower but a little

longer than the internal, is continuous distally with the external

lateral lobe of the plantar pad ; the internal is similarly continuous

with the hallucal lobe of that pad, In the specimen here described,

the depression between these pads or ridges 1s thickly covered

with horny sub-spiniform papille.

The feet of Paradoxurus hermaphroditus and P. niger show no

differences of moment from those of P. larvatus. The underside

of the webs, however, is naked; and the example of P. niger

examined exhibited no sharp horny papille in the depression

between the metatarsal pads of the hind foot.

The Feet of Arctictis.

The fore foot resembles in all essential respects that of Para-

doxurus larvatus, but the web connecting the digits is naked

beneath, and the two elements of the carpal pad are not so sharply

defined from one another and from the plantar pad.

The hind foot is also like that of Paradoxurus in most respects,

particularly, be it noted, in the fusion of the digital pads of the

third and fourth digits proximally. Here also there is no hair on

the underside of the interdigital webs, the division between the

plantar and metatarsal pads is ill-defined, and the metatarsal area

itself is not sharply differentiated into a median depression and

lateral elongated pads. On the contrary, it is comparatively flat

and the naked area extends right back to the heel. In one

example, a full-sized but young male, the skin of the underside

of the heel is covered with a mat of horny, pointed papillee.

No such mat, however, is present on the heel of a young female,

and, judging from the skins in the British Museum, its develop-

ment is variable, the seulpturing being sometimes papillate,

sometimes squamous.

390 MR. R. I. POCOCK ON EXTERNAL

The claws are sharp, strongly curved, and retractile, but are

unguarded by lobes of skin.

In the nakedness of the heel, Arctictis, as has been noticed by

Mivart and others, differs from all other genera of Paradoxurines.

Text-figure 2.

re DONO e

rcenauiite

Gs Lf

YZ Gi re

tie. Nii LZ

GY a j <2

Arctictis binturong.

A. Left fore foot, digits partially stretched.

B. Left hind foot, digits not stretched.

This may be an adaptation to a more thoroughly arboreal life—

a conclusion supported by the prehensile power of the tail *; but

I cannot find any record of the proximal fusion of the pads of the

third and fourth digits. This character is very suggestive of

affinity between Arctictis and Paradoxurus.

The Feet of Arctogalidia.

The feet of an example from Sarawak resemble in a general

way those of Paradoxurus, but differ in one or two interesting

particulars, especially in being longer, narrower, and Jess fully

webbed. In the fore paw the digital and plantar pads are well

* About half an inch of the end of the tail was naked and formed a conical point

to that organ calculated to enhance its grasping capacity.

CHARACTERS OF THE PARADOXURINA. 391

Text-figure 3.

Arctogalidia, from Sarawak.

A. Left fore foot, digits fully stretched.

B. Left hind foot, es

Nandinia binotata.

C. Left fore foot, digits fully stretched.

D. Left hind foot, 4 3

392 MR. R. I. POCOCK ON EXTERNAL

developed in the matter of thickness, but are covered with smooth

integument, and the four lobes of the plantar pad are separated

by shallow grooves and are therefore somewhat ill-defined. he

area between the digital and plantar pads is quite hairless and

the edge of the webbing between the Ist and 2nd, 2nd and 3rd,

and Ath and 5th digits, and to a less degree that between the 3r d

and 4th digits, 1s more deeply emarginate than in Paradoxurus.

The anterior and lateral borders of the plantar pad form a

narrower curve than in that genus, the pollical lobe especially

being set farther back with reference to the median lobe. The

carpal pads are much longer and narrower, but are as wide

throughout as the plantar pad. The claws are short and strongly

curved and not protected by skin-lobes.

The hind foot differs from that of Paradoxurus in characters

similar to those mentioned in connection with the fore foot, but

one additional difference to be noticed is that the pads of the 3rd

and 4th digits are not fused but are separated by a measurable

extent of webbing. The naked area on the metatarsus is of about

the same extent as in Paradoxurus, the heel being thickly hairy

as in that genus; and in the skins examined the area between

the two smooth lateral ridges, or metatarsal pads, is covered with

coarsely squamous integument, the pads themselves, like the

plantar and digital pads, being smooth as in the fore foot.

Lhe Feet of Nandinia.

The feet of this genus have only been briefly described pre-

viously, so far as I have ascertained. Mivart (P.Z.S. 1882

p. 170) says that the tarsus and metatarsus are “‘ about as bald as

in Paradozurus,’ which is true; but Lydekker’s statement that

the tarsus 1s partially bald as in Hemigalus is not in accord with

the facts (Lloyd’s Nat. Hist.: Cats etc., p. 228, 1896).

The fore foot resembles that of Paradoxurus larvatus in the

development of the webs at least to the proximal ends of the

digital pads, in the extent to which the toes are capable of

separation, and in the confluence of the plantar and carpal pads

to form a single mass, of which the component elements are

defined merely by grooves. These pads, however, are longer as

compared with their width than in Paradowurus, and in the more

backward position of the pollical lobe and the closer curvature of

the distal margin of the plantar pad resemblance may be seen to

Arctogalidia. Ue very narrow area of skin, sometimes giving off

short streaks towards the digital pads of the 2nd, 3rd, 4th, and

5th digits, is naked, but otherwise the whole of the area between

the plantar and digital pads is thickly covered with velvety hair

as in Genetta and Viverra. In this respect the feet differ from

those of Paradoxurus *, Arctictis, and Arctogalidia. The underside

of the pollex is, moreover, quite naked, the digital pad of this

* Hodgson, however, described this area in Paradoxurus lanigerus as hairy; but

whether the hair was dey eloped to the extent seen in Nandinéa or in Paradowurus

larvatus does not appear.

CHARACTERS OF THE PARADOXURINAE. 393

digit, which is considerably shorter than in Paradoxurus and a

little shorter than in Arctogalidia, being connected with the

pollical lobe of the plantar pad by a bare strip of integument.

The claws are retractile but are not guarded by small skin-

lobes as in Genetta. The pads are in the main quite smooth, but

the central depression behind the plantar pad is somewhat

coarsely sculptured.

The hind foot, so far as the webbing and the structure of the

plantar pad are concerned, resembles and differs from that of the

other genera described in this paper in the same respects as those

mentioned in connection with the fore foot. The digital pads,

however, of the third and fourth digits are widely separated,

even more so than in Arctoyalidia, and resemble in this respect

the homologous pads of Genetta and other Viverrines. The whole

of the posterior area of the naked metatarsal space is coarsely

ridged transversely, and this ridging is replaced in the elongated

depression behind the plantar pad by coarse polygonal sculp-

turing. This depression is bounded on each side by a thick,

elongated, metatarsal pad, sculptured internally, but these pads

are not so long as in Paradoxurus and Arctogalidia*.

The characters of the feet of the four genera of Paradoxurines

above described may be systematically epitomised as follows :—

a. Pads of 3rd and 4th digits of hind foot proximally united in

the middle line; fore paws broad, their plantar and carpal

pads combined only a little longer than wide; pads of fore

and hind feet with coarse tessellated sculpturing.

a’. Sole of hind toot not naked to heel .............................. Paradoxurus.

b’. Sole of hind foot naked to heel........................0..0........ Arctictis.

6. Pads of 3rd and 4th digits of hind foot sevarated ; fore paws

narrower, their plantar and carpal pads combined much

longer than wide; pads of fore and hind feet smooth or

nearly so; {heel of hind foot hairy as in Paradoxurus].

a?. Area between plantar and digital pads smooth............... Arctogalidia.

62. Area between plantar and digital pads, except of digit 1,

DEG kelives h aire vieetesane semee siiets..-8 watery lociaa eek ees cm uaaenaeies Nandinia.

The tessellated sculpturing of the pads observed in Arctictis

and the species of Paradoxurus examined must be verified for

other species of the latter genus, before it can be definitely

regarded as a generic feature. The fusion of the pads of the

3rd and 4th digits of the hind feet in these two genera—

obviously a specialised feature—may be regarded as evidence of

affinity between them; but the separation of these same pads in

Nandinia and Arctogalidia is a case of the mutual inheritance

of a primitive character, and cannot be considered as evidence of

relationship.

* From the foregoing account of the hind feet of Paradoxurus, Arctogalidia, and

Nandinia it will be seen that Mivart’s description of the tarsus as “ halt-bald ” in

these genera is untrue, since by the term tarsus ke meant the whole area between

the plantar pad and the heel (P. Z.S. 1892, p. 206). He also included Hemigalus

(Hemigalea) in the category of genera with “half bald” tarsus, although his own

tigure of the hind foot of this genus (op. cit. p. 166) shows correctly that the greater

part of the sole behind the plantar pad is covered with hair.

394 MR. R. I. POCOCK ON EXTERNAL

Comparison between the Paradoxurine and Viverrine

types of feet.

The feet of the Paradoxurine genera above described, and those

of the Viverrine genera described in my previous paper (P. Z. 58.

1915, pp. 182-140), may be briefly compared as follows :—

Viverrine.—The pollical and hallucal elements of the plantar

pads are either suppressed or small, and when present lie

altogether behind the internal lateral lobe of the plantar

pads so as to contribute nothing to the width of the latter.

The carpal pads whether single or double are much shorter

and narrower than the plantar pad, occupy only a small

part of the underside of the carpo-metacarpal area, and are

separated from the plantar pad by a tolerably long space,

of which the median portion at least is covered with hair.

The underside of the metatarsus is for the most part covered

with hair; when the metatarsal pads persist, they are |

reduced to a small bilobed pad some distance away from

the plantar pad (O%vettictis), or to two narrow median ridges

of integument in contact throughout the greater part of

their length, slightly separated towards the heel and more

strongly divergent inferiorly where they extend to right and

left to meet the postero-lateral angles of the plantar pad,

leaving a hairy space between (renetia).

Paradoxurine.—The pollical and hallucal elements of the

plantar pads are large and comparable in size to the three

remaining lobes of this pad individually, thus adding con-

siderably to its width. The carpal pads are long and wide,

occupy nearly the whole of the width of the underside of

the carpo-metacarpal area, and conjointly equal or approxi-

mately equal the plantar pad in area; they are defined from

it by a transverse groove which expands mesially into a

depression, but this depression is never hairy. The meta-

tarsal area is naked throughout the greater part of its

length and width; when the metatarsal pads are retained

they form two thick ridges of integument separated by a

wide depressed area.

From this it is clear that the differences between the feet of

these two groups are considerable. The differences in the case

of the Viverrine have arisen, as I have already pointed out

(P. Z.S. 1915, p. 139), by the suppression or reduction in size of

the pollical and hallucal elements of the plantar pads, by the

reduction in size of the carpal pads, the growth of hair between

them and the plantar pad, and by the suppression or reduction

in size of the metatarsal pads and their replacement by normal

hairy integument. In the case of Genetta, which of all the

Viverrine has the least specialised feet, it 1s clear that the two

juxtaposed narrow ridges of skin extending along the middle

line of the metatarsus are the homologues of the two thick ridges

or pads, separated by a median depression, in Paradoxurus or

Arctogalidia.

CHARACTERS OF THE PARADOXURIN &. 395

There is one other genus of Viverride which ealls for attention

here because its feet belong to the Paradoxurine type, though

they differ in certain details “from the feet of the genera to which

that term is here restricted. This is the otter oie amphibious

Civet Cynogale, which Gray first of all made the type of a special

subfamily, afterwards raising it to family rank on characters

held by Mivart to be trivial for that purpose. Mivart regarded

Cynogale as an aberrant genus of his subfamily Viverrinse, which

embraced the Civets, Genets, Linsangs, Paradoxures, and others.

The gland, however, was unknown both to Gray and Mivart.

By this organ, as I have recently shown *, as well as by the

structure of the muzzle and of the rhinarium, Cynogale differs

causiderably from the genera both of the Viverrine and Para-

doxurine sections of Viverridz, and cannot be included in either,

as I define them, without unduly disturbing their homogeneity.

In classifying this genus, therefore, I revert to Gray’s original

idea and regard it as the representative of a special subfamily,

Cynogaline.

The Rhinarium and Vibrisse of Paradoxurines.

In Paradoxurus larvatus the rhinarium is large and prominent.

Viewed from the front its upper edge is markedly biconvex

owing to the depth of the median groove and the curvature of

the lateral angles. This groove extends uninterruptedly from a

point close to the posterior border of the upper surface, over the

anterior surface, where it is very deep, down to the edge of the

lip in the middle line. The infranarial portion is large and

extends laterally beyond the nostrils, where it curls up and nar-

rowly borders them externally throughout their length on the

upper side. Its inferior edge is horizontal with rounded angles.

Just beneath the narial orifice in front, this infranarial portion

is marked with a curved depression which runs downwards and

inwards towards the middle line. This groove appears to corre-

spond to the obliquely inclined infranarial edge of the rhinarium

in Canis and Felis, the area external to the groove being covered

with hair in those genera and other members of the families to

which they belong.

Viewed from above, the convexity of the anterior edge is

interrupted by a deep median angular notch—smaller in the

young—and its posterior edge is concave. The narial slits con-

verge inwards and backwards and are bordered externally by the

narrow naked strip which is continuous in front with the supero-

lateral angles of the large infranarial area. Asa result of the

backward “nm Inward ele ion of the narial slits, the upper

-field of the rhinarium is considerably wider in front than behind.

Finally, it may be added, the hairy portion of the upper lip

below the rhinarium is only about half the height of the anterior

surface of the rhinarium itself,

Comparing the rhinarium of Paradoxurus with those of the

* Ann. Mag. Nat. Hist. (8) xv. pp. 351-360, 1915.

Proc. Zoou. Soc.—1915, No. XXVIII, 28

396 MR. R. I. POCOCK ON EXTERNAL

Viverrine described in my previous paper, it clearly resembles

most closely that of Viverra zibetha, but its anterior and upper

surfaces are much more deeply srooved, its superior angles on

each side of the median notch are more prominent, and its upper

surface is more markedly biconvex.

In P. hermaphroditus the rhinarium is like that of P. larvatus

in all essential respects.

Text-figure 4.

ly 0 1

4, ns ny wy WY Wits ok »

‘1 )

tata ‘iLL

an il RR ep 14,

Rhinarium, seen from the front and above, of A. Nandinia; B. Arctogalidia ;

C. Paradoxurus larvatus; D. Arctictis; EH. Civettictis.

The rhinarium of Arctogalidia from the anterior aspect closely

resembles that of Paradoxurus, but is narrower as compared with

its height, and the lower edge of the infranarial portion slopes

more obliquely upwards. Tiasred from above, the median notch

is wider and somewhat deeper, and the angles that define it are

farther from the middle line and nearer the narial slits. These

CHARACTERS OF THE PARADOXURINE, 397

slits, moreover, are subparallel and only slightly convergent pos-

teriorly, so that the upper surface of the rhinarium is only a

little narrower behind than in front.

In Nandinia the rhinarium, seen from the front, is a little

narrower and higher than in Arctogalidia, its upper edge is less

markedly biconvex, owing to the sulcus being shallower. The

upper surface has a shallower anterior notch, and is a little

longer as compared with its width than in dr ctogalidia and is

quite as wide behind as in front. i

The rhinarium of dAretictis differs from those of the other

Paradoxurine genera here described in one or two points. The

median groove extends over the upper surface, but is sunk in a

much shallower depression. Hence the superior margin of the

anterior surface is not markedly biconvex, but is fairly evenly

convex from side to side, with only a small and shallow median

notch. Seen from above, this margin is similarly convex from

side to side, with a small median notch. The infranarial portion

in front is shallower and has a more evenly rounded inferior

border, and, when seen from above, the narial slits converge

posteriorly as in Paradoxurus.

In my deseription of the rhinaria of the Viverrine genera

Viverra, Civettictis, etc., I said :—* It is impossible to affirm the

existence of any absolute difference between the rhinaria of the

Viverrine collectively and of the Paradoxurine.” Confirmation

of this is supphed by a study of this organ in the Paradoxurine ;

for, although the rhinarium of Paradoxurus, Arctogalidia, and

Nandinia differs from that of the Viverrinze in being very deeply

suleate above and in front, the rhinarium of neal, a genus In

most particulars the least Viverrine of all the Paradloxnraimesy | 1s

much less deeply and widely sulcate, its supero-anterior margin

being evenly convex from side, to side with a quite small median

notch. The convex curvature of this border recalls that of the

rhinarium of the African Civet (Civettictis), although the median

groove of the rhinarium is deeper both above and in front than

in that genus. On the other hand, the rhinarium of the large

Indian Civet (Viverra zibetha), with its upper surface biconvex,

is more like the rhinarium of Paradoxwrus than is the rhinarium

of Arctictis. In the shape of this organ, therefore, the genera of

Viverrine and Paradoxurine intergrade.

The facial vibrissee may be briefly dismissed. The tufts are

without exception normal in number, the mystacials in particular

being long and rigid. Of the two genal tufts on each side, the

inferior is situated in a line With the corner of the mouth and

the superior a little higher up and posterior to it. The least

developed is the interramal, but it is always present and not far

behind the mandibular symphysis.

The Ear of Paradoxurines dnd Viverrines.

There appears to be no accepted terminology for the carti-

laginous ridges which strengthen and support the pinna of the

IR*

398 MR. R. I. POCOCK ON EXTERNAL

ear in mammals, and by interlocking help to close the meatus

when it is capable of being closed. Mivart described the pinna

of the Common Cat (‘The Cat,’ pp. 295-296, 1881) and of the

Genet (P.Z.S. 1892, p. 51, fig. 12), adopting for the several

parts names originally applied to the human ear. An entirely

different nomenclature was, however, proposed by Boas in 1912

(‘Ohrknorpel und fusseres ohr der Saugetiere,’ Kopenhagen).

In the following account I have attempted to show the corre-

spondence between these two systems and have, in the main,

followed that of Mivait as being more familiar and more in-

telligible, without wishing thereby to cast any reflection upon

the excellence of Boas’ work.

In Paradoxurus larvatus the cartilaginous ridges, with their

intervening fosse, which strengthen the lower portion of the

pinna round the auditory meatus, resemble those of the Cat and

Genet in essential features. Two ridges run obliquely upwards

and forwards in front of the inferior orifice of the meatus

(aditus inferior of Boas). The outer of these, the posteron 4 of

Boas, carries a low elevation called the traguws by Mivart. ‘The

inner, the antcron 6 of Boas, has a sinuous edge and runs higher

up the front of the ear than the outer or tragus-bearing ridge.

This inner ridge is called the post-tragus by Mivart in the case

of the Cat, but in his figure of the Genet’s ear it is marked

tragus. Two ridges similarly run obliquely upwards and _ back-

wards from the inferior orifice of the meatus. The outer of

these, the posteron 6 of Boas, is produced inferiorly into a large

angular process, the antitragus of Mivart, which fits into the

lower part of the fossa between the two anterior ridges when

the ear is closed. Above and within the outer ridge lies the

inner of the two posterior ridges, which is much softer and Jess

well developed than the others and shows a small swelling near

its lower end. This ridge, an integumental non-cartilaginous

structure, was not given a special name by Mivart, and was

merely described as a “weiche Falte” by Boas. Nevertheless

it appears to be a constant feature in the ears, at all events, of

the Canide, Felide, and Viverride.

The four ridges above described, with their intervening fosse,

form the anterior and posterior walls of the deep and spacious

fossa lying above the auditory meatus. This large fossa is defined

above by a transverse cartilaginous ridge, the plica principalis of

Boas and the swpratragus of Mivart, the anterior end of which

is overlapped by the inner of the two anterior ridges. ‘Towards

the middle of the ear it exhibits a marked swelling, and behind

this the ridge gradually fades away towards the inner side of the

inner of the two posterior ridges *.

* The cartilaginous thickening in the ear of the Carnivores, named posteron 4 by

Boas and tragus by Mivart, seems certainly to be the homologue of the “tragus ”

in the human ear. But the thickening named “antitragus” by Mivart and

posteron 6 by Boas is not, according to this latter author, the exact homologue of the

human antitragus. This well-developed structure in man is part of another carti-

Jaginous ridge, the posteron 5 of Boas, which is at most feebly developed in the

Carnivores, being merely represented by a weak ridge lying below and on the outer

CHARACTERS OF THE PARADOXURINS. 399

The posterior margin of the ear, nearly in a line with the

supratragus, is doubled to form a definite pocket, called the

* pouch” by Mivart and the ‘“‘ Korbchen ” or “‘ Tasche” by Boas.

For this I have proposed the name “ bursa.” The anterior flap

of this is continuous above and below with the rim of the pinna,

and its edge is deeply, widely, and angularly emarginate. The

posterior flap, on the contrary, is semilunar with a continuously

convex edge, the upper and lower ends of which are attached

behind the rim of the pinna, as in the Genet and, I believe, all

species of Felis.

Comparing the ear of Paradoxurus larvatus with that of the

three genera of Viverrines—Genetta, Viverricula, and Civettictis—

the following points may be noticed. In Genetta dongolana

(Somaliland) the bursa is formed as in P. larvatus, except that

the emargination of the anterior flap is nearly rectangular in the

former and obtusely angular in the latter. In Civettictis civetta

and Viverricula malaccensis, on the other hand, the anterior flap

is less deeply and more widely emarginate and the convex edge

of the posterior flap is continuous above and below with the rim

of the pinna, instead of rising behind it. Also in these two the

inner of the two anterior ridges carries a much larger process

overhanging the anterior end of the supratragus (antihelix or

plica principalis) than is to be seen in Genettw dongolana and

P. larvatus.

In Genetta dongolana and rubiginosa the tragus is more

markedly bilobed and the antitragus provided with a_ better

developed external ridge, giving rise to the “double” condition

described by Mivart in G. tigrina, than in P. larvatus. This

external ridge is very well formed in Viverricula, but is not

larger in Civettictis than in Paradoxurus larvatus.

Apart, however, from the bursa, the ears of the four species

are very much alike, and the recorded differences in the exact

shape of the ridges must be tested in the case of other species of

the genera before, in my opinion, it will be safe to attach

systematic importance to them. They may be due merely to

individual variation.

The ear of Paradoxurus hermaphroditus resembles that of

P. larvatus except that the anterior flap of the bursa is more

deeply and roundly emarginate and its angles, especially the

inferior angle, are more produced.

I did not examine in a fresh state the ears of Arctictis. The

only point of importance that could be made out on a dry skin

was the presence of the bursa and its resemblance to that of

Paradoxurus larvatus, in the origin of the upper end of the

side of the so-called antitragus. Mivart recorded the two ridges as a “ double

antitragus,” a quite intelligible view. On the other hand, Mivart does not seem to

have detected tnat the ridge he named the post-tragus (=anteron 6 of Boas) corre-

sponds to the basal portion of the ascending helix in man, and that the supra-tragus

(=plica principalis of Boas) is the homologue of part of the antihelix of human

anatomists. 5

400 MR. R. I. POCOCK ON EXTERNAL

Text-figure 5,

Base of pinna of left ear af A. Nandinia; B. Aretogalidia; C. Baradorurus

hermaphroditus; D, P. larvatus; E. Viverricula malaccensis; F. Civeétictis

civetta.

5., bursa; o., inferior orifice of meatus; s., superior ridge (=supratragus of

Mivart) ; ae,, external of the two anterior ridges (=¢ragus of Mivart) ; az., internal of

the two anterior ridges (=post-tragus of Mivart) ; pe., external of the two posterior

ridges (=antitragus of Mivart) ; pi., internal of the two posterior ridges; e, supple-

mentary ridge outside pe. in Vivervicula. :

CHARACTERS OF THE PARADOXURIN. 401

posterior flap from the back of the pinna and the continuity of

the rim of the pinna with the upper end of the anterior flap.

This flap, however, is much more widely and less deeply

emarginate than in P. larvatus.

In Arctogalidia the cartilaginous ridges of the ear resemble in

a general way those of Paradoxurus larvatus, but there is a

distinct though small ridge on the outer side of the antitragus.

The bursa, however, differs in one or two points. The margin

of its anterior rim is not so deeply or abruptly excised, and its

posterior rim is continuous above with the rim of the ear and

does not arise behind it as in that genus. Hence the orifice of

the bursa is more closed, and its posterior wall does not form

a flap freely movable on its base of attachment. In other words,

the bursa in Arctogalidia is more like that of Viverricula malac-

censis and Civettictis civetta than of Paradoxurus larvatus and

Genetta.

In Nandinia binotata the inner of the two anterior carti-

laginous ridges differs from that of Paradoaurus larvatus and of

Arctogalidia in the larger size and angular shape of its two

processes. As in Arctogalidia, the antitragus carries an external

ridge. The bursa resembles that of Paradowurus, Arctictis, and

Genetta in the origin of its posterior flap from the back of the

pinna, but the angular excision of its anterior flap is shallower

than in Paradowurus, but deeper than in Arctictis.

From the foregoing account it will, I think, be clear that it is

impossible to make use of the ears in differentiating the Viverrine

from the Paradoxurine. Nevertheless, within the limits of these

two subfamilies the structure of these organs, and especially of

the bursa, will probably be found useful for distinguishing

genera.

I am quite unable to surmise what meaning is to be attached

to the variation in the structure of the bursa, by which the

genera above discussed may be grouped as follows :—

a. Posterior flap of bursa arising above behind the rim of the

ENO Gee ooceos Genetta, Paradovurus, Arctictis, Nandinia.

6. Posterior flap of bursa continuous above with the rim of the

[OUTINGS shes seoec Viverricula, Civettictis, Arctogalidia.

Since the condition of the bursa found in Canide, Felide, and

some of the Arctoid Carnivora is the same as that described

under heading a, this condition is probably the primitive one.

Probably, also, the condition described under heading 6 is the

beginning of the suppression of the bursa, a process which is

completed in some Adluroid and some Arctoid Carnivores.

The Perfume-gland and External Genitalia of Paradoxurus.

One of the earliest descriptions of the gland and genitalia of

Paradoxurus was published under the name Platychista pallasii

(=Paradoxurus hermaphroditus) by Otto (Nova Acta Acad. Leop.-

Car. xvil. p. 1089, pl. Ixxili, 1835). Except in the structure of

402 MR. R. I. POCOCK ON EXTERNAL

Text-figure 6.

Aa pie eel BBX , TN

vat / ; Prot

ASA Aw W/GE

“Ve

/ N

Per ted

oer i

7B NN Wp

G ‘i A Ze

A Lie

AS Wy

C “

Paradoxurus larvatus.

A. Gland and external genitalia of male. g/., glandular area with the position of

the right gland dotted in, the median depression not represented; p., glans

penis ; se., scrotum.

(Continued at foot of neat page.)

CHARACTERS OF THE PARADOXURINA. 403

the penis and its precise relations to the gland, the figure

accompanying the description agrees with my own observations.

A year later Hodgson (Asiatic Res. xix. p. 77, 1836) described

the perfume-glands of three species referred to Paradoxurus,

namely, hirsutus, nipalensis, and lanigerws. According to Blan-

ford the name hirsutus was applied to the two species now known

as niger and hermaphroditus; nipalensis is a synonym of gray,

but the species named lanigerus can only be assigned to the genus

with hesitation. By the system of classification now in use, niger

and hermaphroditus belong to the genus Paradowurus and grayi

to the genus Paguma; and I do not doubt that lanigerus is also

a Paguina.

Hodgson detected no difference between the glands of the

species he examined ; and although he examined both males and

females, his description suggests that the glands in the two

sexes are alike in their relations to the vulva and penis. This,

however, is not the case, as will be explained, since the penis is

situated at the anterior end of the naked glandular area and the

vulva les near its centre.

Mivart’s description of the scent-glands as lying “‘ beneath the

surface of a valve-like antero-posteriorly directed cutaneous

inflection, more or less naked, and situated between the penis

and testes in the male and analogously in the female,” conveys

very little idea of their structure (P. Z. 8. 1882, p. 163).

Turner’s very brief description of the gland of P. niger

(=typus) as an oval, flat, naked space at the base of the prepuce,

although wanting in preciseness as to the situation of the gland,

is peent ite and intelligible (P. Z 8. 1849, p. 25).

The main portion of the gland in the male consists of a pair of

thickened ridges of skin or ‘labia extending between the scrotum

and the penis. These labia form the side-walls of a longitudinal

fossa which is everywhere perfectly smooth ; but the margin and

outer surfaces of the labia are covered with long hair. The

paired glandular thickening, which makes these labia, does not

extend so far forwards as the penis, but a rim of thickened skin,

naked internally, hairy externally, passes forwards in front of

them and encircles the base of the penis like a collar. The

narrow space between the collar and the penis is highly glandular,

Description of text-figure 6, continued.

B. Transverse section of the glandular area of the same; g/., gland of right side;

i., glandular space bordered by the two labia ; p., penis.

C. Gland and external genitalia of female. /., clitoris; J., left labium of gland

folded over glandular area ; @., anus.

D. Same as fig. C, with labia of gland (Z.) pulled apart to show vulva (v.) and

narrow labia of vulva (/v.). a@., anus; cl., clitoris.

E. Transverse section of same behind vulva. gl., gland; ¢., glandular space bounded

by the two labia; v., vagina.

404 MR. R. I. POCOCK ON EXTERNAL

the secretion having a repulsive odour. The secretion of the

paired gland has a ‘‘ mousy ” smell and is poured over the naked

skin of the fossa between the two labia. These labia are capable

of being widely separated so that the glandular area may be

flattened, but they are not closely applied and separated merely

by a narrow rima, like the labia of the homologous gland in

Viverra and Genetta*. The glans of the penis is long, flexible,

subeylindrical, weakly grooved below, and beset with recurved

horny papille. It ends in a styliform process.

The arrangement in the female is tolerably similar. The labia

of the gland arise in front on each side of the clitoris and pass

backwards to the anal area. They are naked on the inner side,

but hairy externally as in the male. Anteriorly they lie outside

the hairy labia of the vulva, which lies just behind the clitoris;

and the labia of the vulva unite posteriorly and form a low ridge

of naked integument which runs along the bottom of the space

between the labia of the gland and spreads to right and left,

forming a short transverse ridge in front of the anal area. The

labia of the gland can be folded over the vulva and the naked

glandular space behind the clitoris, exactly as in the male the

corresponding labia can be folded over the glandular space behind

the penis; but in the female these labia are not closely applied

as they are in Genetia and Viverra.

The anal area in both sexes is tolerably large and naked as in

Genetta and Cwettictis, but the repulsive secretion of the anal

glands is not retained by an annular ring of skin as in Viverra

zibetha.

In male examples of Paradoxurus niger and P. hermaphroditus

the glandular area closely resembles in its general features that

of P. larvatus in consisting of a naked, elongated, tongue-shaped

area of whitish glandular integument extending from the scrotum

behind round the prepuce in front. But there is no thickened

collar of skin round the penis and the labia of the gland are

much less pronounced, so that when the thighs are separated the

glandular surface forms no definite median fossa but is almost

flat from side to side. When, however, the inner surfaces of the

thighs are in their normal position and juxtaposed, the right and

left halves of the glandular surface are brought into contact.

The degree of development of these labia may prove to be a

generic difference between Paguma and Paradoxurus. At all

events, the conditions described hold good in the case of two

species of each of the genera as recognised by Gray. The penis

of Paradoxurus niger, judging from Turner’s description (P. Z. 8.

1849, p. 25), resembles that of P. larvatus, and the same

applies to the penis of P. hermaphroditus that I examined.

In a half-grown female of P. hermaphroditus the glandular

area consists of a nearly flat area of naked skin, without distinct

labia. The vulva lies near its centre, and on each side of the

* The gland in a living example of P. lewcomystax resembles apparently that of

P. larvatus.

CHARACTERS OF THE PARADOXURIN®. 405

vulva there extends backwards a shallow groove beset with a row

of coarse pores. The anterior end of each groove reaches as far

forwards as the corresponding end of the vulva, but the posterior

end reaches considerably farther back, nearly to the hinder end

of the area of naked skin. The low ridges between these pos-

teriorly converging grooves and the vulva are scantily hairy, and

constitute the labia of the latter orifice.

Text-figure 7.

—

SS ZS

Paradoxurus hermaphroditus.

A. Gland and external genitalia of half-grown female. g/., anterior end of glandular

area; v., vulya, surrounded by short hairs; p., row of secreting pores; a., anus,

B. The same of adult male. g/., glandular area flattened; p., prepuce at its

anterior end; sc., scrotum ; a., anus, with orifices of anal glands.

C. Transverse section of glandular area of male. gi., right portion of gland;

p-, penis.

This gland differs from that of the female P. larvatus in the

absence of distinct labia, as in the case of the males of the two

species.

406 MR. R. I. POCOCK ON EXTERNAL

The Perfume-gland and External Genitalia of Avctictis.

The gland of the male was first described by Cantor (J. A.S.

Bengal, 1846, p. 192), and subsequently by Garrod (P. Z.8. 1873,

p. 200). It resembles that of Paradoxurus larvatus in consisting

Text-figure 8.

Arctictis binturong.

A. Gland and external genitalia of male. g/., glandular area showing labia and

central depression; p., prepuce. se., scrotum; @., anus.

B. Gland of the female (vulva omitted) ; lettering as in A.

C & D. Transverse sections of the same with the labia of the gland partially

distended and nearly in contact. gl., gland of one side; 7., glandular space

between the labia (Z.).

of a pair of upstanding labia, hairy externally, smooth internally,

extending trom the scrotum up to the penis, and separated by

CHARACTERS OF THE PARADOXURINA, AQT

a naked fossa which becomes gradually shallower towards the

penis. There is no definite collar of skin round the prepuce but

integument, resembling that of the free edge of the labia in

being scantily covered with yellow hair, encircles that organ.

The posterior end of the fossa is partially divided by a low

membranous partition when the labia are not widely divaricated.

The penis, which I omitted to examine, was described by

Garrod as follows: “The glans penis is conical and pointed,

#inch long, and presents round its base several small, dark brown,

hard flattened papille about ;4, inch long.” From this it may

be inferred that the glans is shorter than in Paradovurus, has

no styliform termination, and is further distinguished by the

restriction of the papille to its base, their smaller number and

flattened, unspine-like shape.

The gland of the female does not appear to have been described.

It is very like that of the mule, consisting of a small longi-

tudinal fossa, with an upstanding labium, hairy without and

naked within, on each side of it. As in the male and in

Paradoxurus, these labia, whick converge anteriorly and poste-

riorly, are capable of being widely divavicated, but they are not

tightly juxtaposed as is the case with the homologous labia of

Viverricula, Genetta, etc. When only partially divaricated, the

floor of the fossa rises into a low median ridge of skin. The

vulva is situated in front of the glandular area, not near its

centre as in Puradowurus. ‘The glandular area is thus wholly

perineal as in the Viverrines.

The Perfume-gland and External Genitalia of Arctogalidia.

The gland in this genus does not appear to have been described.

Temminck and 8. Miiller, as Mivart stated, are silent about it.

Hence Mivart presumed the gland to be as in Paradoxurus

(P. Z. S. 1882, p. 165). On the other hand Blanford (Mamm.

Brit. India, p. 115, 1888) said: “there is no bald space in front

of the scrotum or around the genital orifice ; hence it is probable

that the prescrotal glands, if they exist, are ill developed.”

What material Blanford had whereon to base this opinion

does not appear; but the concluding sentence of the quotation

suggests that the result of his examination, presumably of dried

skins, was unsatisfactory. At all events his remarks do not

justify Lydekker’s statement that ‘there is no glandular tract

in front of the scrotum” (Lloyd’s Nat. Hist. : Cats. etc. p. 230,

1896).

I can say nothing about the gland in the male, but a female

example identified as A. lewcotis, from Sarawak, has a distinct

though small naked glandular tract in front of the vulva; or

rather, since the naked integument narrowly encircles the vulva

behind, that orifice, preceded by a well-developed clitoris, may be

described as situated at the posterior end of the glandular area.

The area itself is antero-posteriorly elongated and is surrounded

408 MR. R. I. POCOCK ON EXTERNAL

laterally and in front by an upstanding flap of quite naked skin

capable of being flattened out externally and anteriorly, or folded

over towards the middle line to form a pair of elongated labia in

contact throughout their length in front of the clitoris and

vulva.

The part of this naked area which appears to be specially active

lies just in front of the clitoris.

Behind the vulva there is a moderately long hairy, non-glan-

dular, perineal tract, so that the anus, situated in the centre of

a small, normal naked space, is remote from the glandular area.

Text-figure 9.

t VY

NWA

Bees WS SS

ies NYS

Gf f fin ‘\ \\\ Se

Ny i} Ih

Arctogalidia.

A. Gland and external genitalia of female. g/., secreting area of gland; 7., partially

distended upstanding labia surrounding it. e/., élitoris; v., vulva ; a., ANUS.

B. Lateral view of the same; letteting as in A.

C. Glandular area showing the labia folded in front of the clitoris ; lettering as in

A and B.

Since glands are present in both sexes of the genera of

Paradoxurine Viverrids described in this paper, and are better

developed in the males than in the females, it can hardly be

doubted that the male of Arctogalidia is also provided with a

gland ; but it appears to me to be impossible to foretell whether

the gland of the male will prove to be in front of the penis or

behind it. Perhaps the balance of evidence is in favour of its

CHARACTERS OF THE PARADOXURINA. 409

being prepenial, instead of prescrotal as in Paradoxurus and

others *.

However that may be, the gland of the female Arctogalidia

differs from that of Paradowxurus in several respects. In Arcto-

galidia the principal secreting area is the depression in front of

the clitoris, encircled by an upstanding flap of thin skin naked

without and within and apparently not specially glandular.

The right and left portions of this flap form two labia confluent

in front and capable of being folded over the glandular depres-

sion, thus meeting in front of the clitoris and vulva. Between

the vulva and the anus there is a tolerably long hairy perineal

area. In the female Paradoxurus the labia are thick, glandular,

and hairy externally. They are not confluent in front but

arise separately at the sides of the clitoris, pass backwards

on each side of the vulva and extend some distance behind

it, almost reaching the anal area, being separated therefrom

by a very short hairy non-glandular area. When folded over

they meet in the middle line behind the clitoris.

The Perfume-gland and External Genitalia of Nandinia.

The gland of the male was described by Flower (P.Z. 8. 1872,

p. 684) as follows :—‘* Nandinia resembles many of its allies in

possessing a... cutaneous scent-gland in the form of a longitu-

dinal median depression an inch in length, with tumid naked

margins and looking very like a vulva, situated in the pubic

region immediately in front of the short, conical retroverted

hairy prepuce.” This description is perfectly correct, but it

unfortunately suggests similarity in position between this gland

and that of the typical Paradoxurines.

Ten years later Mivart (P. Z.S. 1882, p. 170, note) made a

similar mistake when he described the gland as “a bald patch,

no doubt glandular, in the situation of the prescrotal glandular

structure of Genetta.” Itis quite true that the glandin Vandinia

is prescrotal, but it does not lie between the scrotum and the

penis as in Genetta, but in front of the penis as Flower stated.

In 1900 Miss Carlsson (Zool. Jahrb. Syst. xiii. pp. 509-528)

figured and described the gland of the female as a narrow area

of naked skin situated in front of the vulvaand ending anteriorly

in a distinct pouch. This had been previously detected by

Noack, who regarded the pouch as the functional homologue of

the marsupial pouch (Zool. Garten, Frankfurt, xxvii, p. 79, 1886).

In a half-grown female Vandiia I found the glandular area

exactly as described by Miss Carlsson, except that it was separated

from the vulva by a narrow area covered with hair and had

only an indistinct pouch at its anterior end. It consisted of an

elliptical patch of pale naked skin, with the anterior rim slightly

* Tf this proves to be so, a good case might be miade out for severing Nandinia

and Arctogalidia from the Paradoxurime as a separate subfamily Nandiniine.

410 MR. R. I. POCOCK ON EXTERNAL

raised. Probably as age advanced, the pouch would have been

developed and possibly the area would have extended backwards

to join the naked skin surrounding the vulva. The skin over-

lying the gland was thin, and the secretion was brown in colour

and strong in odour.

Text-figure 10.

Nandinea binotata.

A. Gland and external genitalia of adult male. gi., glandular area, flattened;

p., prepuce surrounded by short hair; sc., scrotum concealing anus.

B. The same with labia of gland folded in front of prepuce ; lettering asin A.

C. Transverse section of gland in front ef penis.

D. Glaus penis, its dersal side uppermost.

E. Gland and external genitalia of young female. gl., glandular area; ., vulva;

@., anus. i

In the adult male the naked glandular area extends from the

scrotum a long way in advance of the penis, which is situated

near its posterior end, only a little in advance of the scrotum.

Close to the penis on each side there are some hairs, sometimes

forming a definite crest, sometimes more numerous and less

regularly arranged. Sometimes there are also a few hairs right

in frent of that organ, but for the rest of the area it is quite

smeoth. The anterior half of the area expands and forms,

when the gland is flattened out by the separation of the thighs,

a shallow subcireular depression with tumid margins; but when

the thighs are juxtaposed the right and left halves of the area

approach, or nearly meet, across the middle line, being separated

by a moderately deep fossa the width ef which depends upon the

CHARACTERS OF THE PARADOXURIN&. 411

degree of approximation of the right and left halves of the area.

When its margins meet in front of the penis, these present the

appearance, described by Flower, of two naked ridges of skin

running forwards from the penis.

In longitudinal and transverse sections the gland is seen to

consist of a comparatively thin layer of dermal cells, of tolerably

uniform thickness transversely but gradually increasing in thick-

ness from behind forwards. The secreting cells are not

differentiated into a right and left thickening as in Paradoxurus,

Arctictis, and the Viverrines. The prepuce is hairy, and the

glans of the penis is quite short, a little expanded and rounded

distally, and without spicules.

General Conclusions with regard to the Perfume-glands.

From the account here given it is clear that the gland in the

Paradoxurines varies greatly in position in the genera discussed ;

but it is interesting to note the gradation in the situation it

assumes, especially in the female. In Arectictis it is entirely

perineal, lying between the anus and the vulva as in Genetta,

Viverra, etc., and in Hemigalus and Cynogale, although differing

structurally from the gland in those genera. In Paradoxurus

the glandular area passes anteriorly in front of the perineum, so

that the vulva les near its centre. Nevertheless, the labia of

the gland, as in Arctictis, fold over behind the vulva and clitoris.

In Arctogalidia the external genital organs are situated quite at

its posterior extremity, the secreting area being in front of the

clitoris and the labia also folding anteriorly to it. In Nandinia

the gland is altogether in front of the vulva and disconnected

from it. Thus WVandinia and Aretictis stand at the two

extremes.

In the males the differences between dArctictis and Paradoxurus

on the one hand, and Nandinia on the other, are equally well

marked, but the link in the chain of gradation is wanting,

possibly owing to our ignorance of the gland in this sex of

Arctogalidia.

Another point to consider is which of the two conditions pre-

sented respectively by Arcticiis or Paradowurus and Nandinia is

the more primitive. I incline to the opinion that the latter is

the derivative type; and for this reason—the position of the

prepuce considerably in advance of the scrotum is the condition

occurring in the Canide, Procyonide, and Urside, which are the

most generalised, on the whole, of existing Carnivores. It also

alanine in the Alluroid Cry yptopr octa. In the Pinnipedia, too,

the prepuce is situated a long way in advance of the anus, close

to which the scrotum was pr obably placed before its suppression.

On the other hand, the position of the penis just in front of the

scrotum is a character in which NVandinia reseinbles the Felide,

which are regarded as the most specialised of all the Carnivora.

The opinion above expressed can only be tentatively held,

Proc. Zoou. Soc.—1915, No. X XIX. 29

412 ON EXTERNAL CHARACTERS OF THE PARADOXURINA,

however, because the prepuce is close to the scrotum in the

Mongooses.

The main features of the glands and external genitalia may

be systematically summarised as follows :-—

Males*.

a. Glands paired, situated behind the prepuce, which lies far in

advance of the scrotum at the anterior end of the naked

glandular tract ; hence the labia of the gland when juxtaposed

meet behind the prepuce. Glans penis long or moderately

Fae and spicular.

. Glans penis subcylindrical, long, richly spicular, and ter-

minated by a styliform process ..... Paradoxurus.

6’. Glans penis conical, shorter, spicular ‘at ‘base, and without

styliform terminal process ....... Arctictis.

b. Glands unpaired, situated in front of the prepuce ‘which is

close to the scrotum; hence the labia of the gland when

juxtaposed meet in front of the prepuce; glans” penis quite

SMOKE GTAGL THHTEVATTEC! Sacenonscooasoonce coe snosde cod ane sudeseacsaadnancccens IN GNOGNIDIG

Females.

. Glands paired, behind and sometimes partially alongside the

vulva; labia of the glands wher juxtaposed meeting behind

the clitoris and v ulva:

. Vulva in front of the large naked glandular area ...... Arctictis.

ay Vulva near the middle “of the mae glandular area, . the

secretory area extending behind and alongside the vulva ... Paradoxurus.

b. Glands unpaired, secreting area altogether im front of the

vulva ; labia of the glandular area, when juxtaposed, meeting

in front of the vulva.

a®, Vulva situated at the posterior end of the naked glandular

tract which is bounded laterally and in front by up-

Shichu a ibayen JE opie hl ban aauce adatacese cuauantacanscnsae qaesnas obdeae adabcaded Arctogalidia.

52, Vulva separated from the naked glandular tract which has

a pouch-like depression at its anter doriend: ..235.. kOee asm da. Nandinia.

Considered collectively the glands of these genera differ from

those of the Viverrine in the followmg particulars :—In the

Viverrines the labia of the glands are hairy without and within

and form a pair of tumid masses usually closely juxtaposed or as

in Viverra zibetha, where they are divergent behind, confluent in

front behind the prepuce. In no case does the glandular area

extend in front of the prepuce or of the vulva, nor does it ever

form a flattened naked area covered all over with secretory

orifices. In the Paradoxurine genera, on the contrary, the labia,

when present as upstanding ridges, are naked within, like the

floor of the fossa between them. They are not closely applied

and can be so widely divaricated in both sexes that the secreting

surface, studded all over with pores, forms a flat or nearly flat

area. fs

* The glands of this sex in Arctogalidia are unknown.

ON THE ANATOMY OF GRUIFORM BIRDS. 413

31. Anatomical Notes on the Gruiform Birds Aramus

giganteus Bonap., and Rhinoehetus kagu. By P.

CHALMERS Mircamin, D.Sc, LL.D. ERS. #.Z.S..

Secretary to the Society.

[Received May 11, 1915: Read May 25, 1915.]

(Text-figures 1—5.)

INDEX.

ANATOMY: Page

beryl Osi sienna yee cc secern alee Nee cohen ese, SeLAlta

VAS COLA Renee eee tan cee eerie tent eee LAE

Musclestitnestn tren rate eth UNE et Ore Me me wees ies

OSteolo myers cae cee eee eee ae ee AT

SYSTEMATIC:

A. giganteus close to A. seolopaceus............... 413.

Relations of Aramis) sneccesss- cece eee ee 3

In July 1914 the Society purchased from a dealer a bird of

unknown source, recognised by Mr. D. Seth-Smith, our Curator

of Birds, as being new to our Collection. It turned out to be an

example of Aramus giganteus Bonap., the Limpkin, Clucking Hen,

or Northern Courlan (Aranus pictus Bartram of the Brit. Mus.

Catalogue, vol. xxiii. p. 238). It is the northern form of the

Courlan of 8. America, Aramus scolopaceus, and is a native of

Florida, the Antilles, and Jamaica, somewhat doubtfully accepted

as a distinct species because of its larger size, and of the extension

of the white stripes, confined in the Courlan to the head and

neck, to the back, wing-coverts, and lower parts. The anatomy

of A. scolopaceus has been studied chiefly by Garrod (P. Z. 8. 1876,

p- 275) and by myself (P.Z.S. 1901, p. 629). As I could find no

record of the anatomical examination of A. giganteus, | took the

opportunity of dissecting the Society’s example when it died after

living for a few months in the Gardens; and as | had formerly

dissected A. scolopaceus along with an example of the Ka gu

(Rhinochetus kagu), | renewed the comparison, by dissecting at

the same time another example of the Kagu.

It may be convenient to state at once the general conclusion,

that, so far as anatomical characters are concerned, 4. giganteus

resembles A. scolopaceus very closely indeed. It would not have

surprised me to find such shght differences as I noted between two

individuals of the same species. I do not suggest, however, that

the two species should be merged. The more experience I gain

of avian anatomy, the more I am convinced that systematists

are well advised when they rely, at least with regard to the

discrimination of species and genera, more on those superficial

characters that they can observe in the series of museum

collections, than on the uncertain indications afforded by the

presence or absence of this or that muscle.

29%

Al4 DR. P. CHALMERS MITCHELL ON THE

Pterylosis.— An aftershaft is present. There were ten primaries,

the most distal being smallest, each covert, as usual, being

placed distally to each primary. Dr. Gadow (Vogel, in Bronn’s

Thier-Reich, p. 79) states that there are 11 in Aramus, but in

some examples of A. gigantews and A. scolopacews which J examined

with Mr. W. R. Ogilvie-Grant at the British Museum (Natural

History) we found only 10. There are also only 10 in Lurypyaa

and Lhinochetus. The carpal covert is rather smaller than the

carpal remex, and crosses it in the fashion that the secondary

eoverts cross the secondary quills. A minute plica ties the

carpal remex to the most distal secondary quill, and the general

disposition bears out the view urged by Degen and myself that

the carpal feathers are in series with the secondaries, not with

the primaries. The wing is aquintocubital; the diastaxic gap

is wide and is occupied by a covert in the normal fashion.

The oil-gland is tufted, and there are twelve rectrices.

The disposition of the feather-tracts, as in A. scolopaceus, agrees

very closely with Nitzsch’s account of Psophia.

There were very thick clumps of down on the back, especially

on the rump. In the fresh condition these gave off some powder,

but detailed examination did not show the presence of true

decomposing down. The distribution of powder-downs amongst

birds is, as is now well known, far too irregular to be of use in

systematic classification.

There are no webs between any of the toes, and the claw of the

third digit is unsymmetrical, forming a scoop, slightly notched,

but not constituting a definite comb.

Viscera.—Both carotids are present. The trachea is uncon-

voluted ; the rings are highly ossified, and the normal pair of

extrinsic muscles and the intrinsic muscles are both present.

The configuration of the alimentary tract is of the generalized

type that I have shown (Trans. Linn. Soc., Zool. vol. viii. p. 173,

1901) to underlie the patterns displayed by Charadriiform and

Gruiform birds. The duodenum is a long and narrow loop;

Meckel’s tract is suspended at the circumference of a nearly cir-

cular expanse of mesentery ; between the duodenum and Meckel’s

diverticulum there is a single rather wide loop ; the diverticulum,

which is large and prominent, is placed on the distal limb of the

moderately long axial loop, and the distal portion of Meckel’s

tract is thrown into three or four rather irregular short loops.

The ceca are functional, of moderate length, and slightly

expanded towards their extremities. The rectum is straight and

rather wide. <A ganglionated sympathetic nerve chain passes

round the mesentery, bending a short way into the more

important loops. I have to add to the description I gave of

A. scolopaceus that there are two bridging veins from the

duodenal loop to the distal part of Meckel’s tract. ‘The example

of A. scolopaceuws that I examined had been preserved in spirit,

and very likely I overlooked the presence of bridging veins.

From the evidence afforded by this example of dA. giganteus, which

ANATOMY OF GRUIFORM BIRDS. Al5

I examined before it had been hardened by spirit, I should place

the pattern of Aramus definitely on the Gruiform stem, rather

than at the branching of the Gruiform and Charadriiform

stems.

Muscular Anatomy.

Cucullaris—The cervical portion is much feebler than in

Rhinochetus, but its posterior edge nearly meets the anterior edge

of the rhomboideus externus.

Rhomboideus ewternus.—A. continuous thin sheet of muscle,

distally leaving uncevered a good piece of the rhomboideus

profundus, and proximally stretching far anterior to it. In

Rhinochetus the arrangement is similar, but the muscle thins out

in the middle region and becomes much thicker in front.

Rhomboideus profundus.—Vhis reaches to the extreme distal end

of the scapula and extends forwards almost the whole length of

the scapula ; it is much thicker than the superficial muscle.

Latissimus dorsi metapatagialis—This slender muscle is

present, lying just under the skin, superficial to the latissimus

dorsi posterior and inserted into the skin underlying the humeral

feather-tract. It appears to have similar relations in all the

Gruiform birds.

Latissimus dorsi anterior and l. d. posterior.—These two

muscles are well developed, but do not touch at origin or

insertion. The tendinous insertion of the posterior division is

just deep of, and in contact with, the anchor of the anconeus

humeralis ; its origin is enormous and is partly from the ilium.

In Rhinochetus the origin from the ilium is smaller, and although

the insertion of the posterior division has similar relations with

the anconzeus anchor, it is just in contact with the proximal edge

of the fleshy insertion of the anterior division, In the Crane the

two divisions, especially the posterior, are relatively narrower and

do not touch at origin or insertion.

Serrati. The s. superfic. metapatagialis is large, anising only

from the ribs without fibres from the scapula, and is inserted to

the skin under the dorsal feather-tract. The s. superfic. anterior

is a narrow slip arising from one rib and is inserted by a flat

band to the scapula between the two divisions of the sub-coraco-

scapularis. The s. superfic. posterior is strong, arising from, two

ribs with their uncinate processes, and passing to the posterior

inferior border of the scapula. The s. profundus is in a series of

strong separate slips.

Scapuli-humerales anterior et posterior.—The posterior muscle

(teres major) is large and is pierced by the expansor secundariorum.

I found no trace of the anterior muscle, although Ftrbringer

states its presence in A. scolopaceus, and in the example of that

bird which I dissected I noted a few fibres representing it. It

is absent in Rhinochetus.

Deltoides propatagialis— A rather narrow muscle, not divided

into two peaks, It gives rise to the brevis and longus tendons

416 DR. P. CHALMERS MITCHELL ON THE

after a very short common course; the brevis tendon is very

broad and gives off a distinct distal slip (gamma of Firbringer),

a median patagial fan, and a separate proximal slip (alpha of

Fiirbringer). There is no recurrent anchor to the longus tendon.

The longus tendon is doubled at the elastic part and receives the

whole of the biceps patagialis, the tendon of which is very short.

The pectoralis patagialis is large and runs to the patagial tendon,

where it divides into longus and brevis slips. The condition in

Rhinochetus is as I have already figured (Joc. cit. text-fig. 76),

except that the alpha and gamma divisions of the brevis were

‘ather better separated.

Deltoides major et minor.—The major is a strong muscle reach-

ing down to the end of the third quarter of the humerus; it is

shorter in Rhinechetus. The minor, as in other Gruiform birds, is

quite separate, and is inserted to the other side of the insertion

of the pectoralis minor.

Pectoralis thoracicus.—A very strong muscle with a feeble

fibrous anchor to the humerus, and a single elongated tendinous

insertion. In Rhinochetus the muscle is similar, except that it is

narrower, in association with the shallow keel of the sternum.

Supra-coracoideus ( pectoralis minor), coraco-brachialis externus,

coraco-brachialis internus, and swb-coraco-scapularis.— Practically

identical in their relations in the two species of Aramus and in

Rhinochetus, except that in Rhinochetus the coracoidal origin of

the last-named muscle is relatively very much larger.

Anconeus.—The humeralis division, as in the vast majority of

birds, is a strong muscle arising from the greater part of the

humerus, and is cleft proximally. The scapular head, in addition

to the steut anchor to the humerus described in connection with

the latissimus dorsi muscles, has the origin from the scapula

forked in Aramus, simple in Rhinochetus. The anconeus caput

coracoideum (expansor secundariorum) in Aramus presents the

arrangement described as “‘ ciconine” by Garrod ; that is to say,

it arises in the axilla from a triangular ligament which passes

into a stout tendon running to be inserted to the proximal

secondary quills. I have already stated (P. Z.8. 1901, p. 645)

that this arrangement is usual in Gruiform birds, but that I

found the muscle feeble, although present, in ?hinochetus and

Eurypyga. In the example of Rhinochetus which I used in the

dissections I am now describing, I found no trace of the expansor

secundariorum in either of the wings. This may serve as

another of the many warnings against attaching systematic value

to the absence of an anatomical structure, as it may have been

lost independently by different descendants of a common

ancestor.

Biceps brachti—In Aramus the acrocoracoidal portion of the

muscle gives rise to a tendon which, although closely associated

with the humeral portion, can be traced separately to the ulna,

the humeral portion going to the radius. In Rhinochetus the

division of the tendon of insertion to the radius and ulna is

ANATOMY OF GRUIFORM BIRDS. A417

forked at the extreme distal end. The peculiar accessory head of

the biceps, first noted by Beddard (P.Z.S. 1891, p. 14) and

confirmed by myself (P. Z. 8S. 1901, p. 639) from dissection of

another example of Rhinochetus, was doubttully represented in

the present example, an old and excessively fat bird, by a lump

of fat and fibrous tissue.

Llio-tibialis internus (sartorius),—As in other Gruiform birds,

rather easily separable from the gluteus maximus,

Llio-tibialis (gluteus maximus).—This large muscle, with rather

weak median and very strong post-acetabular portions, had the

disposition found in Gruiform birds generally,

llio-trochanterici posterior, anterior, and medius, were all

distinct and well developed as in other Gruiform birds, but the

small median muscle (gluteus quartus) was absent in 4, giganteus

as in A. scolopaceus; in the present example of Hhinochetus it

was just separable. |

Caud-ilio-femoralis (femoro-caudal and accessory).— The femoro-

caudal was absent in A. gigantews as in A. scolopaceus, present

but small in Rhinochetus ; the accessory was large, but with an

area of tendinous degeneration in A. giganteus, precisely as in

A. scolopaceus, absent in Rhinochetus.

Ambiens.—Present in both A. giganteus and Fhinochetus, its

slender tendon, after passing round the knee, receiving a strong

anchor from the head of the fibula.

Caud-ilio-flecorius (semitendinosus and accessory semitendinosus).

Ischio-flexorius (semimembranosus). Gastrocnemius, middle, or

posterior femoral head.—¥or convenience, to explain the in-

teresting relations of this group of muscles, | have repeated here,

as text-figures 1-4, the text-figures 81-84 in my paper on

Gruiform birds already published in the Proceedings of this

Society (P. Z. S. 1901, pp. 650-651). In 4. gigantews, as in

A. scolopaceus (text-fig. 2), the semitendinosus (C.I.L. 2) is wide,

but not so large as the semimembranosus (I.F.), except towards

its insertion, whereas in Rhinochetus it is enormous, much

_larger than the semimembranosus. In 4A. giganteus the accessory

semitend. (C.I.L. 2) is a broad triangular muscle, meeting the

semitend, in an oblique raphe, exactly as in A. scolopaceus (text-

fig. 2), and then running down to join the middle head of the

gastrocnemius, which it covers; the semimembr., also as repre-

sented in that figure, sends a slip to the conjoined semi-

tendinosus and accessory semitend., and passes on to the tibia

by a flat ligament (A, 2). There was also a thin flat ligament

from the semitend. raphe to the tibia, which I did not find in

A. scolopaceus, but which I found in fallus (text-fig. 1, A. 1).

The condition of this group of muscles in the example of

Rhinochetus that I am now describing, corresponded, except in

one important respect, with what I found on a former occasion

and described as resembling the condition in Otis (text-fig. 3).

The semitendinosus, accessory semitend., and semimembranosus

unite to form a flat tendon of insertion to the tibia (A. 2) plainly

418 DR. P. CHALMERS MITCHELL ON THE

Text-figure 1. Text-figure 2.

Knee-muscles of Rallus longirostris. Knee-muscles of Aramus scolopaceus.

Musculature of knee; right leg, internal view.

FEM.-TIB.-I. Internal separate slip of femoro-tibialis (vastus internus).

AX. Ligament from femur to tibia.

P.I.F. Pub-ischio-femorales (adductors).

C.1.L, Caud-ilio-tlexorius. C.0.0.1. Kemoral insertion of caud-ilio-flex-

orius (accessory semitendinosus). C.I.L. 2. Semitendinosus.

IF. Ischio-flexorius (semimembranosus).

A. 1. Anchor to tibia of C.L.Z.

A.2. Anchor to tibia of IF.

Ge. 1. External femoral division of gastrocnemius.

Ge. 2. Internal femoral divison of gastrocnemius.

Ge. 3. Tibial division of gastrocnemius.

SOT. Soleus.

In text-figure 2 the caud-ilio-flexorius has had a piece removed to exhibit

the deeper lying Gre. 2.

The tendinous areas are dotted.

Text-figure 3. . Text-figure 4.

Knee-muscles of Otis tarda. Knee-muscles of Heliornis fulica.

Description and lettering as in text-figures 1 & 2.

ANATOMY OF GRUIFORM BIRDS. 419

identical with the tendon of insertion of the semimembranosus

in Aramus and Rallus. This tendon arises from the semiten-

dinosus raphe, and the muscular bellies pass under it, meet a

separate muscle (text-fig. 3, Ge. 2) arising from the femur just

proximal to the accessory semitendin., and form a strong tendon

that joins the common tendon of the gastrocnemius. <A glance

at text-figures 1, 2, and 3 will show that this muscle, marked Ge. 2,

appears to be present in Rallus, Aramus, Otis, and Rhinochetus,

but that in the two former it is fused along its edge with the

accessory semitendinosus, whilst in the two latter it is free. If,

then, we identify the muscle marked Ge. 2 in text-fig. 3 with the

middle head of the gastrocnemius, it seems a sufficient statement

of the facts to say that the middle head of the gastrocnemius in

Rallus is either absent, or fused with the accessory semiten-

dinosus. But in both species of Aramus, although the accessory

semitendinosus is sufficiently broad to pass both for itself and a

fused ralline-like middle head of the gastrocnemius, there is, in

addition,a distinct middle head (text-fig. 2,Ge. 2) apparentlyabsent

in Rallus. In Psophia, the middle head of the gastrocnemius 1s

double, one portion corresponding with the Ge. 2 in text-fig. 2,

another with the Ge. 2 in text-fig. 3. In my paper already cited

(P. ZS. 1901) I stated that the external head of the gastro-

cnemius in Oé¢is was trifid, one of the three origins passing under

the biceps. I do not doubt but that the latter is morphologically

identical with the slip marked Ge. 2 in text-fig. 2. In the example

of Rhinochetus that I am now describing, I found precisely the

same condition; not only is the Ge. 2, represented in text-fig. 3,

present, but there is also a separate middle head corresponding

with the Ge. 2 of text-fig. 2. Dr. Beddard (P. Z.S. 1891, p. 16)

has published a description and an incompletely lettered figure

of the gastrocnemius and its relations in RAinochetus. Although

his description is difficult to follow, as it omits reference to the

tibial head of the gastrocnemius which, so far as I know, occurs

in all birds, and refers to a head ‘‘formed by a broad flat tendon

to the head of the fibula,” a disposition which is at least abnormal

in Avian anatomy, I infer that the arrangement of the gastro-

cnemius in the example of Rhinochetus that he dissected was

identical with that which I describe here.

For some time I have been collecting notes on the various

fashions in which the middle head of the gastrocnemius and the

accessory semitendinosus are disposed in birds, but I do not wish

to offer these results for publication until I can present a fairly

‘complete picture of their distribution in the Avian system. Dr.

Gadow (Vogel, in Bronn’s Thier-Reich, p. 184) distinguishes three

conditions of this complex: (i.) when the accessory semitendinosus

cannot be separated from the middle head of the gastrocnemius ;

(ii.) when both muscles are present but quite distinct except in so

far as they may be connected by secondary tendons; (iil.) when

the accessory semitendinosus is absent but the middle head of

the gastrocnemius present. This grouping will not contain all the

facts ; there is a condition, as in Rhinochetus and Psophia, which

420) DR. P. CHALMERS MITCHELL ON THE

corresponds with Gadow’s second group, except that there is, in

addition, a second distinct middle gastrocnemius.

Crotrocnenxine external femoral head, and internal tibial head.

Mio fibularis (biceps, with sling and anchor). Solews.—A]l these

presented no peculiarity, and were similar in the two species of

Aramus and in Rhinochetus.

Pub-ischio-femorales (adductors).—In all three birds both were

present, without trace of tendinous degeneration, and, as in most

Gruiform birds, the internal muscle was much stronger and wider

than the external muscle.

Ischio-femoralis (obdurator externus), obdurator (obdurator

internus); accessorit obduratoris.— With regard to these muscles,

I have to note only that the obdurator externus was notably

small in A. giganteus, and that the area of insertion, to the shape

of which Garrod attached importance, was markedly triangular

in that bird, as in A. scolopaceus and in Rails and Cranes. It is

much more oval in Rhinochetaus.

Peroneus superficiatis and peroneus profundus.—tin both species

of Aramus and in Rhinochetus the superficial muscle is the more

important, with a broad superficial origin, extensive origin from

the fascia of the adjacent muscles and, in Avamzs, almost no deep

origin, but a certain amount of it in Rhinochetus. The anchor

to the sustentaculum of the flexor tendons is broad in all three

and shows traces of being broken into separate slips.. The

tendinous slip to the flexor of the third digit is well developed in

all three birds. The deep muscle is very strong in Rhinochetus,

arising from a large part of the fibula, beginning at the insertion

of the biceps tendon, and from the adjacent surface of the tibia.

It passes in the nor mal fashion into a flattened tendon running

in a groove under the slip to digit Ii] of the superficial apiece.

In ve scolopaceus 1 did not find it, but in A. giganteus it was

represented by a very minute muscular head arising high up from

the fibula opposite the biceps insertion, and passing at once into

a most slender ossified tendon, so closely adherent to the fibula

that, unless it weve specially searched for, it might easily be missed.

It has the usual insertion by a very thin flattened tendon in a

groove under the slip to III. The tendons of both peroneals were

ossified in Aramus, as in many other Gruiform and Limicoline

birds, but in Rhinochetus there was no trace of ossification. I have

shown, in a former communication tothe Society (‘* The Peroneai

Muscles in Birds,” P. Z. 8. 1913), that in Gruiform birds there

is a tencency for the deep peroneal to become reduced or to dis-

appear, 60 that it would not be remarkable to find it present in-

A. giganteus and absent in A. scolepacews, even if these forms

were not specifically distinct.

Tibialis anticus; Extensor digitorum communis ; Hlexores per-

forantes et perforadi of digits IT and I11.—These muscles are all

present in both species ef Aramus and in Rhinochetus and have

the disposition that is normal in birds generally. The most

notable point of difference is that in Aramus the tendons are all

ANATOMY OF GRUIFORM BIRDS. 49)

ossified, above and below the ankle, and are unossified in Rhino-

chetus. ‘The flexor tendon of digit III is connected with the

perforated flexor tendon of the same digit by a strong slip.

Flexores perforati, of digits II, III, 1V.—In all three birds,

as in Gruiform birds generally, there are three heads for this

group of muscles, and fibres from each of the three tendons can

be traced to each of the three heads. ‘The ambiens head is the

tendon of the ambiens muscle, which passes under the biceps

ligament, that is to say, between it and the fibula. The fibular

or external head arises by a tendon from the fibula, proximal to

the insertion of the biceps and passing superficially to the biceps

tendon. The deep or femoral head is fleshy from between the

condyles of the femur.

Flexor longis hallwcis—In all three birds this arises by a

single head from between the condyles of the femur.

Flexor profundus.—Arises from the shaft of the tibia close under

the fibula. In Aramus the tendons of the deep flexors are nearly

equal in size; in Rhinochetus the tendon of the longus hallucis is

relatively smaller. In A. giganteus the relations between the two

deep tendons are much as I represented in A. scolopaceus (P. Z. 8.

1901, p. 654), except that the vinculum to the other deep flexor

is spread over the branches to the digits rather more, as in the

figure of Kurypyga given on thesame page. The present example

of Rhinochetus was exactly similar in this respect to the figure

I have given, also on the same page.

The muscular anatomy affords small ground for separating

A, giganteus from A. scolopaceus. The most notable differences

relate to muscles that are degenerate in the group, and that

might even vary individually—such as the presence of a teres

minor in 4. scolopaceus which is absent in A. gigantews, and the

ptesence of a peroneus profundus in the latter species and its

absence in the former.

The muscular arrangements in which the two species of Aramus

agree and differ from Cranes are more numerous, and are in con-

formity with the separation of a sub-family Aramine.

Osteology.

I have noted only a few osteological points, the systematic

value of which I do not propose to discuss.

Like that of A. scolopaceus, the skull of A. giganteus is strongly

schizorhinal ; the lacrimals are not fused with the skull as occui's

in Rhinochetus and Limicoline birds. The palate is widely

schizognathous with a long pointed vomer, and the pterygoids

are short, and expanded anteriorly asin Grus. Also, as in Grus,

there is a pair of occipital foramina, and strong descending ex-

occipital processes, both absent in Ti inoche tls. The skulls of

both species of Aramus are exceedingly like those of true Cranes,

the likeness extending not only to the larger characters that have

429, DR. P. CHALMERS MITCHELL ON THE

been employed in classification, but to the minute configuration

of the bones.

Sternum.—In Gadow’s valuable synoptic table ( Vogel, in Bronn’s

Thier- Reich, p. 79), he states that the external spine 1s very large

in Grues (in which he includes Aramus) and very small in Ahino-

chetus, and that the internal spine is absent in Grues and in Rhino-

chetus. With regard to the internal spine, I confirm his statement

for Cranes, Rhinochetus, and both species of Aramus. The

external spine is certainly small in Rhinochetus, but it was also

Text-figure 5.

Shoulder-girdle articulations in Gruiform Birds. I. Balearica. II. Grus

(australasiana & earunculata) Il. Aramus giganteus. 1V. Rhinochetus kagu.

A. Acrocoracoidal process of coracoid. C. Coracoid, Cl. Clavicle. P. Procora-

coidal process of coracoid. S. Scapula.

small in both species of Aramus. In the examples of Grus

that I have examined (G. australasiana and G. carunculata) the

external spine was very large and hollowed for a coil of the

windpipe. In an example of Balearica, on the other hand, it was

as minute as in Aramus and Rhinochetus. The sex of the skeleton

had not been noted, but it appears probable that the huge

external spine of Cranes is developed in connection with the

ANATOMY OF GRUIFORM BIRDS. 423

windpipe, and is not to be regarded as a character of systematic

value.

Shoulder-girdle articulation.—In text-fig. 5 I give drawings of

the relations of the coracoid, scapula, and clavicle, to which

Fiirbringer has paid so much attention in his great monograph on

Birds. Fiirbringer has figured the same structures in the case of

Grus, Psophia, Aramus, Hurypyga, Dicholophus, and Otis (Morph.

u. System. der Vogel, 1888, pl.ii., figs. 40-45). Comparison of his

figures with those that I give here, will show that, within the limits

of Gruiform birds, the characters vary considerably and are difficult

to value for systematic purposes. alearica agrees very closely

with the figure Fiirbringer gives for G'rus, except that the pro-

coracoidal process is not quite so large. My figure of Grus, which

applies equally to G.. australasiana and to G. carunculata, shows

much closer articulation of the clavicle and scapula than was

figured by Furbringer, as well as a smaller procoracoidal process.

Aramus giganteus differs from the figure given by Furbringer,

possibly for A. scolopaceus, in that the articular end of the clavicle

forms a broad flattened plate articulating with the scapula.

Rhinochetus has a large procoracoid, and very slight articulation

of clavicle and scapula. According to Furbringer, Psophia shows

the most complete articulation of scapula, clavicle, and procoracoid.

Systematic Position of Aramus.—l have already shown (P. Z.8.

1901, pp. 629-655) that A. scolopaceus fitted naturally with Gru1-

form birds, and the examination of A. gigantews confirms this in

every respect.

ibaa

vivota A

Pee eer dy bt

‘SQONAIONVIAWN SNdOUNTAY ‘6 “INOOVG SNdOdCISuUNTAV 1

PP Bompoom W'9

09 SulAawABU_ 0147001 UVMS

‘T’TId ‘GUYMCAOOM ‘SI6I ‘S'Z'd

ON AN BXTINCT CARNIVORE. 425

On the Skull of an extinct Mammal related to Avluropus

from a Cave in the Ruby Mines at Mogok, Burma.

By A. Suita Woopwarp, LL.D., F.R.S., V.P.Z.S.

[Received May 25,1915: Read June 8, 1915. ]

(Plate I.* & Text-figure 1.)

INDEX.

Page

Atlureidopus bacont, gen. et sp. 0M. .. ...........-........, 428

The rave and remarkable mammal /Juropus, now confined to

the highlands of eastern Thibet, is evidently the survivor of a

group which must have hada wide geographical range in compara-

tively modern geological times. It is so completely intermediate

between the Procyonide and the Urside, that it is sometimes

placed in the one family 7, sometimes in the other £; and its

velationships to the Phocene Hyc«narctos are so obvious §, that it

must doubtless be regarded as a somewhat modified survivor of

the common stock from which the Procyonide and Urside have

diverged. No closely related fossil forms, however; have hitherto

heen recorded ; and the recent discovery of a skull of an allied

extinct species is therefore of interest.

The new specimen (text-fig. 1) was obtained from a cave at the

ruby mines, Mogok, Upper Burma, by Mr. A. L. Bacon, and

brought as a gift to the British Museum by Mr. F. Atlay. The

skull lacks both zygomatic arches and the anterior end of the

palate with the incisors and three of the premolar teeth. It

must, in fact, have lain exposed for some time in the cave; for

the whole of the sagittal crest has been gnawed away by arodent,

evidently a porcupine ||, and there are similar tooth-marks alon

the lambdoidal border and other parts of the occiput. Otherwise

the fossil is well preserved and all its characteristic features are

shown.

Although it is not mineralised, the bone is remarkably dense

and heavy, as in the skull of Blur opus melanoleucus 4]. Nearly

all the sutures between the elements are closed, and the specimen

represents a fully adult individual, which was slightly larger and

more robust than the described examples of the existing species.

It agrees with the latter in all essential respects, such as the

* For explanation of the Plate see p. 428.

+ E. Ray Lankester, “On the Affinities of Hluropus melanoleucus,” Trans. Linn.

Soc., Zool., ser. 2, vol. viii. (1901), pp. 163-172, pls. xvili.—xx.

ab K.S. ‘Bardenfleth, “On the Systematic Position of Aluropus melanoleucus,”

Mindeskrift tor Japetus Steenstrup (1913), art. xvii.

§ H. Winge, E Museo Lundii, vol. ii. (1896), pt. ii. no. 2, p. 62.

\| Similar gnawing of fossil bones has been noticed by R Ly ee “The Fauna

of the Karnul Caves,” Paleeont. Indica, ser. 10, vol. iv. fe p. 2

“| E. Ray Lankester, loc. cit. p. 165.

426 DR. A. SMITH WOODWARD ON

Text-figure 1.

AIX SS

NS ee\ \\ \\

Lp “a }

<= i

Nie

G)Woeodward

A.-C.-— Alureidopus baconi, gen. et sp. nov.; imperfect skull from the left

lateral (A.), palatal (B.), and upper (C.) aspects; one-quarter nat. size.

g. Marks of gnawing by rodents, probably porcupines.

D.— Lluropus melanoleucus A. Milne-Edw.; left upper dentition, outer aspect ;

one-quarter nat. size.

pm. 1-8. First, second, and third premolars.

AN EXTINCT CARNIVORE. 427

disposition of the basicranial foramina, the ending of the palate

posteriorly between the last molars, the position of the infra-

orbital foramen on the cheek, and the absence of a postorbital

prominence. It differs only in its steeper frontal profile and

certain minor characters. ‘The inner face of the stout mastoid

process, for example, is irregularly ridged, not smooth as in

Hluropus. The foramen which pierces the inner wall of the

lachrymal pit in the existing species, is behind this pit in the

fossil.

The molar and premolar teeth, so far as preserved, agree closely

with those of luropus, and exhibit only a less marked crimping

and less tendency to subdivision of some of the cusps. Except

these features, there is nothing to remark about the molars (PI. I.

figs. 1 A-c, m. 1, m. 2). The upper sectorial (pm. 4) is noteworthy

for the simple and robust character of its antero-internal cusp o1

protocone. The third premolar (pm. 3) is preserved only on the

left side and agrees precisely with that of Zluropus, but is rela-

tively a little larger. Hven in the existing genus the space for

the first two premolars is so short that the double-rooted pm. 2 is

thrust crosswise and pm. 1 is reduced to a single minute cusp

(Pl. I. fig. 2); but in the imperfect fossil, which does not retain

either of these teeth, the corresponding space is still shorter.

The bone is broken away on the right; but it seems to be suffi-

ciently well preserved on the left to show that only one single-

rooted premolar originally occupied this position. At least, a

single socket of moderate size fills the whole of the space between

pm. 3 and the canine. The canine (c.), broken on the left, but

completely preserved on the right side, closely resembles that of

Aluropus, even to the faint crimping of its posterior keel ; it is,

however, slightly more robust than in the described specimens of

the existing species.

Some of the principal measurements of the fossil, in fractions

of a metre, are as follows :—

Length from anterior border of nasals to posterior face of occipital condyles. 0°25

Maximum width between outer borders of mastoid processes, about ......... 0°18

Maximum width of basioccipital between omapanie bulllees jest. qateeeas OL043:

Frontal width at postorbital point... - Lue bvantahemaeetaenen O:066

Depth of snout from alveolar border above. pm. 3, about cn. shaonen » OXON)

Depth between alveolar border and beginning of sagittal crest, about ......... 07135

ML, Beane ome ERNE sao goyeqnocqoen ano senaeclapeceandaato casaun condos enined wavesuecdaga OLOBIS

M. 1— 3% JKSSaYSXHO). Bap Bee ap cele nom aaneemear eet ae coc Iai tee ea ee name 0:027

He UR TAGLT tot Wa coer mR ce bata ln.” a te RIM 0}

Pm.4— ,, lene Chime ey epee hy eet bomen ieee gin Lee OOD

a NVC UB Oe peo NE RAC Re CREB SE OnE BCU SCE CORE SnD INC Os SRR SE Seem eam epraemnn (0027

Pm.3— _,, DNSSCEA dO ees ciate Mists oe Babe enacee Gao HORA CORE ES Ee ned Een ON O33

. width ae PRE RS ER AN ace een eR eee ud oe ee ck as wea toeae UOROLA,

Canine— ,, length of base x6 BBBEOA ODE “SO SUEDE AED MIBEER A en RE OEE PEDO con AEK aU OP-3)

55 Wide MORACEAE uM ok tsa tues noel 0:016

The fossil from Mogok, therefore, differs essentially from the

skull of the existing species of dlwropus in the shortness and

bluntness of the snout, with the consequent reduction of space

Proc. Zoou. Soc.—1915, No. XXX. 30

428 ON AN EXTINCT CARNIVORE.

for the anterior premolars. It remains only to decide whether

these differences are to be regarded as of generic or of specific

value. If the anterior premolars were merely reduced in size,

there need be no hesitation in adopting the latter alternative ;

but as pm. 2 differs fundamentally in having only a single root

while pm. 1 was probably absent, I am inclined to refer the fossil

to a distinct genus. I propose that this genus be named Alurev-

dopus, and defined from Zlwropus by the presence of only three

upper premolars of which the foremost has a simple root. The

species represented by the skull now described may be appropriately

named baconi after its discoverer.

EXPLANATION OF PLATE I.

Fig. 1. Hiwreidopus baconi, gen. et sp. nov.; left upper dentition from the outer

(A), lower (B), and inner (C) aspects, partly restored from the right side

of the fossil, nat. size; c. canine; pm. 2, single socket for second premolar;

m. 8, 4, second and third premolars; m. 1, 2, first and second molars.

Fig. 2. Hluropus melanoleucus A. Milne-Edw.; left upper canine (c.) and first to

third premolars (pm. 1-3), lower aspect, nat. size.

ON AVIAN CESTODES. 429

33. Contributions to the Anatomy and Systematic Arrange-

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

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

[Received and Read May 25, 1915.]

(Text-figures 1-6.)

XVII. On T_£NI4 TAURICOLLIS OF CHAPMAN AND ON THE

GENUS CHAPMANLA.

InpDEx. Page

Structure of Chapmania tawricollis .................. 480

Systematic position of C. tawricollis.................. 439

Generarof Udiogenimeyeeeeee eee eee 4.

I obtained, on June 30th, 1914, a large number of examples of

Cestodes from an «American Rhea (Rhea americana) which died

in the Society’s Gardens on the previous day. All these worms

were found in the right cecum, and none of them in the small

intestine. As is known, the cecum is very rarely found to be

inhabited by Cestode parasites, though there are cases on

record *.

From this bird three species of Cestodes have been described.

Of these three, Cittotenia rhece may be set aside at once as having

nothing in common with the species which forms the subject of

the present remarks. ‘This latter species is identical with either

Tenia tauricollis of Chapman or Tenia struthionis Houttuyn

among those which have been found in Rhea, or is another

species altogether. Zenza tauricollis has been placed in a genus

Chapmania (which is not widely different from Jdiogenes), while

Tenia struthionis is now assigned to Davainea. In spite of this

generic separation, which depends upon the not at all full descrip-

tion of Davainea struthionis by Paronat and von Linstowt, I

am by no means certain that they are not actually identical

at least generically.

Davainea struthionis, though found in the Ostrich, is stated by

Fuhrmann § to occur also in Rhea americana; but the statement

of Fuhrmann is not advanced with absolute positiveness ; and it

is only a statement in that no characters of Davainea struthionis

are given by him to confirm the identification. The fullest account

of the structure of Davainea struthionis is that of v. Linstow.

Parona’s notes give no anatomical detail to speak of. It is prob-

ably therefore to the former paper that the responsibility for

including this species in the genus Davainea by all subsequent

* See Bronn’s ‘ Thier-Reich,’ Bd. iv. Abth. 1B, p. 1625.

+ Ann. Mus. Civ. Genova (2a) ii. 1885, p. 425.

{ Arch. f. mikr. Anat. xlii. 1893, p. 447.

§ “ Die Cestoden der Vogel,” Zool. Jahrb. Suppl.-Bd. x. 1908, pp. 6 and 19.

30*

430 DR. F. E. BEDDARD ON

authors is to be referred. Von Linstow himself assigns the worm

to the genus Davainea by reason of the form of the hooks. That

character, however, is now known in other genera—though all of

them of the family Davaineide.

Those who have written subsequently upon this species, Da-

vainea struthionis, have probably accepted its generic rank rather

from the remarks of Max Braun than from the facts detailed in

vy. Linstow’s memoir. The former has pointed out in Bronn’s

‘Thier-Reich’ * that the alleged ovary of v. Linstow, consisting of

a number of separate spherical masses, is not indeed the ovary,

but an instance—largely met with in the genus Davainea—

of the “ Parenchymkapseln ” of German writers, in which the

ripe ova are massed after the disappearance of the uterus (when

that sac is clearly developed in the genus, which would seem to

be not always). Apart from this interpretation of the “ ovary ”

of v. Linstow and the nature and distribution of the set

according to v. Linstow, there is nothing in the description of

that writer which would justify the reference of ‘‘ Tenia struthi-

onis” to the genus Davainea. It will be observed that there is

therefore no reason to distinguish this worm from Chapmania

tauricollis, at any rate so far. The detached masses of ripe ova

correspond with what I shall shortly describe in that species,

where the uterus is more ox less broken up into partly separate

cavities containing eggs. The general form and size of the species

which has been termed Davainea struthionis,as figured by Parona’,

is precisely what I have found to characterise the worm from

Rhea americana, which in other ways agrees positively with the

descriptions extant of Chapmania tauricollis. We have, how-

ever, to assume, if this identification be correct, that v. Linstow

has missed the paruterine organ. This is, however, not evident

in immature proglottids. I can hardly claim to have proved

specific identity ; but I believe it would be difficult on the

facts known to deny generic identity between these two species.

In the meantime, however, I identify the species described in

the present paper with that described by Chapman, Fuhrmann,

and others as Zenia, Davainea, or Chapmania tauricollis. Tt

reaches a length of fourteen inches or so and has thus much the

dimensions given by Chapman {. It is held that Tenia argentina

of Zschokke § is the same species. If so, there would appear to

be a discrepancy in that the measurement of length given by

Zschokke is 8-9 cm. Iam able, however, to clear up this diffi-

culty. The majority of my specimens agreed with Chapman’s in

their dimensions; but in a few the length was not greater than

8 cm. or so.

The anterior segments of the body form a long tract, which

is very slender and widens out more or less suddenly to the wider

* Bd. vi. pt. u. p. 1446.

+ Ann. Mus. Civ. Genova (2a), ii. 1885.

+ Proc. Acad. Sci. Philadelphia, 1876, p. 14.

§ Centralbl. f. Bakt. u. Paras. i11. 1888, p. 2.

AVIAN CESIODES. 431

(3-4} mm.) posterior region. The most posterior segments are

hardly, if at all, longer than wide. ‘There is no neck.

The figure given by Chapman * of the scolex and anterior

segments is quite in accord with the view that the species

described by himself is that which I regard here as “ Tenia

twwricollis.” He does not, however, figure or make any reference

to hooks upon the rostellum (or, for the matter of that, else-

where). Monticelli, however, in his fuller description, found

Text-figure 1,

Scolex and anterior segments of Chapmania tauricollis.

s. Sucker.

the hooks, which he described as “minutissimi uncini.” It

would not be difficult to miss these hooks; I find from a note

made by myself upon the living worm that no hooks were seen.

I have, however, closely examined the scolices of two preserved

examples—the only scolices which I possess—and find traces of

the hooks in one individual only. The other specimen was

* Proc. Acad. Sci. Philadelphia, 1876, p. 14.

+ Nat. Sicil. xii. 1892-3, p. 208.

432 DR. F. E. BEDDARD ON

devoid of hooks. The one which showed them only possessed

a few, which needed a high-power lens and good iUlumination to

show them up. It seems to me that they may easily be shed.

I am quite unable to make any statement as to the shape of

these hooks. ‘They appeared to be little more than acicular in

form, like the hooks of the suckers in some Davainea. The

feebleness of the rostellar hooks seems to me to be related to the

little marked character of the rostellum in this and other forms.

I may remark that there is no reason to believe that the present

species has any hooks upon the suckers *. I shall return later to

the questions of systematic arrangement which depend upon the |

presence of hooks in connection with the genera Zschokkeella and

Inermicapsifer and some others.

In spite of the fact that Chapman described the genital pores

of his species as alternating irregularly, the genus Chapmania

has been defined by some subsequent writers as unilateral, at any

rate in the particular species Chapmania tauricollis +. Chapman

himself remarked that five pores might follow consecutively upon

the same side of the body. I have seen ten segments in series

with the genital pores all upon the same side. It is therefore

obvious where the error may have crept in. The genital pores

lie at about the middle of the lateral border.

I shall now deal with a few points in the internal structure of

this Cestode.

The cirrus-sac does not seem to me to have been fully described

by Fuhrmann. The cloaca genitalis, as he has pointed out, is

deep and expands within the cortex in a funnel-shaped way.

Into the middle of this funnel opens the cirrus-sac, and through

it projects often the protruded cirrus. The cirrus-sac appears to

consist of two distinct regions, or it may be, as I shall point out

later, that the cloaca genitalis consists of two distinct regions. In

any case a bottle-shaped sac opens into what is unquestionably

the cloaca genitalis, the neck of which “ bottle” is much shorter

than the rest. This relatively narrow tube has muscular walls

quite continuous with those of the rest of the cirrus-sac, and in

it lies the cirrus, which nearly fills up its lumen in cirrus-sacs,

which are in an average state of protrusion of the cirrus. The

rest of the cirrus-sac extends into the bodyeof the worm some way

inwards of the lateral water-vascular tube and ends in a retractor

muscle, as has been described by Fuhrmann. The flask part of

the cirrus-sac is divided from its narrow neck by a sphincter-

muscle, which forms a collar within the lumen of the sac, and to

the inside of this a fan-shaped bunch of muscular fibres, provided

with nuclei at their internal ends, further blocks the lumen and

* Fuhrmann, however (Rev. Suisse Zool. iv. 1896, p.111), not only refers to hooks

upon the rostellum, but speaks of having seen the evidence of hooks upon one

sucker. The definitions of the genus Chapmania given by Fuhrmann and Ransom

do not use hooks upon the suckers as a character of that genus.

+ I. e., Ransom, Proc. U.S. Nat. Mus. vol. x]. 1911, p.637. Also used as‘a character

of “Tenia argentina”’ (believed to be synonymous by Zschokke, Centralb. Bakt. u.

Paras. i11. 1888, p. 1).

AVIAN CESTODES. 433

serves, as I imagine, as a retractor of the cirrus. The vagina

opens at the junction of the neck and the flask-shaped region, a

fact which rather tends to prove that the neck region really

belongs to the cloaca genitalis, unless it be remembered that the

male and female genitalia have really a common origin and are

parts of a single sytem. ‘The various facts to which attention

has been called are illustrated in the accompanying figure of the

cirrus-sac and its external orifice (text-fig. 2).

Text-figure 2.

Horizontal section through cirrus-sac.

Tn the lower figure the junction of the two parts of the cirrus-sac is shown

more highly magnified.

g-c. Genital cloaca. c. Cirrus. v. Muscular valve-fold separating the narrow

part of the cirrus-sac from the wider region. 7. Water-vascular tube.

The earliest appearance of the uterus is represented in text-

fig. 3. It is there seen to consist of a modified tract of medullary

tissue lying anteriorly to the male and female gonads, at about

434 DR. F. E, BEDDARD ON

the middle of the proglottid and on a level with the cirrus-sac.

It forms a nearly straight line about the middle of the proglottid,

wider in the middle of its length and tapering off to the side of

the proglottid remote from the cirrus-sac, in which direction it

does not reach the lateral water-vascular tube. At the other end

the growing uterus is limited by the cirrus-sac and the coil of the

vas deferens. In the lateral, most immature part of the uterus

that organ appears to be formed by only a very slight modification

of the medullary tissue. There is simply an increase in the

number of the nuclei, which are thus closer together and become

more conspicuous,

Text-figure 3.

Horizontal section through immature proglottid, to illustrate earliest appearance

of uterus (wé.). ov. Ovary.

In the more central parts of the uterus at this stage, which, as

already said, is wider, the nuclei are more abundant, both actually

and relatively. Between them cavities appear, all of them small

but of varying sizes; the dimensions of some coincide very closely

with the meshwork of the general medullary parenchyma. The

nuclei, at least in many cases, are arranged round these spaces in

a regular fashion and thus constitute a lining epithelium to these

uterine cavities. The several cavities are separate from each other

in this early formed uterus. Fuhrmann* has observed the same

mode of origin of the uterus of Chapmania tauricollis, and

remarks: “ Anfangs erscheint der Uterus gekammert.”

In later stages there is a confluence of these cavities, and in

the fully developed uterus such as is represented in text-fig. 4,

the uterus appears to consist of a larger central cavity which is

prolonged in all directions through the medullary parenchyma

into outgrowths. Jam disposed to think that these latter inter-

communicate and form a network. In horizontal sections near

to the dorsal and ventral surfaces of the uterus the retiform

appearance is very strikingly manifested. In transverse sections

* Rey. Suisse Zool. iv. 1896, p. 121.

_ AVIAN CESTODES. 435

through the mature uterus the appearance of a number of detached

cavities containing ripe eggs is often presented. It is, perhaps,

this condition which has led to the explanation by v. Linstow *

of the clusters of ripe eggs seen in “ Tenia struthionis.”

Text-figure 4.

Horizontal section through mature uterus, showing opening into paruterine

organ.

Ca. Mass of calcareous bedies at orifice of uterus. 7. Lateral water-vascular tube.

Pa, Paruterineforgan. Ut, Uterus.

The paruterine organ of this species has been figured by

* Arch. mikr. Anat. xlii. 1893, pl. xxviii. fig. 14.

436 DR. F, E. BEDDARD ON

Zschokke * in the course of his description of “ Tenia argentina,”

but mistaken by him for a testis, with which identification

Monticelli associated himself +. Fuhrmann {recognised the true

nature of this body, which was certainly puzzling at a time

when the paruterine organ, now known in so many Cestodes,

was hardly or not at all understood. Zschokke has rightly called

attention to the mass of calcareous bodies lying behind the

anteriorly placed paruterine organ. The first beginnings of this

organ are not plain to me, so little differentiated are at first its

tissues from that of the medullary parenchyma of which it is a

part. But I feel safe in saying that it does not put in an

appearance for some time after the uterus has commenced to

develop. I mention this matter as being of importance, since in

the species Rhabdometra cylindrica the paruterine organ appears

before the uterus. There is thus in the present species no

obvious connection in development between thetwo organs. The

paruterine organ in a fairly early stage of development lies, as has

been stated, anteriorly in the segment and quite close to the

anterior edge of the uterus, which is behind it. This surface of

the paruterine organ is capped by a rather dense mass of calcareous

bodies. The calcareous bodies are not, however, confined to

this region of the paruterine organ, or rather to the outside

of it; they also occur scattered throughout its substance, but

not in such great numbers and not, where present, so closely

pressed together. The general outline of the paruterine

organ in these not fully mature segments is shown in horizontal

sections to be somewhat conical, but with a rather convex base,

the latter being anterior in position.

It is, furthermore, to be noted that the paruterine organ is very

closely related to the uterus, which lies behind it. The mass of

caleareous bodies and the margin of the mass forming the

immature paruterine organ are divided by nothing from the

uterine cavity. The uterus, that is to say, has no anterior wall

save that which is furnished by the paruterine organ. There

is thus a distinct relationship between the uterus and the

paruterine organ. At this period in its development the

paruterine body is solid throughout ; there is no trace of a central

cavity. In proglottids at the end of the body, which are rather

longer than broad and apparently quite mature, the paruterine

body has the appearance which is represented in the accompanying

figure (text-fig. 4). This is a representation of a horizontal

section showing the paruterine body rather oblong in form with

rounded angles. It is sharply marked off from the parenchyma

of its proglottid laterally. Posteriorly it is not marked off from .

the cavity of the uterus, that is to say the uterus has no wall of

its own dividing it from the paruterine organ. The two structures

indeed seem to be mutually differentiated parts of one structure.

* Centralb. f. Bakt. u. Parasit. iii. 1888, p. 1.

+ Nat. Sicil. xii. 1892-3, p. 208.

t Rey. Suisse Zool. iv. 1896, p. 122.

AVIAN CESTODES. 437

The paruterine part is about one-third of the total length of the

egg-holding receptacle. The walls of the paruterine body are

thick, and the fibrous-looking tissue has the strands running

along the greater length of the body. Externally it is plainly

marked off from the tissue of the medullary part of the proglottid.

The mass of calcareous bodies is as plain here as in the younger

paruterine organ; but it has got a more definite coherence of its

own; the corpuscles are imbedded in a tissue which forms a cap

to the paruterine organ and protrudes into the cavity of the

uterus behind. The suggestion is that of a valve which, however,

hinders the flow of ova into the paruterine organ rather than the

converse, which is what would be expected.

Text-figure 5.

Transverse section through mature proglottid, showing paruterine organ.

ce. Central cavity of paruterine organ. 7, Lateral water-vascular tube.

The close relationship of the paruterine organ to the uterus

is reminiscent of the conditions which obtain in the not nearly

allied genus Jesocestoides. In this Cestode, which has been

investigated by Hamann *, Fuhrmann7, and some others, the

uterus is divided into two regions which form a continuous tube.

Posteriorly the uterus acquires thick fibro-spongy walls and serves

* Zeitschr. f. wiss. Zool. 1885, + Swedish Exped. Egypt, pt. iii.

438 DR. F. E, BEDDARD ON

as the ultimate receptacle of the ripe eggs, being closed to form a

spherical capsule. It is thisregion which suggests the paruterine

organ of Chapmania, a comparison which Fuhrmann seems, in

the memoir quoted, to hold as possible, though Hamann compares

this swollen and metamorphosed part of the uterus with a shell-

gland (a comparison which seems to be negatived by the discovery

of an ordinary shell-gland by Zschokke and others). In no

genus, however, is there quite so intimate a connection between

the uterus and what is certainly a paruterine organ as in

Chapmania. It is quite likely that JMJesocestoides has preserved

the original relationship between these parts of the reproductive

system.

The access, therefore, of the ripe eggs in the uterus to the

paruterine is thus assured; and the drawing to which I have

referred (text-fig. 4) shows this movement in progress. The

eggs occupy the central hollow region of the paruterine organ,

and are to be seen in transit in various parts of the same and

of the uterus. I have observed eggs entangled, as it were, in

the lax tissue forming the plug of calcareous bodies*. There is

no doubt but that here the transference of ova to the paruterine

from the uterus is quite direct. They could hardly reach it by

another route, in view of the free continuity of the two sacs.

Nevertheless, another view has been advanced by Fuhrmann 7.

in his important résumé of the genera of Cestodes found in

birds, this author remarks in the definition of the genus Chap-

mania—* Die Hier gelangen in einen stark verzweigten Uterus

und von da wie bei Davainea in Parenchymkapseln, worauf sie in

abgelosten Gliedern in ein am Vorderrand gelegenes breites, grosses

Paruterinorgan gepresst werden, das eine Kapsel um sie bildet.”

This definition is accepted by Ransom, who practically translates

it in his general survey £ of the Cyclophyllidea. Ina later and

fuller table of distinctions of the genera of Davaineide § this is

altered. In the latter, Ransom says (as part of his definition of the

genus Chapmania), ‘Eggs pass anteriorly into a paruterine organ

from the uterus either directly or after the disappearance of the

uterine wall and the envelopment of the eggs in individual paren-

chymatous capsules.” This alternative statement as to the fate of

the ova is apparently due to an earlier definition by Fuhrmann of

the genus Chapmania ||, which runs (so far as concerns the matter

under discussion) as follows—‘“‘ Die Hier, statt im Parenchym zu

* It may be pointed out that, in his figure of Chapmania longicirrhosa (later

regarded as identical with Idiogenes flagellum), Fuhrmann (Centralb. f. Bakt. u.

Parasit. Bd. 41, p. 81, fig. 3) represents a mass of calcareous bodies such as occurs

in Chapmania tauricollis, but upon the opposite side of the paruterine organ, 7. e.

upon its anterior face. If this be not an error there is perhaps here an additional

point of distinction between the two genera.

+ Zool. Jahrb. Suppl.-Bd. x. t Bull. U.S. Nat. Mus. no. 69, 1909.

F § “A New Cestode from an African Bustard,” Proc. U.S. Nat. Mus. vol. xl.

p. 646, 1911.

= || Centralb. f. Bakt. u. Parasit. Bd. 41, 1906, p. 83. This memoir, however, is

not quoted by Ransom, which is merely an oversight, as he refers to it in

a footnote enumerating the synonyms of Idiogenes flagellum.

AVIAN CESTODES. 439

zerstreuen, in ganz reifen losgelésten Proglottiden in einem

parenchymatosen Paruterinorgan, das zu einer Uteruskapsel

wird, vereinigen.” I believe, however, that this definition was

made to include the species described in that memoir as Chap-

mania longicirrhosa, a species which Fuhrmann later * trans-

ferred to the genus /diogenes, and to the species J. flagellum.

I have been quite unable to find a trace of anything like the

‘“‘ Parenchymkapseln” of Davainea or any other genus in which

such structures exist. Nor do I think that it would be easy to

miss such bodies were they, at any rate, so conspicuous as in the

genera which are known to possess them (e. g., nermicapsifer, etc.).

My figure is, as I think, decisive as to the direct entry of the ripe

ova into the paruterine organ; in no other possible way can the

facts observed, and there represented, be explained. I can only

suggest that the irregular form of the uterus in ripe proglottids,

as I describe it later, may be responsible for the statement

that the eggs are separately envolved in parenchymatous

capsules before being pushed into the paruterine organ. For in

many sections ova may be seen to lie apparently in closely fitting

capsules, these being in reality the expression of the ramifying

branches of the uterus. Or it may be that both methods occur

in this species, or finally, there is the possibility, which I do not

consider to be very great, that the worm which I deal with in

the present communication is not Chapmania tauricollis but a

new form.

The last word about the paruterine organ of this Cestode is

contained in Fuhrmann’s account of Chapmania tapicat. Here

that author states that “ Bei Chapmania (auch bei Chap. tawri-

collis) geschieht dieser Uebertritt der Hier erst in abgelisten

Gliedern und ist deshalb von andern Autoren noch nie beobachtet

worden.” It is clear, from my own observations, that the eggs

reach the paruterine organ at an earlier period. I think, however,

that in my species, Ofiditenia eupodotidis, plainly belonging to

this subfamily, this late transference does occur.

Systematic Position of Chapmania tauricollis and validity

of genus Chapmania,

There is, of course, no need to argue the position of this tape-

worm so far as concerns its family and subfamily position. It is

clearly a member of the subfamily Idiogenine of the family

Davaineide.

This subfamily contains three genera, viz., Jdiogenes, Chap-

mania, Sphyronchotenia, and very possibly my genus Otiditeniat.

The most recent survey of the characters of the three former

* Zool. Jahrb. t. cit. p. 50.

+ Swedish Exped. Egypt, pt. iii. p. 23.

t See P. Z.S. 1912, p. 194, and ibid. 1914, p. 879.

440 DR. F. E. BEDDARD ON

genera is by Ransom *, who distinguishes the three by the

following salient characters—which I withdraw from his fuller

diagnosis :—

Idiogenes.—Small worms with weak musculature. Genital pores

unilateral (except in J, otidis). Suckers unarmed. Cirrus-

sac large. Eggs pass directly into paruterine organ.

Chapmania.—Larger worms with strong musculature. Genital

pores unilateral (in C. tawricollis). Suckers armed. Cirrus-

sac not large. Eggs first developed in separate egg-capsules

and then passed into paruterine organ.

Sphyronchotenia.—Larvger worms with strong musculature.

Genital pores unilateral. Suckers unarmed. Cirrus-sac

not large. Eggs pass directly into paruterine organ 7.

Many rows of hooks on rostellum.

We may leave aside Sphyronchotenia, which is clearly a

separate genus—not to be confounded with either Jdiogenes or

Chapmania. With regard to the two latter the differentiation

deducible from the above characteristics is altered to some extent

by the new facts recorded in the present communication. In the

first place, the cirrus-pouch of Chapmania is not particularly

small as is alleged by Ransom. I take it that the American

helminthologist has been misled by the absence of any statement

about the cirrus-sac of Chapmania in Fuhrmann’s definition of

the genus, while, on the contrary, /dzogenes is defined by a large

sac. But the latter author, in his description of “ Davainea

tauricollis,” remarks { that the cirrus-sac reaches to the middle

of the proglottid, a statement which I confirm from my own

observations §. There is, therefore, here no difference between

Idiogenes and Chapmania. I have discussed above, in detail, the

statement that Chapmania differs from Jdiogenes in the fact

that the former genus shows a series of egg-capsules in which the

ova are imbedded before their transference to the paruterine

organ, and shown that there is no such difference between the

genera. There remains, therefore, merely the difference of size

and the stouter build of Chapmania, which is caused by the

relatively and actually greater thickness of the longitudinal

muscular layer of the body, to form a basis of distinction from

its ally Jdiogenes. It is not at all impossible to regard these

facts as of generic value; but it must be remembered that

* Proc. U.S. Nat. Mus. vol. xl. 1911, p. 637.

+ This is to be inferred, as the actual transference was not seen in any of its

stages by Ransom.

t Rey. Suisse Zool. iv. 1896, p. 119. '

§ But the exact point to which the cirrus-sac reaches is affected by the degree

and direction of the contraction of the particular proglottid examined. It is

possible that the generic distinction has been chiefly founded upon Chapmania

tapica, where, according to Fuhrmann (Swedish Exped. Egypt, pt. ii. p. 23), the

cirrus-sac hardly reaches the water vascular vessel.

AVIAN CESTODES. 441

variations in the longitudinal musculature are by no means

always accompanied by other structures which imply undoubted

generic difference among the Cestodes. It is possible, however,

that a simple sac-like form of uterus characterises /diogenes *.

I should be, on the whole, disposed to define the genera of the

subfamily Idiogeninz as follows :—

Subfamily /D10GENINE.

A single paruterine organ present in the ripe proglottid.

( SPHYRONCHOTENIA. — Suckers unarmed

| Genital pores unilateral. Longitudinal

muscles thick. Dorsal water-vascular

trunk absent in posterior proglottids.

| Testes numerous, extending far anteriorly.

| Uterus divided into chambers, larger than

| paruterine organ. Lggs transferred late

ie paruterine organ. Currus-sac small.

OrmirantA.—Suckers unarmed. Genital

pores alternate. Longitudinal muscles

thick. Dorsal water-vascular trunk present

in posterior seyments. Testes numerous,

posterior, Uterus divided into chambers,

extends dorsally into cortical layer, much

smaller than paruterine organ. LHggs

transferred late to paruterine organ.

Cirrus-sac small.

CuapuaAnrAt.—Suckers armed. Genital

pores alternate. Longitudinal muscles

thick. Dorsal water-vascular vessel absent

in posterior proglottid. Testes numerous,

posterior. Uterus divided into chambers,

which in mature uterus communicate and

form a network, larger than paruterine

organ. Hggs transferred early to paru-

terine organ. Cirrus-sac large.

Iprocenrst.—Suckers unarmed. Genital

pores unilateral or alternate. Longitu-

dinal muscles slight. Testes few, pos-

terior. Uterus not divided into chambers,

larger than paruterine organ. Cirrus-sac

very large.

A. Ten to twelve

rows of hooks on %

the rostellum.

(

B. Two rows of |

hooks on the ros-

tellun.

* See also footnote to p. 438 for another possible generic distinction.

+ In view of the network which the sperm-duct forms and the tentacles upon the

scolex, I exclude for the present C. tapica from the above genus.

+ It will be noted that the feebly developed longitudinal muscular system, the

small number of testes, and the simplicity of the uterus may be correlated with the

small size of the species of this genus.

442 DR. F. E. BEDDARD ON

I may point out, in conclusion, that the subfamilies Idiogeninz

and Davaineine, have corresponding relations to Cestodes

commonly referred to different families by reason of their lack

Text-figure 6.

Horizontal section through immature proglottids, showing position

of gonads.

c. and ¢. Cirrus-sac. J.v. Lateral water-vascular tube. o. Ovary. Ut. Testes more

thickly disposed on the side away from the pore. 7Z.R. Transverse water-

vessel. v.g. Vitelline gland.

of rostellar hooks. There is, fcr instance, a close resemblance

between Rhabdometra and its allies and /diogenes, which possesses a

single paruterine organ, on the one hand, and between Zschokkeella

AVIAN CESTODES. 443

and Jnermicapsifer and Davainea, with many ‘ ege-capsules,” on

the other. In these instances the loss of the rostellar hooks would

need the reference of Davainea to either Zschokkeella or Inermi-

capsifer, and of Idiogenes to Rhabdometra or one of its near allies.

The relation is quite like that between T'entarhyncha (sens. strict.)

and Tenia. In Teniarhyncha, a genus formed to include the

hookless Tenia saginata, we have a typical Tenia, differing only

by the character mentioned and the correspondingly reduced ros-

tellum. It is, therefore, very important to ascertain positively

the presence or absence of rostellar (and other) hooks in view

of the above relations. This task is—my experience with

Chapmania teaches me—not always easy. I have carefully

re-examined the scolex of my species Zschokkeella gambiana in

order to set doubts at rest. I am still unable to find hooks

thereon, and cannot, sligiaeioe, alter my opinion of its systematic

position.

Proc. Zoot. Soc.—1915, No, XX XT, 31

Pr, Zo S IIB, PIXCTELIL GOODRICH, Pl. I.

MINCHINIA.

P, Zo So 1915, POCEILL-GOODRuGisl, |P ll, IU

MINCHINIA.

ON THE PARASITE MINCHINIA. 445

34. Minchinia: A Haplosporidian. By Hexen L. M.

Prxeti-Goopricu, B.Sc., Beit Memorial Research

Fellow *.

[Received April 9, 1915: Read May 11, 1915.]

(Plates I. & II.7)

The genus Minchinia (Labbé) undoubtedly belongs to the

Haplosporidia, although it has invariably been placed among the

Coccidia. Schaudinn (17, p. 276) went so far as to include the

genus in that of Adelea, and in this he was followed, tentatively

only, by Doflein (7, p. 744).

Minchinia chitonis Lankester is the only well-established

species ; other so-called species will be dealt with later. This

parasite was first recorded so long ago as 1885 by Lankester (11,

fie. 12), who found it in the liver of Chiton and gave a drawing

of the spore under the name of Alossia chitonis. Labbe (9) ex-

amined the same parasite in 1896, and, recognising its distinctive

characteristics, made it the type species of a new genus, which he

named Méinchinia in honour of Prof. Minchin. The spore has a

thick chitinous coat and an outer membranous envelope produced

into a long tail at either end.

In 1897 Leger (18) described a new Coccidian Barroussia

ca1udata whose spore was said to be provided with a long tail at

one of its poles, though, so far as ] can make out, 1b has never

been figured. In a subsequent communication Léger (14, p. 7

footnote) pointed out that this form, parasitic in Lithobius martin,

had affinities with Minchinia, though he did not recommend

connecting the two forms. In spite of this, Labbé in 1899 (10)

included this species of Barrowssia in his new genus as Minchinia

caudata. In 1898 Léger (15) described the flagellated micro-

gamete belonging to this species, which is undoubtedly Coccidian

in type. Therefore Barroussia caudata, a true Coccidian, is no

longer to be included in the genus Minchinia.

Patella and Trochus were also given by Labbé as hosts of

Minchinia, although he seemed undecided whether or not these

contained the same species as Chiton. No figures of these

parasites were given except one of a trophozoite from Patella

which appears to be Coccidian in character. Therefore it seems

likely that this Patella parasite may also turn out to have no

relationship with Minchinia. The specimens of Patella and

Trochus that I have examined have not been infected at all,

although some specimens have been taken from exactly the same

locality as infected Chiton.

* Communicated by the SECRETARY.

+ For explanation of the Plates see p. 456.

446 MRS. H. L. M. PIXELL-GOODRICH ON

OccuRRENCE OF J/INCHINIA IN CHITON.

During the last four or five years I have searched Chiton from

Plymouth and other parts of the British coast, as well as from

parts of the Pacific and Mediterranean, for traces of this parasite,

and it was only in October last that specimens of Chiton at

Plymouth were found to be strongly infected. The species of

Chiton so parasitised was Craspedochilus cinereus (Linn.)* from

Rum Bay, not Acanthochites (Chiton) fascicularis (Linn.), stated

by Labbé to be the host at Roscoff. In fact the latter species I

have never found to be infected. From this locality, out of 135

specimens of Craspedochilus cinereus examined, 85 have had a

more or less advanced infection. This proportion of infected

specimens, roughly two-thirds, does not coincide with Labbe’s

description of M/inchiniaas a rare parasite. However, these figures

only refer to specimens of Craspedochilus cinereus from a small

stretch of coast near Plymouth called Rum Bay. Perhaps the best

ground for the collection of Chiton in the Plymouth district is

the mouth of the river Yealm ; here, however, the specimens are

small only, and out of twenty-five not a single one has been found

to be infected with Minchinia. Chiton (two only) examined from

Wembury Bay have also been free from infection.

The infection is thus shown to be very localised ; whether the

Chiton in Rum Bay are suffering from an epidemic of this

parasite, or whether MZinchinia is endemic to this locality, could

only be determined by an investigation extending over several

years.

The parasites have so far only been recorded from the liver ;

they are, however, not restricted to this organ, but later overrun

all the connective tissue. ‘They are especially numerous between

the lobes of the liver and the interior projections of the walls

of the so-called sugar-glands (Zuckerdriisen). Infection then

spreads to the wall of the gonad, radula sac, gills, and blood-spaces

in the foot. Cases of advanced infection can nearly always be

detected without dissection, because cysts full of spores can be

seen in the gills and through the epidermis of the foot.

Insuch favourable positions, lying immersed in the host’s blood,

the parasite has every opportunity of nourishing itself, and

during its rapid endogenous multiplication the liver becomes

gradually deprived of its reserve food materials. In early cases

of severe infection the presence of numerous colourless plasmodia

and young cysts gives a white appearance to the liver, which is

normally brownish. As the cysts become filled with brown

chitinous spores they appear black by reflected light, and conse-

quently a liver with a severe infection at an advanced stage is

quite black, as well as being enormously enlarged owing to the

bulk of parasites between the lobes. The liver-cells in such

* [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. |

THE PARASITE MINCHINIA. AAT

advanced cases were found to have lost their reserve food and to

be much reduced in size. In spite of the necrotic condition of

this organ, however, the life of the Chiton appeared to be little

affected—at any rate, in captivity infected specimens sometimes

lived longer than non-infected. No observations appear to have

been made on the normal length of life of a Chiton, possibly it

may be great. Quite small specimens are often sexually mature ;

on the other hand, strongly infected specimens are nearly always

very large—some of them the largest specimens ever seen at

Plymouth. Could it possibly be that development of MWinchinia

in a Chiton causes the whole animal to hypertrophy! On the

whole, adult forms were better infected than young ones, as would

be expected. J have not seen the earliest stages of infection, but

presumably the Chiton become infected when feeding: they are’

said to be entirely vegetarian. Labbé has figured three (10, text-

fig. 107) so-called sporozoites in the liver-cells; possibly one of

them is something of the kind, the other two appear to be the

normal contents of the liver-cells. Many experiments by which

J have tried to infect Chiton artificially have given negative

results. Ripe spores have not been induced te open when intro-

duced into the wsophagus or placed in fluid from different regions

ot the alimentary canal. Therefore we seem forced to the

conclusion that there is another host; this, at the beginning,

seemed unlikely: first, because the ,spores as developed in

Chiton are so admirably adapted for an external free existence,

and secondly, it is not easy to see how such an animal as Chiton,

which feeds on plants, could easily become infected from other

animals. Certain other facts in this connection are given below

after the description of the spores.

MeETHODs.

Bouin’s picro-formol-acetic mixture was found to be the best

fixative for infected tissue, since this penetrates to a certain

extent even into ripe spores. The special methods applied to these

spores will, however, be described later when considering their

structure. For general purposes picro-nitric and corrosive

sublimate mixtures were also satisfactory. Owing to the small

size of gametes, sporoblasts, and young spores, 1t was necessary to

have thin sections (3-4 ), and when there were many chitmous

spores this was not easy. Breaking of sections was sometimes

prevented by painting with collodion. For following the

development of the spores it is essential to have living material,

and methyl-green and acetic acid mixture was used for temporary

preparations. Numerous coverslip films and smears stained

with Giemsa were also used for studying the development. For

staining films and sections iron hematoxylin or hematein methods

are of course most useful; water solutions certainly seem to give

better results than the alcoholic ones recommended by Lee (12,

p- 156) and others, though by the latter staining is more speedily

effected. The chitinous substance forming the spore coat stains

448 MRS. H. L. M. PIXELL-GOODRICH ON

very intensely by these methods, which are therefore not always

convenient. In these cases Ehrlich’s, Mann’ s, or Delafield’s

hematoxylins have given better results. Carmine stains have

always proved disappointing. As a rule counter-staining was

not found to be an advantage, the cytoplasm being sufficiently

stained by the hematoxylin.

PLASMOTOMY.

All Chiton examined from October 1914 to April 1915 were

heavily, if at all, infected. Im only two cases was endogenous

multiplication still proceeding, and these specimens were obtained

and fixed during October. One of these (Chiton V) contained

also many stages in sporogony and even some ripe spores; the

other (Chiton X) was crowded chiefly with plasmodia, which gave

a white appearance to the liver. Most of the plasmodia were

irregular in outline, but they varied much in shape and size

(Pl. L. figs. 1 & 2). Scattered through them were nuclei con-

sisting of masses of chromatin, round each of which could be

distinguished a clear space. At this stage the nucleus has no

sign of amembrane. Frequently these nuclei were dividing by

a simple method of mitosis. In vain have J searched through

hundreds of sections to find any trace at this stage of the beautiful

spindles which have been figured for some Haplosporidia (2) (18).

Swarezewsky states that Chatton (5) must have missed them in

Caullerya mesnili; but such does not necessarily seem to be the

case, for I do not think that they can have been passed over in

Minchinia.

When about to divide, the mass of chromatin separates into

two parts, which gradually move away from one another, but for

some time are connected by a slender thread (centrodesmose).

It has not been possible to demonstrate the centrosomes con-

tained in these chromatin masses by means of differential

staining, although one would expect them to be present.

When a plasmodium has attained a certain size, or from other

causes, it breaks up into daughter plasmodia, each with several

nuclei (fig. 2). All these trophic stages are quite naked and

their outlines generally irregular. It seems clear that they flow

along to some extent, pr obably only very slowly, by putting forth

pseudopodia. Sometimes between closely apposed lobes of the

liver a plasmodium becomes very elongated and narrow (fig. 1).

No doubt want of space often determines the breaking up into

daughter plasmodia, It is rare to find one longer than 100 pe or

wider than 50 fle

During this endogenous multiplication of the parasite the

host’s phagocytes become very active and multiply rapidly, some-

times forming clumps. They appear to make a vain attempt to

engulf the parasites, and occasionally one is observed flattened

against a plasmodium.

There is a certain amount of evidence to show that solid

THE PARASITE MINCHINIA. 449

particles can be ingested by the plasmodia as by an ordinary

amoeba. Masses of fibres staining black with iron hematoxylin

may sometimes be seen enclosed in an active plasmodium, They

either disappear before encystment, or sometimes they may be

extruded in a mass directly after the formation of the cyst, and

become compressed between it and the parasite. Further, certain

deeply staining granules are generally contained in the cytoplasm,

round which no clear space can be observed as round a nucleus.

These granules are easily distinguished from nuclei when the

latter assume the vesicular form after encysting (PI. I. figs. 1

& 3, gr.), although it is almost impossible to distinguish them

before this change is affected. Caullery and Mesnil (2, fig. 11)

saw similar chromatic bodies left over after the nuclei had become

vesicular, but interpreted them as degenerate nuclei. In

Minchinia they seem likely to be unassimilated particles, possibly

remains of host-cells, taken in during the active life of the

plasmodium. In one case it was practically certain that the

parasite had ingested a host-cell—apparently a phagocyte! In

many cases these cells apply themselves so closely to a plasmodium

that it is difficult to see any boundary between them (fig. 2).

Cuénot (6) has recently given some useful information about the

activities of the phagocytes of Chiton, and usually there is no

difficulty in distinguishing the nucleus of one of these cells from

a parasite nucleus. In Minchinia, however, certain plasmodia

occasionally appeared at first sight to be giving off buds in which

the nucleus had assumed a vesicular form; but after further

study I am convinced that these specimens were really only

plasmodia to which host-cells were endeavouring to attach them-

selves. Gemmation has, however, been described by Swarezewsky

(18) as a mode of reproduction, in addition to plasmotomy, in

certain Ichthyosporidia.

SPOROGONY.

(1) Formation of gametes.

When the host has become strongly infected most of the

plasmodia draw in their pseudopodia and become rounded off and

encysted. It seems possible, from the consideration of after-events,

that two plasmodia may mingle before encysting, but no direct

evidence has been forthcoming on this point. A thin pellicle is

secreted, but in most cases this is soon covered and obscured by

the host’s amcebocytes, which apply themselves to it, becoming

gradually flattened out to form acellular cyst around the parasite.

This cyst is generally only one cell thick (PI. I. figs. 3 & 4), and its

nuclei become more and more flattened with growth. Occasionally

there may be four or five layers of cells forming a thick cyst

round a parasite, but this is an abnormal condition. The cysts

are spherical, and vary very much in size, but they are not

generally more than 75 y, or less than 40, in diameter. The

nuclei assume a vesicular form, and probably enter upon a resting

450 MRS. H. L. M. PIXELL-GOODRICH ON

stage. This change is affected by the breaking up of a mass of

chromatin into granules, some of the smaller ones of which pass

to the periphery, where a distinct membrane appears. The

central chromatin generally forms two or three distinct masses

often connected with the membrane by fine strands. These

vesicular nuclei are more or less oval, and their longer diameters

are slightly less than 3 p.

Occasionally, forms have been seen in which the nuclei are

massed in the centre, leaving a clear peripheral border of

cytoplasm. Inaddition there is sometimes a cavity in the middle,

but the nuclei are not arranged in a definite single layer round

this cavity as described by Alexeieff (1, p. 36, fig. 4) for his

“‘plasmodes blastuloides” in Jchthyosporium gasterophilum. In

Minchinia the occurrence of the central cavity, at any rate,

appears to be an artifact.

When preparing to divide a nucleus of this vesicular type in-

creases in size and a bundle of achromatic threads appears along

the greatest diameter acting as a kind of spindle. The chromatin,

after arranging itself on this spindle, is drawn to the poles. The

vesicle then constricts in the middle, and the two ends are finally

nipped off as daughter nuclei. ‘This process has been beautifully

figured by Granata (8, Plate 3) for Haplosporidium limnodrili, in

which, according to the magnification given, the nuclei are con-

siderably larger than those of MJ/inchinia.

After a time the cytoplasm separates and collects round these

nuclei either singly or sometimes in clumps of from two to seven

or eight. Occasionally a dozen or so different sized masses have

been seen inside a cyst and have been set free by bursting it.

When compressed, these masses are seen to contain varying

numbers of nuclei. Ultimately, however, such multinuclear

masses are resolved into uninuclear bodies: which a appear at times

to be distinetly ameeboid. ‘These are the gametes. No residuum

is left over except the chromatoid granules, which do not become

transformed into vesicular nuclei as already described.

(2) Syngamy and formation of zygotes.

The gametes proceed to pair. Fusion of their cytoplasm first

takes place to form a single body with still separate nuclei (PI. I.

fig. 5 a) for which the term Prozygote has been proposed (16).

Caullery and Mesnil (2) hesitated to give a definite opinion as to

whetber similar bodies with two nuclei in some of their Haplo-

sporidia represented forms undergoing division or gametes under-

going syngamy, because they also found masses with four nuclei

in some cases. The same difficulty presents itself in MWinchinia ;

but, after prolonged study of living stages and careful measure-

ments of the different sized bodies and nuclei contained in cysts,

the above seems the only possible interpretation. Caullery and

Mesnil’s forms with four nuclei (2, fig. 43) seem to be stages in

the formation of the sporoblasts as described below (figs. 7 & 8)..

THE PARASITE MINCHINIA. 451]

After some time has elapsed the gamete nuclei in the prozygote

approach one another (fig. 5 )and fuse to form the syncaryon

(fig. 5 ce). The zygote so formed appears to be produced by

autogamy. Such may not be really the case, however, for it is

quite possible that the gamete nuclei may have been derived from

distinct parents. Hither two plasmodia during their wanderings

may have come together and their cytoplasm fused before

encystment (plastogamy), or there is some evidence in other

Neosporidia to show that the amebule fuse in pairs on their

escape from the spore. In the latter case probably their nuclei

do not fuse but divide independently, and ultimately syngamy

takes place between gamete nuclei formed in equal numbers from

each parent.

(3) Lormation of spores.

The zygotes generally proceed almost at once to divide twice

(Pl. I. figs. 6 & 7) to give four sporoblasts: division is effected

here again by a simple method of mitosis. As a rule, the nuclei

go through both their divisions before the cytoplasm divides, so

that the stage with four nuclei (PI. II. fig. 8) is quite common,

but sometimes division of the cytoplasm follows after the first

nuclear division. Also,these divisions are not always simultaneous

even in one and the same cyst, therefore there may be enclosed

together with free sporoblasts masses with two, three, or four

nuclei, and even an undivided zygote. This fact added much to

the difficulty of elucidating the life-history of Minchinia.

The sporoblasts when first separated are very small and their

protoplasm rather vacuolar. They soon begin to secrete a mem-

brane which is produced into a short tail at either end (PI. II.

figs. 10&11). The young spore then grows considerably: presum-

ably the membrane is so thin that it does not prevent the

absorption of food. While still quite small its nucleus divides,

giving off a parietal mass of chromatin (figs. 9a & 96) which

sometimes divides into two (fig. 9c) These parietal nuclei or

masses of chromatin, for possibly they are not true nuclei, are

very distinct at this stage. Their prominence, however, is very

transitory, for soon after they are masked by numerous other

chromatic bodies which are given out by the main nucleus.

Probably the parietal ‘* nuclei ” themselves also break down into

similar chromatic granules or globules. These ave highly re-

fringent and pass to the periphery, where they arrange themselves

to form the chitinous spore coat, as will be more fully explained

later.

At the stage when these globules are passing out in a centri-

fugal direction there arises in the cytoplasm, near the nucleus, a

homogeneous finely granular spherule (Pl. I. fig. 14). I have

no evidence that this comes actually from the nucleus as described

by Granata (8) for Haplosporidiwm limnodrili. The spherule is

distinguished by the fact that it stains only slightly and takes up

its position just underneath the operculum. Unripe spores open

452 MRS. H. L. M. PIXELL-GOODRICH ON

fairly easily when treated with certain reagents, methyl-green

acetic mixture forexample. From a young spore under these con-

ditions the spherule makes its way out as soon as the operculum

is raised. Similar bodies have been mentioned as occurring in

many Haplosporidia. In Minchinia the spherule is very large at

the stage when the spore coat is nearly completed (figs. 13 & 14),

and after this appears to gradually diminish, until in the ripe

spore it is generally not to be distinguished at all. Possibly it is

composed of some kind of reserve food on which the developing

spore can feed as soon as it is cut off from the outer world by the

formation of its thick chitinous coat.

During the deposition of the substance which forms the chitin-

ous coat there is a considerable shrinkage of the nucleus (figs. 12,

13, 14). Although the substance appears to corfe directly from

the nucleus it is not presumably similar to chromatin In com-

position. In the living it is more refringent. When the globules,

which arrange themselves at the periphery just inside the spore

membrane (fig. 14), are sufficiently numerous they begin to run

together. Ultimately a continuous layer one micron in thickness

is formed all round the spore (figs. 13 & 15). This is at first

colourless, but later becomes ight brown but remains translucent.

In its behaviour to certain stains this substance also differs from

chromatin. Although it stains densely black with iron hema-

toxylin and red with saffranin, the more selective nuclear stains

such as Ehrlich’s, Mann’s, and Delafield’s hematoxylins do not

stain it nearly so intensely aschromatin. In Giemsa preparations

the membranous covering of the spore generally stains red and

the inner chitinous coat blue, not red like the nucleus.

The chitinous substance is very resistant: like true chitin, it

is not dissolved by boiling in strong caustic potash (30°/,); but,

on the other hand, it does not acquire the characteristic mauve

colour shown by chitin with the iodine in potassium iodide and

zine chloride test. ‘Thus it must be concluded that while closely

resembling, it is not identical with ordinary chitin.

(4) Spores.

The ripe spores vary a good deal in size but are always oval,

and when living generally about 10 p long and 6 p wide. The

largest measured was 13 p long and 8 w wide. The latter giant

spores seem to be distributed promiscuously ; that is, there is no

distinction of cysts into those containing macrospores and micro-

spores. Normally the tails into which the outer membrane of

the spore is produced are about four times the length of the

spore, but they are more or less brittle, and often become broken

off. The chitinous coat is thick and very resistant. It can

be softened in various ways, e.g. immersion in Eau de Javelle

or weak formalin, in order to make it sufficiently permeable

for its contents to be stained. Formalin (4°/, formaldehyde) is

THE PARASITE MINCHINIA. 453

especially good for this purpose, and when followed by Ehrlich’s

hematoxylin enables the spore nucleus to be well stained.

Prolonged immersion in distilled water had no apparent effect on

the spore. Jodine stained the chitinous coat yellow but otherwise

had no result. Adult spores were occasionally induced to open

by pressure and reagents combined (PI. LI. fig. 16). The oper-

culum always opened away from the tail and turned inside out,

but remained attached by the outer membrane, which acted as

a hinge.

Apparently the spores of M/inchinia ave set free only by the

death of the host.

From the structure of the spore one would expect it to be

destined for a prolonged free existence in sea-water. It has been

proved experimentally that the spores can remain for months

apparently unchanged in water, and can also undergo drying to

some extent. No success, however, has been obtained in attempts

to make them infect other Chiton directly. Living spores have

resisted all attempts to make them open. On some occasions

spores have been introduced into the cesophagus of a Chiton by

means of a very fine pipette; on another occasion a few spores in

a tiny piece of blotting paper tied with a fine silk thread were

introduced into the esophagus of an uninfected Chiton, but on

removal after several hours were found to be quite unchanged.

At other times spores, after being in sea-water, have heen mounted

with teased portions from different regions of the alimentary

canal, all with no result. One such preparation sealed up on

November 27th, and another on December 18th, lost all bacterial

infection after a few days, and were on March 22nd quite sterile

with the spores unchanged.

Uninfected specimens of Chiton were also kept during March

and April under conditions as normal as possible except for the

presence of numerous spores of A/inchinia in the water. Although

the Chiton ate the fucus to which spores easily adhere, they did

not become infected during seven weeks nor were any spores

found in their alimentary canals *.

Attempts were then made to infect other animals from the rocks

at the same zone with the spores of JMinchinia, as it was thought

that possibly shore fish, ete. might become infected in this way

with another vegetative stage of Winchinia. Owing to the courtesy

of the Director, these experiments have been carried on in the

Marine Biological Laboratory, Plymouth, and I am indebted to

Mr. A. J.Smith for much valuable assistance. In each case stones

with several Chiton attached were introduced into large tanks

through which water was circulating and containing the animals

being experimented upon. ‘The results of these experiments were

as follows :—

1. Blenny.—These fish attacked the Chiton readily. All stages

* See note at end of paper.

454 MRS. H. L. M. PIXELL-GOODRICH ON

of Minchinia were digested except the ripe spores, which pass

through unchanged. It is of interest to notice that the shell of the

Chiton passed through undissolved, the eight plates reappearing

generally unbroken in the feeces, whereas in the case of Pomatoceres,

which is also readily torn from the rocks and devoured by the

Blenny, the portions of tube eaten were softened if not entirely

dissolved. This difference is, no doubt, due to the greater pro-

portion of insoluble organic matter in the Chiton shell.

2. Rockling.—Chiton left undisturbed for 7 days.

3. Goby.—One Chiton bitten off, but none eaten during 8 days.

4. Motella.—Chiton left undisturbed for 12 days.

5. Crab (Carcinus menas) devoured Chiton readily. Allstages

of Minchinia were digested except the spores, which passed through

unopened. The tails of the spores were often broken, for, as a

rule, the spores were freed from their cysts.

6. Purpura.—Chiton left undisturbed for over a month.

7. Starfish (Asterias glacialis) —6 Chiton out of 8 eaten in

10 days. Jinehinia spores found in alimentary canal showed no

signs of opening.

8. Sea-Urchin (Hchinus miliaris).—Chiton left undisturbed for

13 days.

The passage of the spores through the alimentary canal of the

Crabs and Blennies lasted at most three days. Doubtless in this

way they are disseminated, but they donot appear to be changed.

Experiments to induce spores, recovered from the feces, to open

in the digestive fluids of Chiton were no more successful than

with fresh spores.

From the description given above it will be clear that Winchinia

has affinities with Urosporidiwm and Haplosporidium and belongs

to the family Haplosporiide of Caullery & Mesnil (2), which

Léger and Duboscq (15a) also recognise as a well-defined gyoup.

There is therefore no need to enter here into the controversy as

to whether the Haplosporidia taken as a whole, as conceived by

Caullery and Mesnil, is a rational group. Doubtless, as these

authors were well aware, some rearrangement wiil be necessaiy

when more is known as to the life-history of these strange forms.

Cépéde (3 & 4) in 191] and 1913 briefly descriked a curious Haplo-

sporidian from Donax, unfortunately without giving any figures ;

owing to the presence of a surrounding cell in the young spore he

seemed to think that the term Haplosporidia (azAovs, simple)

was not suitable,and suggested renaming the group Acnidosporidia.

There appears to be no need for such a procedure, the Donax

parasite and also Minchinia, in which the young spore likewise

shows an indication of a parietal nucleus, have simple spores in

comparison with those which possess polar capsules. At any rate

it seems most inadvisable at present to encumber with new names

and classifications the already complicated literature of these

forms when the majority of their life-histories have never been

at all satisfactorily investigated.

3 |

THE PARASITE MINCHINIA, 4:

Summary.

1. The genus Minchinia is here shown to belong to the Haplo-

sporidia, instead of to the Coccidia, among which it has been

placed since it was established by Labbe in 1896.

2. Its life-history in Chiton consists of two stages, a trophic

and a sporogonic.

3. During the trophic stage a multinucleate individual divides

by a process of plasmotomy.

4. During the sporogonic stage a plasmodium becomes encysted,

forms gametes, which fuse in pairs (? autogamy, see page 451) to

give zygotes. The zygote breaks up into four sporoblasts, each

of which acquires an external membrane drawn out into a tail at

each end, and later a thick chitinous coat immediately inside the

membranous one.

5. Crabs, Blennies, and Star-fish eat Chiton, but the spores of

Minchinia pass through unchanged, and are in this way

disseminated.

The Museums,

Oxford.

P.S.—Since writing the above many Chiton have been kept

here in Oxford with free spores of J/inchinia in aerated sea-water.

After three or four weeks numerous unopened and unchanged

spores, often enclosed in fecal pellets, were found in the intestine

of several of the Chiton. hese, which included uninfected and

previously infected specimens, were carefully examined and some

were cut in serial sections, but in no case did the spores show

any sign of opening. All organs of the specimens not previously

infected were quite normal.

This failure of the spores to open when eaten naturally by

Chiton, confirms the negative results obtaimed in the above

experiments, and we seem forced to the conclusion that the spores

of Minchinia do not open in any part of the digestive tract of

Chiton. :

Of course there is the possibility that free spores may be taken

in by some minute animal in which they germinate before passing

back (perhaps by accidental swallowing) into another Chiton.

However, no evidence is forthcoming on this point at present,

although several Neosporidia have been already described from

small marine animals.

Teferences.

1. Auexererr, A.—‘ Sur le cycle évolutif d’une Haplosporidie

(Ichthyosporidium gasterophilum).” Axch. Zool, Exper.

54: Notes and Revue, pp. 30-44, May 1914.

2. Cauttery, M., & Musnin, F.—‘“‘ Recherches sur les Haplospor-

idies.” Arch. Zool. Expér. 4° sér. iv. pp. 101-181. 1905.

3. Chrrpn, C.— “ Le cycle evolutif et les aftinités systématiques

de l’Haplosporidie des Donav.” C.R. Acad. Se. Paris,

WOK) Way FoR Oe IIL.

456 MRS, H. L. M. PIXELL-GOODRICH ON

4. Cipepe, C.—‘ Les Cytopleurosporés ... embranchement

nouveau du Réegne des Protistes.” CO.R. Acad. Se. Paris,

vol. 156, p. 574. 1913.

5. Cuarron, E.—*‘ Caullerya mesnili, n. g.,n. sp.. Haplosporidie

parasite des Daphnies.” O.R. Soc. Biol. Paris, t. lxu.

js O28), ISO:

6. Cunnor, L.—‘‘ Les organes phagocytaires des Mollusques.”

Arch. Zool. Expér. 54, pp. 267-305. 1914.

7. Doriem, F.— Lehrbuch der Protozoenkunde. Fischer, Jena,

1S)

8. Granara, L.—“ Ricerche sul ciclo evolutivo di Haplospori-

dium limnodrili Granata.” Archiv Protistenkunde, xxxv.

pp. 47-79. 1914.

9. Lasspt, A.—‘ Recherches sur les Coccidies.” Arch. Zool.

Exper. 3° sér. iv. p. 033. 1896.

10. Lassi, A.—Sporozoa, in Tierreich, 5. p. 53. 1899.

11. Lanxesrer, E. Ray.—Protozoa. Encye. Brit. vol. xix.

p- 853, fig. 12. 1885.

12. Lez, <A. Bottes.—‘* The Microtomist’s Vade-mecum.”

Churchill, London, 1913.

13. Lieer, L.—‘‘ Nouvelle Coccidie polysporée du tube digestif

des Myriapodes.” C. R. Soc. Biol. Paris, xlix. p. 1082. 1897.

14. Licer, L.—‘‘ Etudes sur les Coccidies.” Bull. Se. France et

Belg. xxxi. pp. 1-22: 1898.

15. Licer, L.—‘ Sur les Microgameétes des Coccidies.” C.R. Soe.

Biol. Paris, 10° sér. v. pp. 639-641. 1898.

15a. Licer, L., & Dusosce, O.—* Selenococcidium intermedium

et la systématique des Sporozoaires.” Arch. Zool. Exper.

5e sér. v. pp, 187-239. 1910.

16. PrixeLu-Goopricu, H. L. M.— The Sporozoa of Spatangoids.”

Quart. Journ. Micros. Se. vol. lxi. 1915.

17. Scuaupinn, F.—‘‘ Untersuchungen iiber den Generations-

wechsel bei Coeccidien.” Zool. Jahrb. Anat. vol. xi.

pp- 197-287. 1900.

18. Swarczewsky, B.—‘* Ueber den Lebenscyclus einiger Haplo-

sporidien.” Archiv Protistenkunde, xxxii. pp. 49-108.

1OWA:

EXPLANATION OF THE PLATES.

Unless otherwise stated, the figures were drawn with a camera lucida from pre-

parations stained with iron hematoxylin.

Lettering.

e., cyst; ch., chitinous coat of spore; ad, daughter plasmodium; g., gamete;

gr., chromatoid granule; /., host-cell; ., nucleus; 0., operculum; s., sporo-

blast; sp., spherule; €., tail of spore.

Puate I.

Fig. 1. Section of a small Plasmodium of Minchinia with pseudopodia and several

nuclei, some dividing. XX 1000.

2. Another specimen showing Plasmotomy : d., small daughter plasmodium

partly covered by dividing parent form; #., host-cells (phagocytes).

< 2000.

Fig.

Ole

THE PARASITE MINCHINIA. 457

. Portion of a section of an encysted parasite beginning to break up into

gametes: c., cyst with host-cells flattened against it; g., amoeboid gamete.

Stained Ehrlich’s hematoxylin. > 2000.

. Portion of a cyst contaiing free and conjugating gametes. 3000.

. (a) Prozygote formed by fusion of the cytoplasm of two gametes. Stained

Ehrlich’s hematoxylin. (+) Zygote showing formation of syncaryon.

. Zygote showing first division of the syncaryon. 3000.

. Later stage: second division nearly complete. > 3000.

Prarn II.

. Portion of a cyst showing two stages of division into the four sporoblasts :

s., free sporoblasts. > 3000.

. Stages in early spore-formation from sections. X 3000. (a) Showing be-

ginning of nuclear activity; (6) showing single parietal nucleus ;

tail cut through.

. Young living spore from a cyst: the clearer space indicating position of

nucleus. X 2000.

. Slightly older spore also drawn from the living. > 2000.

. Young spore with large nucleus giving off chromatic bodies into cytoplasm.

x 3000.

. Optical section of a spore nearly full grown and with chitinous coat (ch.)

almost complete. Membranous tails (¢.) cut through spherule (sp.).

x 3000.

. Optical section of a free spore at stage between those represented by figs. 12

and 13, showing chromatic granules arranging themselves just inside

membrane to form chitinous coat. Slghtly flattened by cover-glass.

x 2000.

. Adult spore drawn from the living. 1000.

. Spore opened by means of pressure and reagents. X 1000.

a . 7 nave

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Stage 1. Perameles obesula (12 b).

PZ.S. 1915: PARKER” Pia

Stage!. Dasyurus viverrinus (7.5mm.).

Wiles Qe

3 M.M.P. Al.

F.G.

A. P—

Stage II.

care Nidan

SR OR

Dasyurus viverrinus (8.5mm. A.).

MENPES PRESS, WATFORD.

HEART OF MARSUPIAL EMBRYOS.

Ee ee ae mera’ Nén«

ISS, JPNRKOR, JI,

IP, aS

“MOL/A [B1]Ud/\,

: ‘OPO JO MOA [es10G

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MODELS OF EMBRYONIC MARSUPIAL HEART. |

E. A. STEELE, DEL.

ON THE DEVELOPMENT OF THE HEART IN MARSUPIALS. 459

39. The Harly Development of the Heart and Anterior

Vessels in Marsupials, with Special Reference to

Perameles. By Karuarine M. Parker, B.Sc. (Lond.),

Assistant in the Department of Zoology, University

of London, University College *.

[Received May 25, 1915; Read June 8, 1915. ]

(Plates I., II.f and Text-figures 1-25.)

INDEX. Page

Imtyoditctionh iyo rcteae eee cece, 469

Description of SHaees eSciawsnacotaenennmeey AGO.

Summary and Diao See Bec keascwous pAOO

(a) Development of the Soa} « vee 486

(6) Development of the Cardinal Vigne eee) CAO:

Final Summary and Conclusions .. A cadiauaaneanercahry (OT

References to Titcarare een eee) Oy i aK 497

INTRODUCTION.

The following piece of work was undertaken at the suggestion

of Professor J. P. Hill, for whose invaluable help in carrying it

out I am deeply grateful. The embryos studied are all from his

collection and text-figs. 1 and 6 were made from photographs

taken by him.

I wish also to thank Miss EH. A. Steele for her beautiful

figures of the model (Stage V.) and Mr. F. C. Pittock for much

help in making the model.

The material affords an excellent opportunity for the study of

the mode of development of the pleuro-pericardial canals, the

origin and differentiation of the endothelial heart-tubes, and the

method of fusion of the lateral primordia of the heart, and it

was in the hope of making some progress towards the solution of

the interesting problems of early cardiac development that the

work was undertaken.

The general arrangement of the pericardium, heart, and aorta

in the early stages (viz. Stages I. to IV. inclusive) can be

accurately determined by graphic reconstruction. Figures 1 to 5

(Pl. I.) were all obtained by this method, and are intended to

give some idea of the relations of the pericardium and heart to

the gut and the brain-plate in successive stages.

In the next stage (Stage V.) the curvature of the heart makes

it impossible to represent the relations of the parts accurately in

two dimensions. A wax-plate reconstruction was therefore made

(see Pl. IT.), and as all the vessels of the head which were

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

+ For explanation of the Plates see p. 499.

Proc. Zoou. Soc.—l915, No. XXXII. 32

460 MISS K. M. PARKER ON THE

recognisable as such were included in the model, a certain amount

of light was shed on the development and early relations of the

cardinal veins and aortic arches. The following account there-

fore deals not only with the development of the heart, but also

with certain facts relative to the early development of the

vessels of the head.

DESCRIPTION OF STAGES.

Srace I. Perameles obesula (1 Z).

Dasyurus viverrinus (7°5 mim, vesicle).

The material for this stage consists of four embryos of Pera-

meles obesula, one cut longitudinally and three transversely, and

several of D. viverrunvus. —

(a) Perameles obesula (1 Z).

* Total length of embryo A in curved condition =6-08 mm.

In this stage there is a flat brain-plate with an extensive neural

crest proliferation in the cranial region (text-fig. 1, V.C.P.(Z.G.)).

Text-figure 1.

Perameles obesula (1Z). Dorsal view of embryo A.

Av. Avea vasculosa. 2... Lateral mesoderm. N.C.P. (2.G.). Neural crest

proliferation (primordium of trigeminal ganglion). S. Somite. A, A). Level

of section represented in text-fig. 2.

There are four distinct somites (S.): behind the fourth is a

somitic thickening with an indistinct posterior limit, and in front

of the first, another imperfectly defined mass, representing

presumably a transitory first somite.

* All measurements of embryos and certain descriptive details are taken from

Professor Hill’s original notes and photographs of the material.

DEVELOPMENT OF THE HEART IN MARSUPIALS. 461

Laterally to the somitic mesoderm and the medullary plate is

a zone appearing clear in surface view, in which the mesoderm

forms a thin sheet. Outside this again is an opaque zone of

lateral mesoderm (Z.J/.), completely surrounding the embryo.

This is bounded peripherally by a clear zone which separates it

from the area vasculosa (Av.). The entoderm forms a thin

continuous layer, while a small incipient head-fold definitely

marks the anterior margin of the brain-plate and involves also

the protochordal plate lying in the middle line immediately

below it.

In the lateral mesoderm on each side a horizontal cleft has

appeared, separating the mesoderm into a dorsal somatic and a

ventral splanchnic layer. These clefts, commencing on both

sides of the embryo and extending forwards, constitute the

leuro-pericardial canals, the form of which can readily be seen

in Pl. I. fig. 1 (P.p.C.) The canals form a horseshoe the median

Text-figure 2.

Perameles obesula (1Z, B). Transverse section in plane A, Aj.

(See text-fig. 1.)

Ent. Entoderm. J.P. Medullary plate. P.p.C. Pleuro-pericardial canal,

S.M, Splanchnic mesoderm. So.M. Somatic mesoderm.

anterior portion of which lies underneath the anterior margin of

the brain-plate (JZ.1/.P.), while the lateral limbs extend back

into the somitic region. In the anterior region, the dorsi-ventral

extent of the pleuro-pericardial canal is very small (text-fig. 2,

P.p.C.), and in fact the continuity of the canal is actually inter-

rupted on the right side of the embryo (PI. I. fig. 1).

The cleft increases markedly in size in the region of the

trigeminal neural crest proliferation (text-fig. 1, V.C.P. (7.G.))

and attains its greatest size in the hind-brain region (text-fig. 3 A

& B). Behind this it becomes reduced in size until, opposite the

posterior somites, the ccelomic cavity is represented by irregular

clefts in the lateral mesoderm.

The pleuro-pericardial canals throughout their extent have a

thin somatic and a slightly thicker splanchnic wall (see text-figs. 2,

3A & B, Sod, S.M.), which in the anterior region is in close

32*

462 MISS K. M. PARKER ON THE

contact with the entoderm (text-fig. 2). In the region of the

maximum size and development of the pleuro-pericardial canals

(Pl. I. fig. 1) the endothelial primordia of the heart are

differentiating (text-fig. 3, Hnd.) between the entoderm and the

splanchnic mesoderm, which therefore projects as a prominent

fold into the pleuro-pericardial canal.

Text-figure 3.

Perameles obesula (1 Z, B).

A. Transverse section m region of greatest width of pleuro-pericardial canals,

with endothelial tubes developed. B. Transverse section showing origin of

angioblast cells from the splanchnic mesoderm.

A.C. Angioblast cell. Hnd. Endothelium. M.P. Medullary plate. P.p.C. Pleuro-

pericardial canal. S.J£. Splanchnic mesoderm. So.1Z. Somatic mesoderm.

The endothelial heart primordia are best developed in their

posterior portions, where they are actually tubular in some

embryos of this stage (see text-fig. 3 A). Anterior to the tubular

portion, the primordia are represented by solid cords of angio-

blast cells, isolated examples of which are found scattered along

the length of the pleuro-pericardial canals in the positions indi-

eated in Pl. I. fig. 1(4.C.). It may be concluded from this that

the endothelial tubes differentiate postero-anteriorly, and that

their increase in length is brought about not by direct forward

growth of the first formed parts of the tubes, but by the

DEVELOPMENT OF THE HEART IN MARSUPIALS. 463

progressive differentiation of angioblast cells in the cephalic

portions of the pleuro-pericardial canals.

The evidence of this stage does not justify any definite state-

ment with regard to the origin of the endothelium of the heart.

From text-fig. 3 B it will be seen that the splanchnic mesoderm

(S.J) shows distinct traces of proliferative activity ; its ventral,

indented margin has an irregular outline and there are indications

of loosening of the cells. On the other hand, there is no definite

evidence of entodermal proliferation, though in an earlier stage,

which will form the subject of a separate paper, the appearances

by no means exclude the possibility of the entodermal origin of

the endothelium, whilst there is clear evidence of proliferative

activity on the part of the entoderm of the area vasculosa.

(b) Dasyurus viverrinus (7°5 mm. vesicle).

The material on which the following description is based

consists of two embryos, one cut transversely, the other longi-

tudinally.

Greatest length of each embryo=7 mm.

In this stage there is a flat brain-plate, the anterior margin of

which is marked in the middle line by a thickened terminal ridge

(text-fig. 4, 7.R.). There is a well developed neural crest proli-

feration, the anterior portion representing the primordium of

the trigeminal ganglion, and the posterior that of the facial,

glosso-pharyngeal and vagus ganglia. No somites are yet

differentiated.

The outline of the anterior end of the brain-plate and the

pleuro-pericardial canals and endothelial heart-tubes are shown

in Pl. I. fig. 2. From this it will be seen that the pleuro-

pericardial canals (P.p.C.) extend continuously round the head-

end of the embryo and lie anterior to the anterior margin of the

brain-plate (J7.1/.P.).

The. pleuro-pericardial canals attain their greatest size in the

hind-brain region (#.B.)at this stage, and here also the primordia

of the heart are well established in the form of endothelial tubes

lying between the entoderm and the thickened splanchnic

mesoderm. The endothelial tubes terminate anteriorly at the

level of the posterior limit of the trigeminal primordium. In

front of this, however, there are scattered angioblast cells and

strands of cells extending forwards as indicated in Pl. I. fig. 2.

In the condition of the heart primordia the Dasyurus embryos

are in advance of those of Perameles described above, for the

endothelium is definitely tubular throughout a great portion of

its extent and the myocardial fold is consequently well developed.

From the longitudinal section (text-fig. 4) it will] be seen that

the antero-median portion of the pleuro-pericardial canal lies

some distance in front of the anterior margin of the brain-

plate (7.2.).

In Stage I., then, we have a horseshoe-shaped pleuro-pericardial

464 MISS K. M. PARKER ON THE

cavity extending round the embryo and with the lateral limbs

prolonged into the somitic region. The endothelial primordia of

the heart are represented by more or less continuous tubes, solid

cords, and scattered angioblast cells differentiating in the postero-

anterior direction and lying between the entoderm and the

splanchnic mesoderm.

The endothelial heart-tubes are the only vessels yet established.

Text-figure 4.

Dasyurus viverrinus (75 mm. vesicle). Longitudinal section, median

through the anterior margin of the brain-plate.

Amn. Head-fold of amnion. Ent. Entoderm. P.p.C. Pleuro-pericardial canal.

T.R. Terminal ridge.

Srace II. Dasywrus viverrinus (8'°5 mm.).

The material for this stage consists of one embryo (A) of

Dasyurus viverrinus cut transversely and one (Aa) cut longi-

tudinally. A graphic reconstruction of the gut and pericardium

together with the endothelial heart-tube and aorta of the left.

side of embryo A will be found in Pl, I. fig. 3.

DEVELOPMENT OF THE HEART IN MARSUPIALS. 465

Total length of embryo A, 8°5 mm.

This embryo possesses a flat brain-plate with well marked

optic grooves; somites are not as yet distinctly differentiated.

The position of the anterior margin of the brain-plate is indi-

cated in fig. 3 (J2.1.P.) as well as the outline of the gut (7.G.).

On comparison with fig. 2, it is clear that the progress of the

head-fold has brought about considerable alteration in the

Text-figure 5.

Dasywrus viverrinus (8°5 mm.). Longitudinal section of embryo Aa,

median through anterior end.

C.F. Cardiac fold, #.G. Fore-gut. M.M.P. Margin of medullary plate.

P.p.C. Pleuro-pericardial canal.

relations of the gut, brain, and pericardium (see Pl. I. fig. 3 and

text-fig. 5). In embryo A (fig. 3), which is slightly in advance

of embryo Aa (text-fig. 5), the anterior margin of the brain-plate

now marks the actual anterior limit of the embryo, while at the

same time the crescent-shaped fore-gut (/.G'.) hasdeveloped. Its

anterior limit lies immediately behind the anterior margin of the

brain-plate, while closely applied to its posterior and lateral

466 MISS K. M. PARKER ON THE

margins lies the continuous pericardial cavity which shows a

marked increase in size as compared with the previous stage.

(Compare PI. I. figs. 2&3, P.p.C.) This increase is greater in the

median anterior limb of the pericardium than in its lateral

portions. (Compare text-figs. 4 & 5.)

The differentiation of the endothelial primordia of the heart

has progressed considerably and they now extend as actual tubes

even to the middle line of the pericardium. This fact is some-

what remarkable, as such a condition, in which the lateral heart-

tubes are actually in contact at their extreme cephalic apices

and diverge widely and abruptly from this point of contact, is

not found in any other stage.

Some distance behind its cephalic extremity, the lateral heart-

tube gives rise to the first aortic arch (fig. 3, 4 1), which follows

the antero-lateral margin of the gut almost to the middle line

and there becomes continuous with the corresponding dorsal

aorta, the two aorte being completely established in this stage.

It will be unnecessary to go into further details of the structure

of this embryo, as the sectional appearances found in it are

exemplified yet more clearly in the Perameles embryo to be

described next. It is, however, an important stage with regard

to the processes of growth and folding which bring about the

relations of the brain, gut, and pericardium which are found in

subsequent stages,

Stace IIT, Perameles nasuta (18).

The material in this stage consists of two flat embryos with

widely open brain-plates and lateral heart-tubes. Embryo A has

eleven somites, the first being small and indistinctly limited;

embryo B has nine, the most anterior being indistinct here also.

The following description is based mainly on embryo A, which

was cut transversely,

Total length of embryo A after partial flattening under cover-

glass, from the anterior margin of the brain-plate to the hinder

extremity of the primitive streak : 7-5 mm.

Vascular area, 8°96 x 5°8 mm. in diameter.

Text-fig. 6 represents a dorsal view of embryo A, while the

outlines of the gut and pericardium and the endothelial heart-

tube and aorta of the left side are given on PI. I. fig. 4. From

these figures it will be seen that the stage shows a considerable

advance in general development on the preceding. The head-fold

has progressed back as far as the region of the auditory pit.

The brain, though flat and widely open in the fore- and mid-

brain regions, is deeply grooved in the hind-brain region. Lying

lateral to the brain-plate are two pairs of mesodermal masses,

one the maxillo-mandibular process, the other the hyoid arch.

Between these two arches is the first visceral pouch, and behind

the hyoid arch the second visceral pouch is already developed.

DEVELOPMENT OF THE HEART IN MARSUPIALS. 467

Shortly behind this, the paraxial mesoderm is differentiated

into somites.

The maxillo-mandibular process (text-fig. 6, W/m. arch.) forms a

dense mass of mesoderm not distinctly marked out into maxillary

and mandibular portions. It lies laterally to the gut on the

outer side of the Jateral and dorso- and ventro-lateral walls of the

gut, and extends antero-posteriorly from almost the anterior end

of the gut back to the first visceral pouch, which is situated level

with the anterior intestinal portal. The entoderm of the first

visceral pouch reaches the ectoderm, but the closing membrane

is not perforated.

Text-figure 6.

Perameles nasuta. Dorsal view of embryo A.

B, B, indicates the level of the section represented in text-fig. 7.

Aw. Auditory vesicle. F.N. Facial neuromere. H.arch. Hyoid arch. Mm. arch.

Maxillo-mandibular arch. 27.P. Medullary plate. S. Somite.

The hyoid arch is a mass of mesoderm only slightly smaller in

surface view than the maxillo-mandibular arch, but situated

entirely dorsal to the gut. Behind it is the second visceral

pouch, which is small and does not reach the ectoderm

Gels ties 4 Ve 2).

The form of the pericardium and its relations to the gut,

468 MISS K. M. PARKER ON THE

as well as the topography of the endothelial heart-tubes and the

aorte and aortic arches, will be understood best by reference

to fig. 4 and text-fig. 12. From fig. 4 it will be seen that the

pericardium (P.) has increased markedly in size. As in the

previous stage, its inner wall is closely applied to the entoderm

of the crescentic or U-shaped anterior intestinal portion (4./.2.)

(cf. text-fig. 12). The antero-median portion of the pericardium

is somewhat rectangular in transverse section (text-fig. 7); its.

dorsal wall is slightly thicker than its ventral, and between

the former and the floor of the gut lie the endothelial heart-

tubes (text-fig. 8) and the first aortic arches (text-fig. 7),

Text-figure 7.

eae,“

Perameles nasuta (18, A.). ‘Transverse section through median

pericardium and first aortic arch.

A. First aortic arch. D.A. Dorsal aorta. F.G. Fore-gut. 2.P. Medullary

plate. 2f.m.A. Maxillo-mandibular arch. N. Notochord. PP. Pericardium..

V.C.M, Vena capitis medialis.

In the antero-median limb of the pericardium, the endo-

thelial heart-tubes, enclosed in a fold of splanchnic mesoderm,

lie separate from each other below the closed gut (Pl. I. fig. 4

and text-fig. 8), while at the level of the anterior intestinal

portal they diverge in the manner indicated in fig. 4 and lie on

either side of the open gut in the dorso-medial wall of the peri-

cardium (text-fig. 9). In their posterior portions the heart

primordia lie on the ventral side of the pericardium.

The endothelial heart-tubes throughout their length are almost:

completely enclosed by the layer of splanchnic mesoderm con-

stituting the primordium of the myocardium. In the posterior

region of the heart the myocardial layer is closely applied to the

outside of the endothelial tube (text-fig. 10), but in the greater

part of its extent there is a considerable space between the two.

layers of the heart primordium (text-fig. 9).

From the cephalic extremity of each endothelial tube, there

arise two vessels, one of which runs forwards and outwards

DEVELOPMENT OF THE HEART IN. MARSUPIALS. 469

towards the lateral margin of the gut and then parallel with‘this

margin (Pl. I. fig. 4,41). It loops round the anterior limit of

the gut, joins the dorsal aorta, and thus constitutes the first or

mandibular aortic arch. It is impossible here to fix any exact

limit between the endothelial heart-tube and the aortic arch.

Text-figs. 7 and 8 represent typical sections through each. The

transition from the heart primordium to the aortic arch is

indicated by the gradual reduction in the dorsi-ventral extent of

the space surrounding the endothelial tube and the rotation

of the vessel, so that its greatest diameter comes to lie parallel

Text-figure 8.

Perameles nasuta (18, A.).

A. Transverse section through heart primordia and second aortic arch.

B. Complete outline of same section.

Ay. Second aortic arch. D.A. Dorsal aorta. Hd. Endothelium of heart.

M.m.A, Maxillo-mandibular arch. 2£.P. Medullary plate. My. Myocardium.

P. Pericardium. V.C.M. Vena capitis medialis.

with the floor of the gut. (Compare text-figs. 8 and 7.) Outside

the limits of the pericardium the aortic arch runs in the mesoderm

of the maxillo-mandibular process (text-fig. 7). It is connected

with the dorsal aorta by one main loop and by several smaller

vessels which were omitted from fig. 4 for the sake of clearness.

The second vessel, which arises from the anterior end of the

heart, is small and runs backwards and outwards, lateral to and

almost parallel with the heart-tube, and has precisely the same

relations to the gut and mesoderm.

470 MISS K. M. PARKER ON THE

Comparison with the succeeding stages shows that this repre-

sents the ventral portion of the future second aortic arch (Pls. L.,

II., figs. 4 & 8, A 2), while a small vessel arising from the dorsal

aorta and running outwards on the dorsal wall of the gut ventral

to the auditory vesicle, corresponds with the dorsal portion of the

completed arch in later stages.

Text-figure 9.

een, és

ae.’ : f

ig © oe. Eo

é Re 7» OS

Gag Otters

bas ‘eh 226 oo wg te, L

See

Fs. g

Se iS

hae ee Ae TORY

BE OE

Perameles nasuta (18, A.).

A. Transverse section in region of lateral heart primordia and open gut.

B. Complete outline of same section.

Ag. Second aortic arch. D.A. Dorsal aorta. Hind. Endothelium of heart.

M.F. Medullary fold. My. Myocardial layer. P. Pericardium. V.C.M. Vena

capitis medialis, V.P.1. First visceral pouch.

In the median space between the anterior ends of the endo-

thelial heart-tubes are a number of scattered angioblast cells

lying between the splanchnic mesoderm and the entoderm

(Pl. I. fig. 4 and text-fig. 7). These cells possibly represent the

primordia of the capillaries found in the corresponding position

in the next stage. They afford an instance of the origin of

angioblast cells from the splanchnic mesoderm after the establish-

ment of the definitive endothelial heart-tubes.

_ Posterior to the region represented in fig. 4, the heart-tubes

DEVELOPMENT OF THE HEART IN MARSUPIALS. 471

gradually curve outwards and pass imperceptibly into vitelline

veins.

The dorsal aorta is paired and runs back continuously, the two

vessels keeping approximately the same distance from the middle

line (Pl. I. fig. 4, D.A.).

Immediately dorsal to the dorsal aorta on each side there is

situated a series of apparently isolated sections of a minute blood-

vessel (text-fig. 7, V.C.W/.). These capillaries lie close against

the medullary tube, medially to the neural crest proliferation in

the region of the trigeminal ganglionic primordium. From the

position of this vessel relative to the dorsal aorta and nerves,

it is evidently the vena capitis medialis of Grosser (5). A brief

summary of some of the literature on the subject of the vene

capitis medialis and lateralis and their relations to the anterior

cardinals will be found below, together with a review of the facts

of development of these veins in Perameles.

Text-figure 10.

M Fig

‘s ot Seat

ae Seo ane

Perameles nasuta (18, A.). Transverse section through posterior

portion of lateral heart primordia.

D.A. Dorsal aorta. End. Endothelium of heart. I/.F. Medullary fold.

My. Myocardial layer.

Although it is not within the scope of this paper to deal

in any detail with the posterior vessels, it may be noted here that

in this stage, in the region of the posterior somites, there occur

intersegmental offshoots from the dorsal aorte. These inter-

segmental offshoots are shown very clearly in the longitudinal

series (text-fig. 11). It will beseen from the figure that between

each two successive somites (S.) there is a small dorsal offshoot

from the aorta. Dorsal to the somites there are a few scattered

endothelial cells (Hnd.). The longitudinal vessel connecting the

offshoots is not continuous, but portions of. it are present in the

next section.

At the level of the second somite there occurs a pair of small

vessels lying in the somatopleure immediately dorsal to the heart-

tubes. Each consists of a single vessel with a few minute

A72 MISS K. M. PARKER ON THE

branches and is blind at both ends and not connected as yet with

any other capillaries. From these vessels the Cuvierian ducts

are later developed.

In this stage we have, therefore, lateral heart-tubes which,

while they approach one another anteriorly, are widely separate

in the posterior region. Paired dorsal aortz and the first pair of

aortic arches are developed, whilst traces of the second arch are

also present. This stage accordingly agrees inits general features

with the 83 days rabbit described by Bremer (1). In addition

to the heart and arterial vessels there are also present in the head

the first traces of the venous system in the form of disconnected

portions of the vena capitis medialis and the primordia of the

Cuvierian ducts.

Text-figure 11.

Perameles nasuta (18, B.). Longitudinal section through somites showing

dorsal offshoots from the dorsal aorta,

D.A. Dorsal aorta. D.Br.D.A. Dorsal branch of samo. End. Endothelium

of longitudinal vessel. SS. Somite,

Stack IV. Perameles nasuta (2 P).

The material consists of four embryos, A and C cut transversely,

B and D longitudinally.

Each of the four embryos has fifteen or sixteen somites, The

neural tube is still unclosed throughout its length, but the folds

are closely approximated in the hind-brain region. The mid- and

fore-brain segments are widely open as in Stage III., but the

primary cranial flexure has occurred so that the fore-brain is

bent forwards and downwards (see text-fig. 15),

The relations of the gut, pericardium, and heart are indicated

in Pl. I. fig. 5. It is well to note at this point that the

outline of the brain-plate as indicated in figure 5 is not

strictly comparable with the corresponding line in fig. 4

(compare text-figs. 12 and 15). The fact that the primary

cranial flexure has occurred, renders it impossible to plot the

morphological anterior end of the brain in the same plane recon-

struction with the hind-brain, gut, ete. The difference in the

relations of the brain to the pericardium in the two stages may,

however, be judged by the position of the auditory neuromere

(4..V.), which lies at the posterior margin of the pericardium in

DEVELOPMENT OF THE HEART IN MARSUPIALS. 473

Stage III. and at the anterior margin thereof in the present

stage.

The first and second visceral pouches are now well marked

(fig. 5, VP. 1 & 2), while the relations of the maxillo-mandibular

process and hyoid arch show little advance on the preceding

stage.

The antero-median portion of the pericardium has increased

very considerably in the antero-posterior direction. Furthermore,

it may be noted that as the portion of the gut lying anterior to

the first visceral pouch has remained the same length (compare

figs. 4 & 5) and the anterior margin of the pericardium is now

situated in the same plane with the first visceral pouch, the

pericardium must have moved backwards as a whole.

Text-figure 12,

PP “hie

Sac mS fates x

ee Bee... eel | ww

EEE me a

sree r

(LAE Rc

A 88 GP ape 59550 Ong.

ets cunyyssee VIS

Perameles nasuta (18, B). Longitudinal section, median through

the anterior end.

A.I.P, Anterior intestinal portal. C.F, Cardiac fold. 2.1M.P. Margin

of medullary plate. P. Pericardium. P,P. Protochordal plate.

The form of the endothelial primordia of the heart is shown

in fig. 5. They have fused at their cephalic extremity, the fused

portion extending through some eighteen sections and represent-

ing the most closely approximating portions of the endothelial

tubes in Stage ITI. (Pl. I. fig. 4). From it is derived the bulbus

' (conus) arteriosus of the next stage (PI. II. fig. 8, B.A.). Posterior

to this fused portion, the endothelial tubes lie close together but

unfused for a considerable portion of their length (Pl. I. fig. 5

and text-fig. 14), and then diverge widely and pass into vitelline

veins, The endothelial tubes throughout their length are

enclosed by the myocardial wall, which shows characteristic

thickening and prolongations of its cells throughout the greater

part of the length of the heart (text-fig. 14, My.).

The myocardium of the right and left sides is united from the

cephalic extremity of the heart primordium to the point of

divergence of the right and left endothelial tubes, but the line

of fusion is not marked by any groove. In the posterior region

where the endothelial tubes separate from each other, each is

surrounded by its own myocardial layer, so that for a short

distance in front of the anterior intestinal portal, the two heart-

tubes lie below the closed gut, each surrounded by an independent

fold of splanchnic mesoderm. The primordia of the heart

474 MISS K. M. PARKER ON THE

are prolonged into the lateral gut-folds and pass gradually into

vitelline veins.

The heart as a whole is somewhat asymmetrical (see fig. 5),

being curved over to the right side of the embryo.

The aortic arches, two of which are developed, arise from the

median bulbus arteriosus. The endothelial heart-tube bifurcates

in front, and each half runs forwards and slightly outwards as a

relatively wide vessel situated between the two layers of the

splanchnopleure. These vessels, which constitute the first aortic

arches, run forwards and outwards in a course similar to that

of the same vessels in the next stage (compare figs. 5 and 6).

Anteriorly they loop round the gut to join the dorsal aorte.

From the anterior convexity of this first aortic arch are given off

capillaries which form a network surrounding the primary optic

vesicles. From the lateral margin of the fused tip of the heart

is given off on each side a small vessel which runs outwards and

backwards, loops round the gut, and constitutes a continuous

though slender second aortic arch.

Text-figure 13.

Si8g)

é ee

Peas pet As

Perameles nasuta (2 P, A). ‘Transverse section through bulbus arteriosus.

End. Endothelium. G. Gut. My. Myocardium. P.C. Pericardial cavity.

Rt.A. Root of aortic arch.

The vena capitis medialis, which was just recognisable in the

preceding stage, is now considerably further developed. It is

represented by an irregular and not perfectly continuous series

of capillaries, situated dorsal to the aorta on either side of the

medullary tube, medial to the cranial ganglia. These capillaries

are connected by very fine sprouts with the dorsal aorte (text-

fig. 14, V.C.ML.); ventro-lateral to the auditory vesicle and lateral

to the nerve-roots, there is another line of scattered capillaries

connected with the vena capitis medialis. These are the first

traces of the vena capitis lateralis. Portions of the vena capitis

medialis can be traced in the region of the lateral heart-tubes as

far back as the Cuvierian ducts (7.e¢., the region of the third

DEVELOPMENT, OF THE HEART IN MARSUPIALS, 475»

somite). Behind this again there are, as in Stage III, inter-

segmental offs hoots from the dorsal aorta, but as yet no continuous:

vessel in this region.

Text-figure 14,

My 68

ae ~Bo ba

ay Ey Ceara

WSs repeat 4SS ee:

SENN eal

you Z

Transverse section through ventricular

Perameles nasuta (2 P, A).

region of the heart. Ss

Ag. Second aortic arch. A.V. Auditory vesicle. D.A. Dorsalaorta. nd. Endo-

V.C.M. Vena capitis:

thelium. J.P. Medullary plate. My. Myocardium.

medialis.

Text-figure 15.

Re, ; Zwei

£0... 5, ws! ‘ts ie? 7

i is

e Saaleies! 2 sa"

Perameles nasuta (2 P, B).

Longitudinal section, median through the anterior end of the embryo.

End. Endothelium. #.B. Fore-brain. F.G. Fore-gut. UB. Mid-brain.

U.M.P. Margin of medullary plate. My. Myocardium, O.P. Oral >plate..

P. Pericardium.

In this stage the Cuvierian ducts are recognisable lying in

the somatopleure opposite the third somite and immediately

Proc. Zoo. Soc.—1915, No. XX XITI, 33

476 MISS K. M. PARKER ON THE

dorsal to the heart-tube. The main trunk of each ends blindly

anteriorly and posteriorly and is of considerable size, causing a

bulging of the mesoderm of the somatopleure, which is thus

brought into contact with the mesoderm surrounding the heart-

tube. As yet, however, neither of the Cuvierian ducts opens

into the heart-tube. From the medial side of each Cuvierian

duct a few small capillaries are given off. They run towards the

middle line and represent that portion of the anterior cardinal

vein which at a later stage connects the vene capitis medialis and

lateralis with the Cuvierian duct. (Compare Pl. II. fig. 7,

Ae VW)

Stage IV., therefore, possesses a heart in which the endothelial

tubes have fused anteriorly and curvature has already com-

menced. ‘Two complete aortic arches, an incomplete vena capitis

medialis, traces of a vena capitis lateralis, and Cuvierian ducts

are present.

Stace V. Perameles obesula (10. viii. 03).

Macropus sp.

The material for this stage consists of five similar embryos of

P. obesula, three cut transversely and two longitudinally, and one

embryo of Onychogale frenata (? Macropus sp.) cut transversely.

In several respects, e. gy. curvature of the heart, the Onychogale

embryo represents a slightly earlier stage than Perameles obesula

(10. viii. 03). For purposes of description, however, it will be

convenient to deal first with the Perameles embryos, as a wax-

plate reconstruction was made of the heart and anterior vessels

of embryo A (PI. II. figs. 6-8).

As regards general development, Perameles obesula (10. viii. 03)

shows only a slight advance on P. nasuta (18) Stage III. The

brain has practically not changed ; the gut is in the same condition

except that the first visceral pouch is more extensive and closure

of the fore-gut has progressed back to slightly behind the auditory

vesicle.

In the vascular system, however, we find a most marked

advance, the heart having assumed a definite form with ventricular

and auricular divisions recognisable. Figs. 6 to 8 illustrate the

model of the heart and anterior vessels in this stage. The gut,

included to form a building base, is coloured white, heart endo-

thelium and arteries red, veins and most of the capillaries blue,

myocardium yellow. The myocardium is left intact on the left

half of the model from the roots of the aortic arches back to the

level of the anterior end of the Cuvierian duct, but has been

omitted on the right side so that the whole of the endothelial

tube is here exposed to view. The capillaries surrounding the

gut are also left intact on the left side of the model, but on the

right have been omitted in order that the aortic arches might be

seen more clearly. Study of the actual sections shows that the

capillaries of the right side closely resemble those of the left.

DEVELOPMENT OF THE HEART IN MARSUPIALS. AT7

The first and second visceral pouches are seen in the model as

lateral projections from the gut (figs. 6-8, V.P.1& 2). The

heart, which is median and ventral anteriorly, still consists of

separate lateral primordia posteriorly, the two halves diverging

in the lateral lips of the anterior intestinal portal (fig. 6, A./.P.).

By the great enlargement of the heart itself, the pericardial cceelom

has become relatively considerably reduced, and now simply forms

a space surrounding the heart ventrally and ventro-laterally

(text-fig. 18). The median pericardium extends from the

cephalic extremity of the heart to the anterior intestinal portal.

At their cranial ends the endothelial tubes (exposed in the

model by the omission of the myocardium) unite to form a broad

conical portion, the bulbus (conus) arteriosus. The first and

second aortic arches (A. 1 & 2) are given off from the dorsal side

of the bulbus (fig. 8, B.A., A. 1 &e.). In this region, the myocar-

dium simply forms a continuous layer covering the endothelium,

Text-figure 16.

oe a. |

Sawer

x is

Peery

Perameles obesula (10.viii.03).

A. Transverse section through bulbus arteriosus.

B. Complete outline of same section.

End. Endothelium of bulbus arteriosus. G. Gut. My. Myocardium.

but separated from it by aspace (text-fig. 16). The bulbus is the

only portion of the heart in which the endothelial tubes have

actually fused. Behind it, the tubes are in contact as far back as

the anterior intestinal portal, but the wall between them is every-

where complete. The myocardium of the two sides has fused

throughout the region of the closed gut and there is no ventral

mesentery (text-figs. 16-18). There is, however, a very well

marked groove on the ventral aspect of the myocardium, which

marks the line of junction of the right and left halves. At the

opening of the gut the right and left heart primordia separate

completely.

From the ventral view of the model (PI. II. fig. 6) it is obvious

that already the heart has begun to bend between the two points

(a) the roots of the aortic arches and (6) the opening of the fore-gut

33*

478 MISS K. M. PARKER ON THE

The curvature, however, does not affect both sides equally, and a

marked asymmetry results. The anterior ventricular portions of

the heart-tubes are already being pushed backwards so as to lie

ventral to the auricular portions. It is clear that by continuation

of this curvature with accompanying fusion of the two halves,

the typical embryonic relations of auricle and ventricle will

ultimately be achieved.

In the ventricular region of the heart, the right and left endo —

thelial tubes are approximately equal in size, but where there is

an inequality the right is the larger (text-fig. 17, Hnd.)

Text-figure 17.

Perameles obesula (10.viii.03). Transverse section through the ventricular region

of the heart.

Ag, Second aortic arch. A.V. Auditory vesicle. D.A. Dorsal aorta. Hnd. Endo-

thelium. J.P. Medullary plate. My. Myocardium. V.C.L. Vena capitis

lateralis. V.C.M. Vena capitis medialis.

In addition to the curvature which is bringing the ventricular

region into position ventral to the auricular region, there is a

certain amount of curvature in the horizontal plane of the

embryo. re.

In the right half of the heart, a definite constriction of the

endothelial tube marks the limit between the ventricular and

auricular portions. On the left side there is no such constriction.

Posterior to this constriction the right endothelial tube widens out

suddenly, reaching about three times its width in the constricted

region. The left endothelial tube in this region widens only very

slightly. ‘The right and left heart primordia furthermore show

considerable asymmetry as regards curvature, for while the

portion of the left tube lying lateral to the open fore-gut is prac-

tically straight, the right tube in this region shows well marked

DEVELOPMENT OF THE HEART IN MARSUPIALS. 479

curvature. (See Pl.II. fig.6.) Correlated with this difference in

the size and curvature of the endothelial tubes, the opening

of the fore-gut is also asymmetrical (fig. 6, AJ.P.). At the

anterior intestinal portal, the right and left primordia of the

heart separate, and both tubes become reduced in size, the right,

however, more markedly than the left. The endothelial heart-

tubes pass imperceptibly into vitelline veins.

A further distinction between the ventricular and auricular

portions of the heart lies in the fact that in the anterior region

the myocardium is separated from the endothelium by a con-

siderable space crossed by fine strands of protoplasm (text-fig. 17),

while in the posterior portion the myocardium is closely applied

to the endothelium (text-fig. 18). The transition between these

two conditions takes place gradually in the region of the atrio-

ventricular constriction of the right side.

Text-figure 18.

Perameles obesula (10.viii.03).

A. Transverse section through auricular portion of the heart.

B. Complete outline of same section.

End. Endothelium. G. Gut. My. Myocardium.

Turning now to the blood-vessels, two aortic arches are complete.

Their relations are seen most clearly in the side view of the model

(PI. II. fig. 8, 4.1 & 2). From the ventral view (fig. 6) it will be

seen that there are a number of capillaries lying against the gut-

wall, between the roots of the right and left mandibular arches.

These are probably derived from the scattered angioblast cells in

the corresponding position in Stage ITI., and are doubtless destined

to contribute to the formation of the median ventral aorta which

is established in the next stage (Stage VI.). The first aortic arch

runs forwards to the anterior end of the gut and is there connected

by a well developed loop, situated laterally to the apex of the

fore-gut, with the corresponding dorsal aorta. The aorta is paired

480 MISS K. M. PARKER ON_THE

throughout its length (Pl. II. fig. 7, D.A.). From the anterior

convexity of the loop of the first arch there is given off on each

side a vessel which runs outwards and forwards and then breaks

up into series of capillaries which form a cup surrounding the

primary optic vesicles. These capillaries run round the postero-

lateral face of the optic vesicle to become connected dorsally

with the veins of the head (fig. 7).

The second aortic arch arises from the dorsal side of the bulbus,

runs backwards and outwards to loop round the gut in the hyoid

arch and to join the dorsal aorta (see Pl. I. figs. 6-8, Ap 2).

From the dorsal aorte posterior to the second aortic arch,

there arises a pair of small sprouts running outwards on the gut-

wall. These are the dorsal elements of the third aortic arch.

The best developed venous trunk of the head, viz. the vena

capitis medialis, is clearly seen in the dorsal view of the model

(fig. 7, V.C.M.). Each isa small vein lying dorsal to the dorsal aorta

Text-figure 19.

Perameles obesula (10.viii.03). Transverse section through the Cuvierian ducts.

A.C.V. Anterior cardinal vein. ©.D. Cuvierian duct. .D.A. Dorsal aorta. End.

Endothelium. 22.P. Medullary plate. My. Myocardium. V.C.M. Vena

capitis medialis.

close against the neural tube (text-fig. 17, V.C.M.). Anteriorly

this vein runs into the dorsal aorta. In the model this is only

shown on the left side, but high-power examination of the

sections reveals a very fine capillary completing the connection

between the right vein and the aorta. Arising from the vena

capitis medialis in the anterior half of the mandibular arch is a

series of capillaries which are continuous with those surrounding

the optic vesicles. The brain here is widely open and its margin

is situated just to the outer edge of the capillaries. The latter

would accordingly lie medial to the neural crest were such present

in this region. In the region of the first visceral pouch there is

another line of capillaries lying lateral to the vena capitis medialis

and connected with it. From the anterior end of these capillaries

there runs outwards and forwards a vessel connecting them with

a group of capillaries lying in the mandibular mesoderm lateral

DEVELOPMENT OF THE HEART IN MARSUPIALS. 481

and ventral to the gut. The vena capitis medialis runs back

alone for a short distance, and is then again connected with a

more laterally situated capillary. This capillary runs back from

this point to the level of the incipient third aortic arch, and then

runs laterally to be connected with the Cuvierian duct which is

now well developed (fig. 7 & text-fig. 19). On the right side of

the embryo this transverse connection is very incomplete, but not

quite so incomplete as would appear from the model, since the

difficulty of building up these fine capillaries caused some to be

lost in this region. This vein, lying lateral to the vena capitis

medialis and to the primordia of the nerve-ganglia, is the vena

capitis lateralis of Grosser (6) and Salzer (17). From the vene

capitis medialis and lateralis the anterior cardinal vein is derived.

The auditory vesicle lies in the space between the venz capitis

medialis and lateralis immediately dorsal to the second aortic

arch. Running in the hyoid arch area few capillaries apparently

corresponding with the much more conspicuous group in the

mandibular arch.

Text-figure 20.

Perameles obesula (10.viii.03). Transverse section showing the connection of the

vena capitis medialis with the dorsal aorta.

D.A. Dorsal aorta. M.P. Medullary plate. V.C.M. Vena capitis medialis.

V.O.L. Vena capitis lateralis.

- The Cuvierian ducts have increased considerably in size, and

that of the left side opens direct into the lateral heart-tube (text-

fig. 19, C.D.). Behind the opening of the Cuvierian duct a single

small capillary runs posteriorly in the somatopleure representing

the future umbilical vein. The vena capitis medialis, it should

be noted, continues on after the lateral bend of the vena capitis

lateralis (fig. 7). The two are closely connected in the region of

the incipient third aortic arch. ;

One important point which is difficult to observe in the figures

of the model is shown in the sections of the embryo (text-fig. 20),

and that is the fact that the vena capitis medialis at irregular

intervals opens into the dorsal aorta.

In this stage, then, we find the ventricular and auricular

482 MISS K. M. PARKER ON THE

portions of the heart differentiated. Fusion of right and left

primordia has only affected the myocardium and the cephalic ends

of the endothelial tubes. The two halves of the auricular portion

of the heart are wide apart.

Two complete aortic arches are present and one is in process of

formation. Vene capitis medialis and lateralis and Quvierian

ducts are all present.

It is not necessary to give a detailed description of the embryo

of Macropus sp., which is included in this stage, as it differs only

in certain points ‘from the Perameles obesula embryo described

above. In the degree of development of the gut and pharyngeal

pouches, as well as of the nervous system, the two embryos very

closely resemble each other.

Text-figure 21.

Macropus sp. Transverse section through the root of the second aortic arch.

D.A. Dorsal aorta. End. Endothelium. I.P. Medullary plate. My. Myocardium.

Rt.A. Root of aortic arch. .G. Primordium of trigeminal ganglion.

V.C.L. Vena capitis lateralis. V.C.M. Vena capitis medialis.

Two complete aortic arches are present, but there is no trace of

a third.

The vena capitis medialis resembles that of the _Perameles

embryo. The vena capitis lateralis, however, is slightly less

advanced, being only recognisable in the region of the trigeminal

and facial neural crest proliferations, and not extending back as

far as the auditory vesicle. Cuvierian ducts are present, and the

right one at least opens into the heart-tube. The sections are

somewhat broken in this region, so that satisfactory observations

on the openings of the Cuvierian ducts and their relations to

the anterior cardinal veins are impossible.

The heart differs in several respects from that of the Perameles

embryo. The general relations of heart and pericardium and the

mode of origin of the aortic arches are exactly the same in the

DEVELOPMENT OF THE HEART IN MARSUPIALS., 483

two embryos. In the Macropus embryo, as in Perameles, the

myocardium is fused in the middle line throughout the length of

the closed gut. A slight groove marks the line of fusion in the

posterior portion of the heart, but there is no indication of a

ventral mesentery at any point. The ventricular region of the

heart is distinguished from the auricular by the fact that in

the ventricular portion a considerable space intervenes between

endothelium and myocardium, whereas in the auricular portion

the two layers are close together. The limit between the two

divisions is further indicated by a constriction of each endo-

thelial tube, which then widens out abruptly to form the auricle.

Right and left endothelial tubes are united anteriorly in the

region of the bulbus arteriosus, just as in the Perameles embryo.

Behind this again they diverge around the opening of the gut.

It may be noted that in the region of the widest divergence of

the endothelial tubes a fine bridge runs across and connects

the two.

The curvature, so far as it can be made out without reconstruc-

tion, is similar to that of the Perameles embryo. The asymmetry

appears to be less marked than in the latter embryo, but on this

point it is impossible to make a positive statement without recon-

struction. Throughout a considerable portion of its length,

however, the right endothelial tube is larger than the left, just

as in the Perameles embryo.

The Macropus embryo then differs from the Perameles embryo

of a similar stage mainly in the configuration of the endothelial

tubes, which are joined at their cephalic extremity, then widely

separate for some distance (text-fig. 21), and then again approxi-

mated, though not joined. This difference, as well as the slight

differences in the myocardial wall, may very probably be due to

slight dissimilarity in the positions of the endothelial tubes and

the myocardium before union of the latter.

Srace VI. Macropus ruficollis.

The material for this stage consists of a single embryo of

Macropus ruficollis, cut transversely.

Greatest length of embryo, 5:2 mm.

Dorsal perimeter, about 13°56 mm.

The embryo is sharply bent in front of the fore-limb buds, so

that the head, invested by proamnion, is sank down into the yolk-

sac and forms an acute angle with the trunk. No trunk amnion

is yet developed. The brain, though open in the fore- and mid-

brains, is closed in the region of the hind-brain.

The gut is closed as far back as the third well-developed somite.

Three visceral pouches are present.

The heart shows a considerable advance on the preceding stage.

The right and left heart-tubes are fused except in the region of

the sinus venosus, where they remain separate, while the Cuvierian

484 MISS K. M. PARKER ON THE

ducts, which are now established as wide vessels, pass across the

coelomic cavity to open into the right and left heart-tubes (text-

fig. 22),

pan of the median portion of the heart has resulted in

the definite establishment of a U-shaped ventricular limb and an

auricular portion extending from the left dorsal side of the ven-

tricle posterior to its apex and separating, at the opening of the

gut, into the right and left halves of the sinus venosus.

Text-figure 22.

gt og

"Rig

Ceara wie

ako,

"385

o7*

eles

+ a,

“4

f ti

806%

Macropus ruficollis. Transverse section through the Cuvierian ducts.

A.O.V. Anterior cardinal vein. C.D. Cuvierian duct. .D.A. Dorsal aorta. End.

Endothelium. 2.7. Medullary tube. My. Myocardium. Py.A. Proamnion.

V.C.M. Vena capitis medialis.

The cephalic portion of the S-shaped heart is somewhat curved,

so that, as in the preceding stage, the bulbus (conus) arteriosus

lies dorsal to the cephalic extremity of the ventricle. The bulbus

arteriosus is continued into a short median ventral aorta which

bifurcates to form the first pair of aortic arches. The second and

DEVELOPMENT OF THE HEART IN MARSUPIALS. 485

third pairs of aortic arches arise from the median ventral aorta.

immediately posterior to its bifurcation. The second arch is.

large, the third very small.

Tn correlation with the rapid development of the fore- and mid-

brains at this stage, the head-plexus found in Stage V. (see Pl. Il.

figs. 6-8) has become extended into a long slender vessel, destined

to form the anterior part of the internal carotid artery and lying

on either side of the mid-ventral line in the fore-brain region.

As in the preceding stage, it anastomoses anteriorly with capillaries

arising from the vena capitis medialis.

Both vene capitis medialis (text-fig. 22, V.C.1/.) and lateralis

are present, though neither can be traced continuously throughout

the head-region. The vena capitis medialis extends to the ante-

rior end of the brain, lying close to the medullary tube, dorsal to

the dorsal aorta and the internal carotid artery. It is discon-

tinuous in the region of the auditory vesicle, where no veins are

recognisable. Posterior to that, it runs back as a continuous

trunk to the level of the Cuvierian duct, and beyond this is

recognisable as a minute vessel lying close to the neural tube in

the trunk region.

Lying lateral to the primordium of the trigeminal nerve, there

are a few scattered capillaries which represent discontinuous

segments of the vena capitis lateralis. Immediately posterior to

the primordium of the trigeminus, the vena capitis lateralis arises

from the vena capitis medialis and runs back as a small vessel

lying lateral to the root of the facial nerve. Venez capitis lateralis

and medialis are interrupted in the region of the auditory vesicle,

but both are present immediately posterior to it. At irregular

intervals on their course there are transverse communications

between the two veins. The vena capitis lateralis does not form

a continuous trunk in the region posterior to the auditory vesicle,

but immediately anterior to the point of separation of right and

left heart-tubes it increases markedly in size and is connected

by a wide anastomosis with the vena capitis medialis, which

becomes very small posterior to this level. The enlarged vena

capitis lateralis, or, as it may here be called, anterior cardinal

vein, passes ventro-laterally and, running alongside the dorsal

aorta for a short distance, finally opens into the Cuvierian duet

in the manner seen in text-fig. 22.

The umbilical vein is now present, running in the somatopleure

and opening into the Cuvierian duct.

In this stage, then, we have a heart in which fusion of the

right and left primordia has occurred except in the region of the

sinus venosus, and curvature has carried the auricular limb into.

position dorsal to the ventricle. Three aortic arches are present,

vene capitis medialis and lateralis are well established though

discontinuous and open vid the Cuvierian ducts into the sinus

venosus.

486 MISS K. M. PARKER ON THE

SuMMARY AND Discussion.

A. Development of the Heart.

From the foregoing description it is evident that the early

development of the heart in such Marsupials as Perameles and

Dasyurus proceeds along essentially the same lines as in Kutheria.

The early stages of the heart development in the latter have been

described by a number of investigators (e.g. Mollier (15) ); but

although the broad outline of the process may be said to be well

known, there is still considerable difference of opinion with regard

tocertain points. It will be useful, therefore, before summarising

the preceding observations, to briefly review the literature on the

subject.

With regard to the lateral paired primordia of the heart little

need be said at this point. The heart endothelium arises between

the entoderm and the splanchnic mesoderm, from which latter

it is either partially or wholly derived. The primordia of the

heart-tubes are first recognisable in the hind-brain region and

grow forwards at the expense of angioblastic cells proliferated off

from the splanchnic mesoderm, which is itself thickened and

indented to form the primordium of the myocardium. It should

be noted that in the earliest stages examined (PI. I. figs. 1 & 2)

the endothelium lies to the medial side of the pleuro-pericardial

canals in the anterior region and to the lateral side in the pos-

terior region. To this point reference will be made subsequently

in connection with the discussion of the reversal of the peri-

eardium which, according to some authors, takes place at the

time of formation of the head-fold.

The processes which bring the lateral heart primordia into

position below the fore-gut relate primarily to the formation of

the head-fold, and it is therefore necessary to get a clear idea of

the mode of closure of the gut before considering the problems

relating to the fusion of the lateral primordia of the heart.

Some authors (e.g. Robinson (13) ) hold that the formation of

the fore-gut is due mainly, if not entirely, to the rapid growth

of the embryo over the relatively stationary line between the

embryonal and extra-embryonal areas. Thus Robinson (18) says:

‘“ The orifice (of the umbilicus) is not reduced in size during the

early stages of development by the convergence of its margins

towards a central point. This being the case, no tucking off of

the embryo from the surface of the ovum can occur; on the con-

trary, what does occur is almost the exact opposite of such a

process, for the margin of the area remains as a relatively

slow-growing region, whilst the embryonic and extra-embryonic

portions of the wall of the ovum rapidly increase in extent.

Under these circumstances, it follows that the margin of the

embryonic area will soon appear as a ring between the upper or

embryonic and the lower or extra-embryonic parts of the ovum,

both of which have expanded beyond it in all directions.”

DEVELOPMENT OF THE HEART IN MARSUPIALS. 487

While the forward growth of the brain-plate doubtless plays

an important part in the initiation of the formation of the fore-

gut, this explanation does not account for the conditions revealed

by reconstructions of the gut and pericardium in the early stages

of head-fold formation.

On the other hand, various investigators (e.g. Rouviére (14),

Graper (4) ) contend that there occurs, in addition to the forward

growth of the brain-plate, a backward progression of the anterior

intestinal portal, whilst in older accounts an actual fusion of

lateral folds in the mid-ventral line was assumed. Both Robinson

(13) and Rouviere (14) give excellent reasons for regarding this

assumption as erroneous. They point out that if gut-closure

were effected by the fusion of lateral folds (such as are shown in

text-fig. 9), the heart would remain in connection with the gut by

a dorsal mesocardium and with the yolk-sac wall by a ventral

mesocardium. Robinson denies the existence of a ventral meso-

cardium in mammals, and quotes this fact in support of his theory

that the separation of the gut from the yolk-sac is due to growth

of the embryo rather than to fold-formation. Rouviére, on the

other hand, while he agrees with Robinson as to the absence of a

ventral mesocardium in mammals, gives a different account of

the process of gut-closure. He describes the formation of lateral

pleuro-pericardial canals which grow forwards round the anterior

end of the brain-plate and fuse to form a continuous cavity.

The splanchnopleure forming the posterior wall of the pleuro-

pericardial cavity now forms a continuous fold which Rouviére,

following Tourneux, calls the ‘‘cardiac fold” (compare text-fig. 12,

C.F.) and which he describes as growing actively backwards as

a whole.

In the chick, on the other hand, a ventral mesocardium is

present, but this is due, as Robinson points out, to the relatively

late penetration of the mesoderm in the head region. The pleuro-

pericardial canals do not extend round and unite in front of the

medullary plate in early stages, but only at a later stage do they

penetrate into the floor of the fore-gut after that has been formed.

The lateral cavities therefore do not at once become continuous,

but remain separated from each other by a double layer of

mesoderm constituting the ventral mesocardium.

With regard to mammals, Rouviére, while he does not discuss

the influence of the forward growth of the brain-plate, concludes

that the crescent-shaped cardiac fold grows backwards as a whole,

and that the free edge of the splanchnopleural fold progresses

always in advance of the primordia of the heart, so that no fusion

of the splanchnopleure is involved and no ventral mesocardium

is formed.

Griiper, in a description of the growth processes in the

developing chick, which he worked out by staining the living

embryos and keeping them under observation while still alive,

shows that there is considerable evidence in support of the view

that the margin of the fore-gut (anterior intestinal portal) moves

488 MISS K. M. PARKER ON THE

backward concurrently with the forward growth of the brain-

plate. He gives a series of comparative measurements which

show that the rate of removal of the lip of the anterior intestinal

portal from the anterior end of the brain is greater than the rate

at which the brain-plate grows forward from a given fixed point ;

hence it is evident that the anterior intestinal portal must be

moving backwards.

Concurrently with the formation of the fore-gut, the lateral

heart-tubes come to lie ventrally to it, but do not at once fuse.

Wilson (20), in a paper on young human embryos, draws attention

to this fact and refers to the embryo of Perameles nasuta, described

in this paper as Stage III., as exemplifying this condition ; but

he does not discuss the question as to how these lateral hearts

approach one another.

We may now consider the evidence afforded by the material

described above, and will endeavour to show that it is entirely

in accord with the view that there is actual backward growth of

the anterior intestinal portal, and that it is this process, and

not fusion of lateral folds, that brings about lengthening of the

fore-gut.

It may be noted here that in Perameles, as in the rabbit

(Rouviere), no ventral mesocardium is present at any stage, a fact

which, in itself, is a strong argument against the theory that gut-

closure is effected by the fusion of lateral folds.

If we compare a stage in which the head-fold has not yet

appeared (Pl. I. figs. 1 & 2) with one in which a small portion

of the fore-gut is differentiated (figs. 3 & 4), we see that the pleuro-

pericardial ccelom not only moves backward relatively to the

brain-plate but also increases very considerably in width. It is

obvious that such an increase in size must either cause the

pericardium to extend peripherally or to close in towards the

axial line, and it is perfectly clear on comparison of figs. 2 & 3

that it is this latter process which is taking place. From a

longitudinal section, such as is shown in text-fig. 4, it is evident,

moreover, that such an expansion of the pericardium must involve

the backward growth of the splanchnopleural floor of the fore-

gut. If the lengthening of the fore-gut were due entirely to the

rapid forward growth of the brain-plate, there would be no such

inward closure of the pericardial region. Moreover, if we compare

figs. 2 & 3 (Pl. L.), we see that in the earlier stage, the pericardial

colom is situated peripherally to the margin of the brain-plate,

while in the second stage, the inner margin of the pericardium

lies in the lip of the anterior intestinal portal. Now the growth

in length of the brain-plate im the period between these two

stages would naturally give rise to a fold round its anterior

margin, but would not bring the pericardium into the position it

occupies in Stage II. (fig. 3), unless there occurred concurrently

with such growth in length either an increase in width of the

brain or an inward closure of the pericardium. Comparison

of figs. 2 & 3 again shows that while no increase in width of the

DEVELOPMENT OF THE HEART IN MARSUPIALS. 489

brain-plate has occurred, the pericardium has actually closed

in towards the axial line. Precisely the same conclusion may be

reached from a comparison of figs. 1 and 4, but as the interval

between the stages is greater and the embryos are not of the

same species, less importance attaches to them in this con-

nection. ;

It has already been remarked that, in the chick, the develop-

ment of the pleuro-pericardial canals occurs at a later period

than in the mammal, so that in this type the form of the head-

fold in early stages cannot be affected by growth of the

pericardium. If we compare the shape of the fore-gut in a chick

of two somites with that of Dasyurus Stage II. (text-fig. 23 A &

Pl. I. fig. 3), we see that in the first stage of head-fold formation

in the bird, the outline of the anterior intestinal portal is broadly

Text-figure 23.

Anterior end of chick of (A) 2 somites, (B) 4 somites, to show the relations of the

head-fold, brain-plate, anterior intestinal portal and _ pleuro-pericardial

cavities.

A.J.P. Anterior intestinal portal. F.G. Fore-gut (outline). JF. Medullary fold.

M.M.P. Margin of the medullary plate. P.p.C. Pleuro-pericardial cavity.

erescentic, while in Dasyurus it is U-shaped. This difference

I conceive to be due to the fact that in the chick, no factor but

the forward growth of the brain-plate is operating at this stage,

while in the mammal, in addition to this process, the expansion

of the pericardium is already bringing about the formation of

lateral folds and the consequent narrowing of the anterior

intestinal portal. A slightly later stage of the chick (text-fig.

23 B) shows an approximation to the mammalian condition, for

the pleuro-pericardial canals have appeared and are progressing

towards the middle line; lateral folds have therefore arisen and

the outline of the anterior intestinal portal is U-shaped.

' We may therefore conclude, that while the forward growth of

490 MISS K. M. PARKER ON THE

the brain-plate initiates the formation of the head-fold, there

occurs concurrently with this process in the mammal, and ata

slightly later stage in the chick, a rapid expansion of the peri-

cardium and a consequent backward and inward growth of the

fold of splanchnopleure which constitutes the inner margin of

the pleuro-pericardial cavity.

Additional evidence is afforded by the study of the longitudinal

sections and reconstructions of Stages III. & IV. (text-figs. 12 &

15; Pl. 1. figs. 4&5). From the longitudinal sections, it is evident

that a great increase in length of the brain has occurred in the

mid- and fore-brain regions. If, therefore, the increase in length

of the fore-gut were due to elongation of the brain-plate, a corre-

sponding increase should occur in the portion of the fore-gut

lying below these segments, 7.e. the portion anterior to the first

visceral pouch in Stage III. (fig. 4). Comparative measurements

of the gut in figs. 4 & 5 show, however, that no increase in length

has occurred anterior to the first visceral pouch. Moreover,

growth of the medullary plate would not necessarily bring about

lengthening of the fore-gut unless it occurred along a straight line

representing the longitudinal axis of Stage III, 7.e. unless the

brain remained unflexed. The positions of the auditory neuro-

mere in fig. 4 (opposite the second visceral pouch) and fig. 5

(opposite the first visceral pouch), show that the brain-plate

has moved forward relatively to the gut between Stages IIT. &

1V. If, now, we study the longitudinal section of Stage IV.

(text-fig. 15) we see the conditions resulting from the increase in

length and forward growth of the brain-plate. The gut has

increased in dorsi-ventral extent, the medullary plate projects

considerably anterior to the cephalic limit of the gut, and

flexure has occurred at two points; that is to say, the rapid

growth of the fore- and mid-brains, instead of involving a longi-

tudinal stretching of the portion of the gut lying ventral to

them, has caused little or no increase in length of the embryo

along its straight long axis: the additional extent of the brain-

plate is accommodated within the limited space by flexure.

We see, therefore, that although the brain-plate lengthens

rapidly after the first establishment of the head-fold, we can safely

conclude that this does not cause elongation of the gut, for the

regions of greatest growth of the gut and brain-plate are not

correlated and the value of the forward growth of the brain as a

factor in the lengthening of the fore-gut is largely rendered

nugatory by the occurrence at this period of the cranial flexure.

On the other hand, there is little difficulty in interpreting the

progressive closure of the gut at this stage as being due to an

entirely different cause, for the median pericardium has extended

rapidly, its antero-posterior length in the middle line having

more than doubled in the short developmental period elapsing

between Stages ITI. and IV. (PI. I. figs. 4 & 5). As the anterior

margin of the pericardium is in contact with the ectoderm of the

head-fold, the rapid expansion of the cavity naturally involves a

DEVELOPMENT OF THE HEART IN MARSUPIALS. 49]

closure inwards, towards the axial line, of the fold of splanchno-

pleure limiting the gut.

It is necessary now to consider the early development of the

heart in relation to the mode of closure of the gut described

above. It is evident, as Rouviere points out, that if gut-closure

be effected by the backward movement of the cardiac fold, no

ventral mesocardium will be formed at any stage in forms,

such as mammals, in which a continuous pericardial cavity is

present prior to head-fold formation. It remains to be considered

how the lateral primordia of the heart reach their position in

the dorsal wall of the median pericardium. Various authors, e. g.

Robinson (13), have assumed that as the head-fold forms, the

pericardium undergoes a complete reversal, so that its anterior

Text-figure 24.

Transverse sections through embryos of Dasyurus viverrinus, (A) Stage I. (7°5 min.),

(B) Stage II. (85 mm.), to show the direction of extension of the pericardial

cavity.

D.A, Dorsal aorta. Had. Endothelium. 22.P. Medullary plate.

P.p.C. Pleuro-pericardial canal.

wall becomes posterior and its ventral wall, dorsal. Of such a

process of reversal, the longitudinal sections figured here (text-

figs. 4, 5 & 12) give no evidence. Moreover, in the anterior

region of the pericaruium, the primordium of the heart on each

side of the embryo hes at or near the inner, medial margin of

its pleuro-pericardial canal, so that a reversal which affected the

anterior limb of the pericardium would indeed bring the heart

primordia into position ventral to the gut, but would carry them

also to the lateral margins of the gut, a position which they do

not occupy.

On the other hand, if we take into consideration the fact that

Proc. Zoou. Soc.—1915, No. XXXIV. v4

492 MISS K. M. PARKER ON THE

the heart primordia lie at the medial margin of the pleuro-

pericardial canals, we see that the inward progression of the edge

of the splanchnopleural fold in the direction indicated by an

arrow in text-fig. 24 A, will bring about the conditions shown in

text-fig. 24 B. “(Compare also text-fig. 8, where the relations are

essentially the same and probably approximate more closely to

those in the living embryo.)

From this stage, it is evident that when the lateral limbs of

the pleuro-pericardial canals become incorporated in the median

pericardium by the backgrowth of the cardiac fold, the heart

primordia will lie in the dorsal wall of the pericardium and will be

situated towards the middle line of the gut. (See text-fig. 8.)

Thus, as Rouviere (14) indicates in describing similar con-

ditions in the rabbit, the position of the heart primordia in

such a stage as is represented in text-fig. 24 B, is brought about

‘by the inward extension of the lateral prolongations of the

cardiac fold.” There is no evidence of reversal of the peri-

eardium, nor is there adequate ground for assuming that such

a process occurs

We can, therefore, gain a clear conception of the way in

which the lateral heart primordia attain the position they occupy

in Stage III. (Pl. I. fig. 4; text-fig. 8) lying side by side below the

closed fore-gut.

In order to complete the history of the early development of

the heart, it is now necessary to consider the mechanism which

brings the heart-tubes into contact in the middle line.

If we compare figs. 4 and 5 (Pl. I.) we see at once that while

the pericardium has increased rapidly in the antero-posterior

direction, it has not increased in transverse width and, in fact, at

the point of closest approximation of the heart-tubes, an actual

decrease in width has occurred; that is to say, the pericardium

at this stage is growing in the antero-posterior direction at the

expense of its transverse width. This fact suggests a simple

explanation of the approximation of the heart- tubes after gut-

closure, for it may be supposed that if the total width of the

pericardium is reduced by this process of stretching, the distance

between the heart-tubes will decrease until they meet each

other in the middle line *.

The heart-tubes, in the period following immediately on their

reaching the middle line, grow very rapidly, so that, in the next

stage (V.); we find various forms of curvature which serve to

accommodate the increased length of the heart. In the embryo

of Perameles obesula described in this stage (PI. II. fig. 6), the

heart-tubes are in contact through a great portion of their length

and here follow a parallel curved course. The separate heart-tubes

lying in the lips of the anterior intestinal portal, however, show

a marked difference from each other both in their length and the

* The suggestion that the approximation of the heart-tubes is due to such a

erowth in length without compensatory growth in width was made to me by

Professor Hill.

DEVELOPMENT OF THE HEART IN MARSUPTALS. A9Q3

form of curvature, the right primordium being larger and more

markedly curved than the left. This asymmetry occurs to a

somewhat less extent in the Onychogale embryo of this stage,

and also in a number of embryos of Dasyurus viverrinus of about

the same stage, so that evidently at this period the right and

left primordia of the heart develop independently of each other.

It may be suggested that the greater length of the right heart-

tube is to be accounted for by the fact that it is destined to

form the convex, longer side of the completed ventricular

limb.

B. Development of the Cardinal Veins.

Before summarising the results of my observations on the

development of the cardinal veins, more particularly the anterior

cardinals, it may be useful to give a short resumé of previous

work in this field.

Hoffmann (7) in 1893 described the development of both

anterior and posterior cardinals in Selachians by the formation

of a series of offshoots from the dorsal aorte. These oftshoots

become connected on each side to form a continuous longitudinal

trunk. He figures capillaries lying on both medial and lateral

sides of the auditory vesicle, but makes no comment thereon.

Salzer (17) in 1895 descr ibed the dev elopment of the anterior

cardinal veins in the guinea-pig. According to him, the first

vein of the head arises on the medial side of the cranial ganglionic

primordia. A vein next arises lying lateral to the ganglionic

primordia of nerves VIT., 1X. & XS aad to the auditory estes.

This vein, which Salzer calls “‘ vena capitis lateralis,’ communi-

cates with the medial vessel and seems to be formed from a series

of lateral outgrowths from it. The medial vessel degenerates in

the region of nerves VII. to X., so that for a time there is a

condition in which the vein of the head runs medially to the

trigeminal nerve, then, passing laterally, runs outside nerves VII.,

TX. -& X. and the auditory vesicle, and finally passes round the

medial side of nerve XII. before opening into the Cuvierian duct.

In subsequent stages, the process of development of the lateral

trunk is continued anteriorly and posteriorly in the region of

the trigeminal and hypoglossal. Thus the definitive anterior

cardinal vein runs laterally to all the cranial nerve-roots.

Grosser (6) in 1907 gave a similar description of the develop-

ment of the anterior cardinals throughout the vertebrate series.

He calls the vein lying medial to the nerve-roots the vena capitis

medialis, and the lateral vessel the vena capitis lateralis. The

former develops first and lies close against the neural tube.

From it are given off lateral vessels which become connected on

the outer side of the nerve-roots to form the vena capitis lateralis.

The vena capitis medialis persists only at its anterior end, the

rest of the anterior cardinal being derived from the vena capitis

lateralis,

34*

494 MISS K. M. PARKER ON THE

Turning now to the facts revealed by the foregoing study of

Perameles, we find both the vene capitis medialis and lateralis

present. In Stage ITI. the first traces of the venous system

of the head are present in the form of isolated segments of the

vena capitis medialis. Further, in this stage, in the region of

the somites, there is on each side a series of dorsal offshoots from

the dorsal aorta (text-fig. 11) partially connected to form an as

yet incomplete longitudinal vessel lying, like the vena capitis

medialis, close against the neural tube. This vessel Hoffmann

(7) described as representing the primordium of the posterior

cardinal vein. It is worthy of note that whilst the origin of this

vein from the dorsal aorta is thus clearly demonstrated, no

connection between the anterior segments of the vena capitis

medialis and the dorsal aorta could be observed, even after

careful study of the individual sections under the high power.

In the next stage (IV. P. nasuta 2 P), however, the vena capitis

medialis, though not forming a continuous longitudinal trunk, is

recognisable throughout the head region and is connected at

irregular intervals with the dorsal aorta. Furthermore, the

vena capitis medialis in this stage gives off lateral capillaries

which anastomose to form the primordium of the vena capitis

lateralis. In the somitic region we find again a series of inter-

segmental offshoots from the dorsal aorta. The vene capitis

lateralis and medialis continue to develop side by side, giving rise

to the condition shown in Pl. II. fig. 7 (V.C.L. and V.C.IL).

(See also text-fig. 25.) In this stage the dorsal aorta and the

vena capitis medialis are connected by small capillaries (see text-

fig. 20) whilst anteriorly the two vessels pass into continuity with

each other*. From the material available it is not possible to

say definitely how these connections arise, but the facts suggest

that the vena capitis medialis is derived from the dorsal aorta.

This view is further supported by the existence in the trunk

region of a longitudinal vessel which is undoubtedly formed from

a series of outgrowths from the dorsal aorta (text-fig. 11). This

vessel apparently bears the same relation to the posterior cardinal

that the vena capitis medialis does to the anterior cardinals, 7. e.

it gives origin to capillaries which contribute to the formation of

the posterior cardinal. The origin of the vena capitis medialis

from the dorsal aorta cannot, however, be regarded as proved, for

in the first stage in which it is recognisable, no connection with

the dorsal aorta could be traced ; in the two following stages (IV.

and V.) the connection is established and is lost in all subsequent

stages (e.g. VI.).

From the descriptions of Salzer (17) and Grosser (6) it seems

that in the forms which they have studied, the vena capitis

medialis fuses in its entirety with the vena capitis lateralis, and

* Professor Hatta tells me that he has found this condition also in the embryo

ot the lamprey.

DEVELOPMENT OF THE HEART IN MARSUPIALS. 495

the anterior cardinal vein formed by the fusion of these two

vessels passes ventro-laterally to open into the Cuvierian duct.

This description is not, however, applicable to Perameles. In

Stage V. (Pl. Il. fig. 7) the vena capitis medialis is continued

backwards into the trunk region of the embryo. The vena capitis

lateralis lies parallel with it and communicates with it repeatedly

in the anterior region; it then diverges from it and constitutes

here the vessel usually known as the anterior cardinal vein,

opening into the Cuvierian duct. The posterior prolongation of

the vena capitis medialis continues as a small vessel lying aiongside

Text-figure 25.

Diagram of the relations of the ven capitis medialis and lateralis to the

primordia of the nerves. Viewed from the dorsal aspect.

A. Stage V. Perameles obesuéa (10.viii.03). B. Stage with twelve cranial nerves,

Perameles nasuta (43.vii.05).

A.C.V. Anterior cardinal vein. A.V. Auditory vesicle. ©O.D. Cuvierian duct.

HB. Hind-brain. V.C.L. Vena capitis lateralis. V.C.M. Vena capitis

medialis. V. Trigeminal nerve. VII. Facialnerve. LX. & X. Common

root of glosso-pharyngeal and vagus nerves. XII. Hypoglossal nerve.

the neural tube in the trunk region. The Cuvierian duct arises

as a relatively large vessel lying in the sematopleure dorsal to

the posterior portion of the lateral heart-tubes. From its

posterior extremity there runs back a series of capillaries which

anastomose with capillaries arising from the prolongation of the

vena capitis medialis. From this line of capillaries, which thus

496 MISS K. M. PARKER ON THE

shows a double origin, the posterior cardinal vein is undoubtedly

derived, but the details of the process of development of the

postcardinals cannot satisfactorily be worked out in the material

available. The above account, however, agrees with that of

Evans (2) for the chick with regard to the origin of the post-

cardinal from capillaries derived partly from the Cuvierian duct

and partly from a vessel lying close to the neural tube.

The vena capitis lateralis in Stage V. is connected anteriorly

with the primary head capillaries arising from the first aortic

arch and also with groups of capillaries in the mandibular and

hyoid arches (see Pl. IT. figs. 6-8).

In subsequent stages, the development follows the course

described by Salzer (17) and by Grosser (5). Thus in a stage in

which twelve cranial nerves are established (Perameles nasuta,

13. vil. 05), the anterior cardinal vein runs medial to nerves V.

and XII. and lateral to VII., [X., and X., and to the auditory

vesicle (text-fig. 25 B); i.e., the portion in the region of and

anterior to the trigeminal nerve and also that posterior to the

vagus, is derived from the original vena capitis medialis, the

intervening portion from the vena capitis lateralis. Traces of

the vena capitis medialis are, however, still present on the medial

side of nerves VII., [X., and X.

Florence Sabin (16), in a recent note on the development of

cardinal veins in the chick, supports the view that the cardinal

veins are derived from the dorsal aorta, She, however, states :

“The part of the head vein which lies close to the neural tube

arises from the arch of the aorta and is a part of the vascular

system of the central nervous system; the caudal part of the

head vein arises directly from the aorta.” In this respect my

results differ somewhat from hers, for in Perameles the vena

capitis medialis (7. e. ‘the part of the head vein which lies close

to the neural tube”) is present before there is any trace of the

capillaries arising from the arch of the aorta (Stage III.). It is

indeed secondarily connected with these, but as is shown in

Pl. II. fig. 7 (V.C.M.) it also extends up to the extreme anterior

end of the head in close relation to the dorsal aorta with which,

in fact, it fuses. Since this vein exists before the formation of

the head capillaries which connect it with the arch of the aorta,

it obviously cannot be derived from that arch.

In seems, therefore, that in Selachians (Hoffmann (7)), the

chick (Evans (2), Florence Sabin (16)), and also in Perameles and

Macropus, there exists in the primary condition a continuous

vessel lying close to the nerve-cord throughout its length. This

vessel is derived in the posterior and probably also in the anterior

region from the dorsal aorta. It contributes to the formation of

both anterior and posterior cardinal veins.

It may be concluded that the presence in early stages of a vein

lying close against the neural tube throughout its length is

correlated with the relatively great importance of the central

DEVELOPMENT OF THE HEART IN MARSUPIALS. 497

nervous system in these stages. The brain and spinal cord, being

the first organs to attain any considerable degree of development,

are naturally the first to receive a vascular supply, and both venze

capitis later: alisand medialis persist for some time, forming a rich

supply of capillaries to the brain and surrounding the developing

cranial nerves.

KINAL SUMMARY AND CoNCLUSIONS.

The facts revealed by the study of early stages in the

development of Marsupials point to the conclusion that while

the initiation of head-fold formation is in all probability due to

the forward growth of the brain-plate, there occurs also an active

backward growth of the anterior intestinal portal. This process

is associated with the rapid expansion of the pericardium which

occurs at this period of development, and which brings about the

backward and inward growth of the layer of splanchnopleure

limiting the pericardium.

In the course of this inward closure, the pericardial cavity

extends to the ventro-lateral and finally to the ventral side of the

lateral primordia of the heart, so that when the lateral portions of

the pericardium become incorporated in its median limb, the

heart primordia lie in the dorsal wall of the pericardium.

The approximation of the heart-tubes after gut-closure is

due to the fact that, at this period, the pericardium grows

rapidly in length and decreases in width so that the heart-tubes

are brought together by longitudinal stretching of the pericardial

wall lying between them.

Curvature of the heart is due to its rapid growth at a period of

less active extension of the pericardium.

The first two aortic arches in Perameles are typical, and the

development of the veins of the head resembles that process in

other mammals in that the anterior cardinal vein is derived from

persistent portions of two primitive head-veins, the venz capitis

medialis and lateralis. The posterior continuation of the vena

capitis medialis also contributes to the formation of the posterior

eardinal vein and is itself derived from the dorsal aorta.

REFERENCES To LYrERATURE.

1. Bremer, J. L.—‘“‘ The Development of the Aorta and Aortic

Arches in Rabbits.” American Journal of Anatomy,

vol. XXX.

2. Evans, H. M.—‘‘ On the Development of the Aorte, Cardinal

and Umbilical Veins and other blood-vessels of Vertebrate

Embryos from Capillaries.” Anatomical Record, vol. 111.

3. Evans, H. M.—Development of the Vascular System, in

Keibel and Mall’s ‘ Text-book of Human Embryology.’

498

MISS K. M. PARKER ON THE

GrarEr, L.—“ Beobachtung von Wachstumsvorgingen an

Reihenaufnahmen lebender Hithnerembryonen nebst

Bemerkungen uber vitale Firbung.” Archiv fiir Ent-

wicklungsmechanik der Organismen, iB deixexocnil

. GROSSER, O. & BrREzINA, E_* Ueber ‘die Entwicklung der

Venen des Kopfes und des Halses bei Reptilien.” Morph.

Jahrb. Bd. xxxiii., 1895.

. GrosseR, O.—‘‘ Die Elemente des Kopfvenensystems der

Wirbeltiere.” Werh. d. Anat. Ges. Erg.-Heft. z. Anat.

Anz. Bd, xxx., 1907.

. Horrmann, C. K.—“ Zur Entwicklungsgeschichte des Venen-

systems bei den Selachiern.” Morph. Jahrb. Bd. xx.,

1893.

. Hocusrerrer.—‘ Die Entwicklung des Blutgefiasssystems.”

Hertwig’s Handbuch d. vergl. u. exper. Entwicklungs-

lehre d. Wirbeltheire, Bd. 11. 2, 3.

Keiser, W.—“ Untersuchungen iiber die erste Anlage des

Herzens, der beiden Liingsgefiissstamme und des Blutes

bei Embryonen von Petromyzon planeri.” Jena. Zeitschr.

fix Naturw., vol. lvii.

. Lewss, F. T.—‘“‘ The Intra-embryonic Blood-vessels of Rabbits

from 84 to 13 days.” Amer. Journ. Anat. vol. iii.

. Matt, F. P.—‘* On the Development of the Blood-vessels of

the Brain in the Human Embryo.” Amer. Journ, Anat.

vol. iv.

. Mitten & McWuorrer.—“ Experiments on the Development

of Blood-vessels in the area pellucida and embryonic body

of the Chick.” Anat. Record, Aprii 1914.

. Rosrnson, A.—‘‘The Early Stages of Development of the

Pericardium.”” Journal of “Anatomy and Physiology,

vol. xxxvii., 1902.

. Rovuvibre, H.—‘‘ Etudes sur le développement du péricarde

chez le lapin.” Journal de Anatomie, vol. xl., 1904.

. RtcKxerr & MouiiiEer.—‘ Die erste Entstehung der Gefiisse

und des Blutes bei Wirbeltieren.” Handb. d. vergl. u.

exper. Entwicklungslehre d. Wirbeltiere, herausg. von

O. Hertwig, Bd. 1.

. Sasin, F. R.—‘‘ On the Origin of the Duct of Cuvier and

the Cardinal Veins.” Proceedings of Amer. Assoc. of

Anatomists, Anat. Record, vol. ix. No. 1, 1915.

. Sauzer, H.—‘* Ueber die Entwicklung der Kopfvenen des

Meerschweinchens.” Morph. Jahrb. Bd. xxiii., 1895.

. Scoutrr, H. von W.—“ Early Stages of Vasculogenesis

in the Cat with especial reference to the mesenchymal

origin of endothelium.” Memoirs of the Wistar Institute

of Anatomy and Biology, No. 3, 1914.

. Tanner, J.—‘‘ Zur Entwicklungsgeschichte der Kopfarterien

bei den Mammalia.” Morph. Jahrb. Bd. xxx., 1902.

. Witson, J. 'T.—“Observations upon Young Human Embryos.”

Journal of Anatomy & Physiology, vol. xlvii., 1914,

DEVELOPMENT OF THE HEART IN MARSUPIALS,

499

EXPLANATION OF THE PLATS,

Lettering.

A, Auricle. l G. Gut.

A.1, A.2, A.3. First, second and third | H.B. Hind-brain.

aortic arches. | I.C. Internal carotid artery.

A.C. Angioblast cell.

A.C.V. Anterior cardinal vein.

A.T.P. Anterior intestinal portal. | My

A.N. Auditory neuromere. JP

A.V. Auditory vesicle. P.p.C

A.V.C. Auriculo-ventricular con- | NG:

striction.

B.A. Bulbus arteriosus.

C.D. Cuvierian duct.

D.A. Dorsal aorta. VEPs V.P2.

End. Endothelium of heart.

. Margin of the medullary

plate.

. Myocardium.

. Pericardium.

. Pleuro-pericardial canal.

. Trigeminal ganglion,

. Ventricle.

. Vena capitis lateralis.

M. Vena capitis medialis.

First and second visceral

pouches.

F.G. Fore-gut.

Puate I.

Figures 1-5 represent graphic reconstructions of the anterior ends of five embryos,

viewed from the ventral aspect and all drawn at the same magnification.

The outline of the gut is indicated by a broken line and its area is stippled,

except where it is covered by the pericardium. ‘The limits of the peri-

cardium are indicated by a fine black line and its area is coloured grey.

The endothelial heart-tubes, the aorta and the aortic arches are coloured

red. ‘The outline of the brain-plate is shown as a heavy black line where-

ever it was possible to determine its limits. In the region of the neural-

crest proliferation in Stage I., the margin of the medullary plate is

indistinct ; a dotted line here indicates its probable outline.

The outlines of the gut, pericardium, and brain-plate were plotted on one

side only, and the second side is a duplicate of the first, except in the case

of figure 1 (Perameles obesula 1 Z), where the pericardium was plotted on

both sides, as in this instance only was any marked asymmetry observed.

Fig.1. Stage I. Perameles obesula (1 Z).

a I. Dasyurus viverrinus (775 mm.).

3. ,, Il. D. viverrinus (8°56 mm.).

4, ,, Ill. BP: nasuta (18).

5. IV. P. nasuta (2 P).

Prate II.

Figures 6-8 represent three views of the wax model of the heart and gut of

Perameles obesula (10.viii.03) Stage V. The gut is painted white, heart

endothelium and arteries red, veins and most of. the capillaries blue, and

myocardium yellow.

These three figures were painted by Miss H. A. Steele.

Fig. 6. Ventral view of model of P. obesuda (10.viii.03).

7. Dorsal view.

8. Lateral view.

a rng: } van week

AROS ee:

ON SPIDERS FROM DUTCH NEW GUINEA. 501

36. On Spiders of the Family Salticids collected by the

British Ornithologists’ Union Expedition and the

Wollaston Expedition in Dutch New Guinea. By

El. R. Hoge, M.A., B.Z.S.

| Received and Read May 25, 1915. |

(Text-figures 1—11.)

INDEX. Page

Boethoportia ocellata, gen. et sp. n. ..............04......., 501

Dioleniws alboprceus, Spe Me sc. .ceies seis eeeaeeeeene se) OOM

LACH OOP QUE Ly FD>. Woso.o0.so nciscnocodoeavanesevenascessossoasccs | O05)

DE, GOGHEMIDUK TAT, SO> Ws soacnonnbsaeoocnes coo doodes jon sboacossancs Ole}

CO RUSHAG GUS S85 1016-39043. abd ad noise bbonap dopeanuee se aenae to bocmod sit 40)

JUG DUS JUCQNEOMMWOG oq sc00s0ecnconéuancosoadaseoongescanooseecnn DLL?

ZACUDODFIS CF CUPOMUCR, coe cic canosnooceedage eeaea pensesesenss ova ILB}

ZR CANE SDe Ms). eee tec Mer eEe Th. Vana eR ee ee eeeteoor aay OLE

LL ROU DOSING 1s one shee ddl dion eocisdae subes sagdepnce“eacdencaeeenee | lY/

L1 SYP UES SP-Glls eee eee sk ae eee OLD)

MO DSUSUNOLIMON tenth peas eet eR ose ee eee COL

Bathippus montrouzieri var. papuanus 0.20.00... 522

TETORTHPOMS UGE, SD. We “oagosegonoan opaas9 9nd euoa0e soacsadeaece GS}

COORG SOAIGI5, Ob Who Gab ddasqnoseqoodeaccssobeoorsonseseoseooes dao | BE

ONUGOM ALANS Dealers tasacvacce ok. os sare eee EAE OO

The spiders herein described complete the record of the

collections made by the above named Expeditions kindly entrusted

by the promoters to my care. The paper in which I described

the bulk of the collection appeared in Vol. XX. Part 14, of the

Transactions of this Society.

This interesting addition to our previous knowledge includes

some very beautiful forms of the exquisitely spangled varieties

which in their small bodies rival the coloration of the brilliant

bird fauna in which New Guinea is so rich. In spite of the

considerable amount of work devoted of late years by many able

contributors to collections brought from that country, it is evident

that much remains for future explorers before its numerous

species are exhausted. The proportion of new forms to those

already known among these is about three to one.

Family SALTICID 4.

Section Pleuridentati.

Group BorrHEs.

BoETHOPORTIA, gen. nov.

Boethoportia differs from Portia Karsch in having the front

row of eyes straight along the upper edges and the rear row as

02 MR. U. R. HOGG ON SPIDERS

(Sy

wide as the front row, and from Soethews in having the femur,

patella, and tibia of the first pair of legs stouter than the others,

fimbriations of long bristles at least on the tibia of the same, and

the metatarsi and tarsi of all legs very fine, the former at least

as lone as the tibie.

BoETHOPORTIA OCELLATA, sp. n. (Text-fig. 1.)

1 male and 1 female. (Z'ypes of the species.)

Female. The cephalothorax is bright chestnut-red, black round

the eyes with sparse white hairs; mandibles red-brown with pale

red fangs; lip and maxille red-brown, paler at the front edges,

with yellowish-grey fringes. Sternum also red-brown with

Text-figure 1.

Boethoportia ocellata, gen. et sp. n., 2.

a., epigyne; 6., profile; c., male palp; d., mandible showing teeth.

yellowish-grey hair. Coxe red-brown; femora brown under-

neath, with yellow patches on the upper side ; tibiz brown, with

a yellow band in the middle and long brown bristly fringes.

Metatarsus and tarsus yellow, with here and there patches of

white hair. The abdomen pale yellow at the base, the remainder

with large greyish-yellow spots on a brown gound ; the underside

is also brown with yellow spots, the hairs greyish-yellow on the

pale parts, brown on the darker portions.

The cephalic part of the cephalothorax is flat over the eye-space,

FROM DUTCH NEW GUINEA. 503

slightly sloping forwards, from the eyes it slopes steeply down

to the margin at the sides and rear. The thoracic part is barely

one-half the length of the cephalic, and the rear slope is deeper

than the whole length of the cephalothorax.

The front median eyes project forward on black rims. They

are three times the diameter of the laterals, which stand back

somewhat, but are level with the former along the upper margins.

The eyes of the rear row are of the same sizeas the front laterals.

The usually small eyes of the median row have a diameter two-

thirds the length of these; they are rather nearer to the front

lateral than to the rear eyes and are somewhat closer together

than either.

The abdomen is broadly ovate, straight and widest in front ;

the spinnerets are terminal, the superior having a conical second

joint. The epigyne is roughly triangular, rounded at the apex

and incurved in the middle of the base-line, with horizontal oval

apertures at the lower corners.

Male. Similarly coloured to the female.

The measurements (in millimetres) are as follows :—

Female.

Long. Broad.

( 22 in front.

Cephalothorax... 3d sae ts ROUND

2 | 33 4 high at rear.

INioclomven seen nae 6 4

Mandibles ...... 2

Trochanter Patella Metatarsus

Coxa. & femur. & tibia. & tarsus.

lees cea i 12 AL 5 da = 105%

De ] 4 43 42 = 14}

‘ 1 : me

3}, il Bt 33 4 = Iie

4, il 5 D4 7 = 18

Rall plenty. 4 Dab 2; a (ie

Mate.

Long Broad

Cephalothorax... 3 2i

Abdomen......... 4 12

Mandibles ...... 12

Pat. Metat.

Coxa. Tr. & fem. & tib. & tars.

We Osh ead. Ie 4 34 4 4 = 12}

OR 7 3 32 2 1]

Be 2 24 3 3} — 93

4 1 34 4 6 = 142

JE Or ee een aoe J ls 5 ls 4

504 MR. H. R. HOGG ON SPIDERS

Group DIoLENIES.

Genus Diotentus Thor.

Thorell, European Spiders, 1870, p. 203.

EK. Simon, Hist. Nat. des Ar, vol. 11. 1901, p. 480.

DIOLENIUS ALBOPICEUS, sp. n. (Text-fig. 2.)

1 female. (Type of the species.)

Female. The ground-colour of the cephalic part is black-brown

with white hair at the sides and round the eyes, greyer and finer

inside the eye-square. The outer margin of the thoracic part is

dark yellow-brown, but across the middle is a broad white-haired

transverse stripe.

Text-figure 2.

é.

Diolenius albopiceus, sp. n., 9.

a., Sternum, cox, and trochanters ; 6., eyes from in front; c., eyes from above;

d., epigyne; e., profile. ; :

There are also three unbroken wide transverse stripes across

the abdomen, one near the front, one in the middle, and one at

the posterior end.

There are seven pairs of spines on round roots on the under

side of tibia i., but no row of long hairs between them as in

D. amplectens, and five pairs of spines under metatarsus i.

FROM DUTCH NEW GUINEA. 505

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 22 \ 8 nn roam

Abdomen......... 33 2"

Mandibles ...... 3

Tr. & fem. Pat. Metat.

Coxa ~—— &tib. Stars.

Hees as.-a2 1. 1 P22 34 3 =, ule

2. + Ze 2 ge Os

3. 4 2 2 2) =,» G2

4. i 22 3 3 eS) Os

12 Oy Eee uit eee ries = il 2 3 => 23

Of the half-dozen somewhat similar species described by Thorell

from New Guinea and the neighbourhood, this is nearest to

D. amplectens Thor, (Ragni Aust.-Malesi, 1881, p. 412 et seq-).

It differs therefrom in having the front row of eyes more re-

curved, the line touching the upper edges of the median cutting

the side eyes at a point near the lower margins; the diameter

of the median is slightly more than twice that of the side eyes.

It differs from D. phrynoides Walck. in having the tibia of the

front pair of legs cylindrical instead of club-shaped, and without

any fimbriation on the under side.

D. fasciatus Thor., the coloration of which is somewhat similar,

has (according to that author) eight pairs of stout spines under

tibia i. and only two pais under metatarsus 1.

Section Unidentate.

Group CHRYSILLES.

Genus TreuamoniA Thor.

Thorell, Ann. Mus. Gen. ser. 2, vol. v. 1887, p. 385.

EK. Simon, Hist. Nat. des Ar. vol. 11. 1901, p. 552.

TELAMONIA VIDUA, sp. n. (Text-fig. 3.)

1 female. (Type of the species.)

Female. The cephalothorax 1s black, sparsely covered with white

squamous hairs and upright black and white bristles over the

eye-space, on the clypeus they are greyish-white. The mandibles

are black, paler about the falx-margin, the fangs dark red-brown.

Lip, maxille, and sternum black-brown with greyish hair. The

legs yellow-brown underneath, the cox quite bright yellow, but

the first three pairs of femora are dark yellow-brown on the

upper side. The abdomen is black above with upright grey hairs

and bristles, a white area of squamous hairs at the base, and two

506 MR, H. R. HOGG ON SPIDERS

similarly haired transverse bands broken in the middle, one

about the middle and the other near the posterior end. On the

under side, a black oblong area with a pale yellow-brown border

reaches to the sides.

The eye-space slopes forward and covers about two-fifths the

length of the cephalothorax. The latter is convex, high, and

slopes from the rear eyes to the posterior margin.

The rather large triangular tooth on the inner margin of the

falx-sheath, though not far from the base of the fang, terminates

the chitinous rim, the margin being cut away below it. The two

teeth on the outer margin are further down and stand at the

normal end of the same. The abdomen is ovate, rounded in

front, and tapers at the posterior end.

Text-figure 3.

Telamonia vidua, sp.n., °.

a., male?; b., mandible showing teeth; c., male palp; d., epigyne;

e., profile.

The rear row of eyes is not quite so broad as the front row,

and narrower than the cephalothorax at the point where it

crosses the latter. The front row is recurved; the side eyes

half the diameter of the median, standing a little away from the

latter. The lip is longer than broad; the maxille, upright,

rounded anteriorly, and thickly covered with bristly hair, are less

than twice as long as the lip. The sternum is ovate, truncate in

front, where it is twice as wide as the base of the lip.

Under metatarsus i. are two pairs of spines, one near the base

and another at the anterior end. There are two spines above

on femur iv., one small on the outer side of the patella, two

pairs on the tibia, two single ones on the outer side and a bunch

at the anterior end of the metatarsus, all of moderate size.

FROM DUTCH NEW GUINEA. 507

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 4 aa win MOSS

Abdomen ....... 5 3

Mandibles ...... 2

Pat. Metat.

Coxa. Tr.&fem. &tib. . & tars.

Legs 1 1 33 4 on ee Le

2. ik 3 34 7 10

on 1} 3 33 a 1 12

4, le 33 At a 133

Pallipiree aes: af = 2 4 iP = D+

Male. A somewhat broken specimen, almost similar in colour-

ing to the above, appears to be of the same species but, as in

other cases, the first pair of legs is the longest instead of the

fourth. The shape of the cephalothorax and positions of the

eyes are similar as also the falx-sheath teeth, but the lip is rather

broader, making it as broad as long. The coxee have thick white

hairs at the ends adjoining the trochanters.

The white pattern on the back of the abdomen is formed of

scales in two rows of rather large spots disposed longitudinally

instead of transversely, and the pale bordering on the under

side is less distinct than in the female. There is a row of

eleven spines on the anterior half of tibia i. underneath as in

T. trabifera Thor. and allied species.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 34 1 i Lot

Abdomen......... 32 12

Mandibles ..... 1

Pat. Metat.

Coxa, Tr.& fem. &tib. & tars.

hess tee IL. ile 33 34 oh | ae alll

2 1 3 24 24, = 83

3 i 25 24 a 83.

4 1 3 23 3 = 93

Balai tet swat cee ys = 11 it 1 = 32

In ‘ Ragni di Selebes,’ p. 251, Dr. T. Thoreil described a male

from Kandari which ne ane Telamona (Moevia C. L. Koch,

1848) latruncula. This is apparently very closely akin to the

above female, but differs in having a white marginal fillet round

the cephalothorax, and elsewhere being brightly coloured red,

blue and green, and having the lip 13 longer than broad. The

Proc. Zoo, Soc.—1915, No. XXXV. 35

508 MR. H. R. HOGG ON SPIDERS

legs are shorter, but in about the same proportion as in this

female.

Hither that or the above described male appears near enough

to belong to the female, but of course the pair above described is

known to be from the same neighbourhood.

TELAMONIA MANDIBULATA, sp. n. (Text-fig. 4.)

Il male. (Z'ype of the species.)

Male. Cephalothorax black-brown, with white squamules over

the eye-space interspersed with a few upstanding black bristles ;

the mandiblesare black-brown. The lip and maxille brown with

reddish fringes. The sternum dark brown with greyish-white

Text-figure 4.

Telamonia mandibulata, sp.n., ¢.

a., eyes; b., mandibles, lip, and maxille ; ¢., male palp.

hair. Of the coxe, which are all contiguous, the first pair are

brown, the remainder bright yellow. The abdomen is black-

brown on the upper side, with upstanding white bristly hairs and

a fillet of white squamules round the base and sides; inside this

are two rows of large spots of the same arranged longitudinally

from a little before the middle to the rear end; underneath it is

a dull yellow-grey. The front pair of legs is brown all over

FROM DUTCH NEW GUINEA. 509

with the exception of a pale yellow ring at the distal end of

the tibial joint. The other legs are yellower. They are all

sparsely furnished with white almond-shaped scales and up-

standing pale yellow-brown hairs. The palpi are as dark as the

first pair of legs.

The cephalothorax is about one-fifth longer than broad, flat on

the eye-space, highest at the level of the rear row of eyes, whence

it slopes slightly forward and steeply to the margin all round.

The eye-space, broader than long, extends to nearly one-half the

total length ; it is slightly narrower at the rear than at the

front. The eyes are all ringed with wide black margins. The

small eyes of the median row are about midway between the

rear and the front side eyes. The clypeus is one-third as wide as

the diameter of the front median. The front row is recurved,

the side eyes being slightly separated from the median which

- are close together,

The mandibles are perpendicular, convex on their outer side.

At a short distance from their insertion below the clypeus they

begin to widen out towards the anterior end, where they are

twice their width at the base. The total thickness at the anterior

end, in addition to the space between the inner and outer

margins of the falx-sheath, comprises a superimposed raised area

reaching from near the outer margin to the corner farthest away

from the base of the fang, where it ends in a large prominence.

Between this and the normal falx-sheath margin it is hollowed

out. On this secondary outer margin is a small fringe. The

usual falx-sheath is wide and deep, the inner margin terminating

in a large conical tooth at its lower end, from here to the base of

the falx the thickness is considerable.

The lip is longer than broad ; rounded anteriorly and narrowed

at the base for about one-fourth of its length, it is remarkably

convex, the middle being a considerable height above the level

of the margins. The maxille are upright, rounded anteriorly,

narrowed at the base, about twice as long as the lip, and as broad

as the latter is long. They are also very convex, being raised to

the middle of their area in successively smaller layers, and termi-

nating at their greatest convexity in a small boss.

The sternum is ovate, truncate anteriorly, where it is wider

than the lip; the front coxe are clearly longer than the others.

The abdomen is ovate, with rather long terminal spinnerets. ©

There are two spines above on femora i. and ii. One weak

spine on the inside of each patella, one pair at the base of tibia 1.,

two single on the outer side, and a row of eleven on the anterior

half of the inner side of the same. Four pairs under tibia u.,

one pair and an anterior bunch on metatarsus 11.

The tibial apophysis of the palp is broad at the base but tapers

to a fine point. The distal joint is short and broad near the

base, rounded at the sides and square at the anterior end, with

a short stigma springing from a hollow above the bulb. The

patella is as long as the tibia.

35*

510 MR. H. R. HOGG ON SPIDERS

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 4l | : an front.

Abdomen......... 44 23

Mandibles ...... 22

Pat. & Metat.

Coxa. Tr. &fem. tib. & tars.

Tags) 3.088 ll 13 a 4) 42 = 153

2. 1 34 34 Syn ky eet oe IL

3 1 33 33 AS —— oral

4, i 4 + 4i =) Ie2

AP aallioLagseeeta ste geste 4 2 13 el

The legs are of about the same proportion as those of 7’. scalaris

Thor. from Ternate. The species resembles 7’. trabifera Thor.

and some others in the row of 11 spines on the side of tibia 1.

(Thorell, Ragni Austro-Mal. 1881, pp. 477 & 480), but differs

from all in the extreme convexity of the lip and maxillz, and the

shape of the mandibles.

Group SAITEz.

Genus Jotus Koch.

Jotus L. Koch, Die Arach. Austr. 1881, p. 1243.

JOTUS IGNEUS, sp.n. (Text-fig. 5.)

1 male, 1 female. (Types of the species.)

Female. The cephalothorax is pale red-brown, moderately

thickly covered with recumbent yellowish-white hair and on

‘the eye-space with upright white bristles ; round the margin of

the front eyes are fillets of curly white bristles and much longer

ones of the same colour on the clypeus.

The mandibles are yellow-brown, darkest on the anterior

margins, with the fangs brown at the base and pale red at the

anterior half.

The lip and maxille are orange with dark yellow fringes, the

sternum and coxe paler yellow with yellowish-grey hair. The

legs are bright yellow, the front pair being rather the darkest.

The abdomen above is pale yellow, with smooth silky white

hair. Along each side a row of large brown spots forms a broken

line nearly continuous at the rear end: on the under side is a

wedge-shaped brown area extending from the genital fold to just

above the spinnerets. The front and sides are the same colour as

the upper side. The spinnerets and basal portion of the epigyne

are darker yellow.

The front row of eyes is recurved, slightly wider than the

rear row, which is narrower than the cephalothorax at that part,

and the small median are midway between the rear eyes and the

front laterals.

FROM DUTCH NEW GUINEA. 511

The single flat triangular tooth, about the middle of the inner

falx-sheath margin, is particularly large, and the anterior tooth

of the outer margin is also rather large.

There is a row of three short spines in front of each femur

above, with two single spines about the middle. On the under

side of tibia i. and ii. are three pairs of long powerful spines

and two smaller spines on the inner side; on metatarsus i. and ii.

underneath are two pairs of long spines ; on metatarsus iii. and iv.

are a pair near the base and a bunch at the anterior end, and

on tibia of same two above, two on the side, and a pair under-

neath. On the femoral joint of the palp are two spines, on the

patella one on each side, on the tibia one above, and on the distal

joint one below.

Text-figure 9.

Jotus igneus, sp.u., 3d.

a., female; 6., eyes and mandibles of female; c., mandible showing teeth of male;

d., epigyne of female; e., male palp; f, Plexippus paykulli, epigyne of

female.

Male. Similarly coloured to the female, the sides of the cephalo-

thorax being, however, darker brown and the brown stripes on

the abdomen continuous; the brown area on the under side also

begins quite at the base. On the front pair of legs the femora

are darker than on those of the others. The distal end of the

tibiz and metatarsi are also rather darker.

The male palp is of the same pattern as that of J. auripes

L. Koch, but the epiphysis is not serrated, nor are the bristly hairs

512 MR. H. R. HOGG ON SPIDERS

nearly so thick and long. The pattern of the cephalothorax

differs in having pale median and marginal stripes, with a

darker yellow-brown area between, instead of black, and the

median area on the upper side of the abdomen pale instead of

dark. Its larger size also distinguishes it from the former.

It will be seen from the measurements below that in the male

the first pair of legs is longest, while in the female it isthe fourth

pair. There is no appreciable difference in the length of the

tibia and patella ii. and iv. in the female, while in the male those

joints in the fourth pair are only very slightly longer.

The measurements (in millimetres) are as follows :—

Male. Female.

Long. Broad. Long. Broad.

Dachalothomes.. 42 a front. 42 { ae front.

Abdomen..... ... 6 24 5 24

Mandibles ... .. 24 lf

Male.

Pat: Metat.

Coxa. Tr.&fem. &tib. & tars.

Tie esr oe ee 1 13 4 5 Le

2 il 34 4 Sy eS US

3 1 4 34 SS = 12

4 1 4 32 le

Palisa eels can 4 2 13 eS ys

Female.

ers: 1 13 4 4 Bie ol

2 12 4 3 23 = 103

3 17 4 33 ss = 122

4 1} 4 33 eo = 1

LEY O10 Cane, So eereete ilaee 2 12 11 its = 22

Group PLEXIPPEs.

Genus Piexippus C. Koch.

Plexippus C. Koch, Ueb. Ar. Syst. v. 1850, p. 51. (Ad part.

P. ligo= P. paykullii Aud.)

PLEXIPPUS PAYKULLII Aud. (Text-fig. 5, f.)

Attus paykullai Aud. in Sav. Desc. de Egypte, 2° Edit. 1827,

vol. xxii. p. 172; lc. pl. vii. fig. 22, folio. 21826.

Salticus culicivorus OC. L. Doleschall, Tweede Bijd. Arach. van

den Ind. Archip. p. 14, pl. ix. fig. 5: Act. Soc. Sci. Indo-Néerl.

vol. v. 1858-9.

Menemerus culicivorus T. Thorell, Ragni Selebes, 1877, p. 228;

id., Ragni Amboina, 1878, p. 237 ; id., Ragni Austro-Mal. 1881,

p. 508.

FROM DUTCH NEW GUINEA. 513:

Menemerus paykulli Keyserling, Die Arach. Aust. 1883, p. 1425,

pl. exxiii. fig. 4.

Plexippus paykulli T. Thorell, Ragni Indo-Mal. 1891-2 =2, p. 369.

1 male and 3 females.

Collected from Spain and Africa; also, according to various

authorities, round the world eastwards and, according to F. O. P.

Cambridge, in 8. America.

It is impossible to compare Doleschall’s drawing of S. culi-

civorus with a recognised female specimen of P. paykulli without

feeling that it must have been drawn from the same, and Thorell

apparently arrived at this conclusion (Ragni Indo-Mal. 1892,

p. 370). In his description, which is quite short, Dr. Doleschall

says that the legs are in the order 43 2-1, while it is certain that

the second pair is the shortest.

The patella and tibia iii. are of the same length as patella and

tibia iv.,and the rear row of eyes in the female is slightly shorter

than the first row, while in the male it is quite clearly so.

Von Keyserling gives an excellent drawing of the male and

adds what he says is the epigyne of the female; the latter, how-

ever, is quite unlike those of specimens in the British Museum

(Natural History), which resemble the above. (See text-fig. 5, f.)

Group ZENODORES.

Genus ZENoporwus Peckham.

Ephippus T. Thorell, Ragni Aust.-Mal. iii. p. 643 (1881).

Zenodorus G. & E. Peckham, Proc. Nat. Hist. Soc. Wisc. vi.

p. 287 (1885); E. Simon, Hist. Nat. des Ar. vol. ii. 1901, p. 656.

ZENODORUS D’URVILLIL Walck.

Attus Wurvillii Walck. Hist. Nat. des Ins. Apt. 1. p. 459

(1837).

Ephippus @urvillet TY. Thorell, loc. cit. p. 653.

Zenodorus @urvilles G. & E. Peckham, loc. cit.; id., E. Simon,

loc. cit.

1 male.

This male seems clearly to belong to this species, agreeing

with Thorell’s elaborate description, and showing the first pair

of legs longest, though not so long in proportion as in some later

described species. The mandibular tooth on the inner margin is

conical and quite large if the soft basal portion is taken into’

consideration, and only “very minute” if you reckon the point

alone, which is darker and harder. The mandibles are hollowed

out a the middle of the inner side, much corrugated, and the

stout base of the short curved fang occupies the whole of the

anterior end of the falx.

A raised flat rim at the front of the clypeus seems a feature im

this genus Zenodorus; it is not quite so well defined in the female

as in the male.

514 MR. H. R. HOGG ON SPIDERS

The measurements (in millimetres) are as follows :—

Long. Broad.

Csttaloiions , é { - ni Ror

Nbdomeny anaaer 5 24

Mandibles ...... D

Pat. Metat.

Coxa. Tr. & fem. & tib. & tars.

hegs Lee Ie 1 4 2, 34 Ze ae

2. le 3 24 24 = 9

Be 1 4. Di ehetee 23,0 p=elle

4, Wa 3 3, 24 201th tle

JEG) nee rasennenanacaed a 2 1 13 a

ZENODORUS DANAE, sp. n. (Text-fig. 6.)

8 males and 10 females. (Including the types of the species.)

Female. Cephalothorax black-brown, with very brilliant green,

red, and yellow iridescent scales at the sides and rear of the

eye-space, with a few scattered here and there between the eyes

Text-figure 6.

Zenodorus danae, sp.u., 3.

a., female; 6., epigyne; ¢., male palp; d., mandible showing teeth; e., profile.

and on the clypeus. The mandibles, lip, and maxille are dark

red-brown with brown bristles and upright, flat, club-shaped,

pearly-white iridescent hairs. The sternum, though dark, is

more yellow-brown, with pale grey hair.

The legs and palpi are bright orange-yellow, with brown spines

FROM DUTCH NEW GUINEA. 515

and bristles. The two rear pairs of legs darken into yellow-brown

on the upper surface of the tibial and metatarsal joints. “he

abdomen is black-brown on the upper side; on the basal area,

along the sides of the rear half, and in a procurved fillet across

the middle, is a pattern formed of the green, red, and golden

scales,

The under side is pale yellow-brown without any pattern.

The eye-space is two-fifths of the total length of the cephalo-

thorax; the rear row is as broad as the front row, which is

recurved; the eephalothorax at the rear row of eyes, its broadest

part, considerably exceeds them in breadth. The clypeus is less

than one-half the diameter of the front median eyes and termi-

nates in a thick flat marginal rim. The mandibles are short,

stout, and conical, the fang very thick at the base; on the inner

margin of the falx-sheath the chitinous rim is continued only a

short way down as far as a conical tooth; this, though rather

small, is not “very minute,” and below it the side is hollowed

away. On the outer margin are two points on a single base and

a thick fringe of bristles. The lip is as broad as long, rounded

anteriorly, half the height of the maxille, which are upright,

convex, broad, and rounded on the outer side. The first pair of

coxe, slightly wider apart than the breadth of the lip, are longer

than any of the others.

The abdomen is oval, about twice as long as broad.

There are two pairs of stout spines on the under side of meta-

tarsus 1. and similar smaller ones under metatarsus 11.

Three pairs of spines under tibia i. with two single spines on

the inner side. One on each patella. On femora iil. and iv. there

are 1,1 spines above, and numerous spines on the tibie and

metatarsi.

On the inner side of the patella, tibia, and metatarsus i. there

is a thick fringe of bristles.

The front pair of legs are stouter than the others.

The vulva consists of two broad oval depressions side by side,

separated by a narrow ridge with two other shallower and

smaller fovese below the first pair, the whole on an elevated,

rather square area.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 5 | ‘ He fa

Abdomen......... 6 33

Mandibles ...... 2

Rate Metat.

Coxa. Tr. & fem. & tib. & tars.

J DiGi caueeeetoo ik iz 4 4 3 ele

2 1 33 3 25 =) 10

3 1 4 4 og = 125

4 1 33 3 a es 1G

IPapl assesses 4 12 13 = 5

516 : MR. H. R. HOGG ON SPIDERS

This species would seem to be rather close to Z. julia Thor.

(l. ec. p. 650), which it resembles apparently in the pattern of

brilliant scales on a black ground and the form of the epigyne,

but differs from it in the much more even lengths of the legs,

the third pair in Z. yulia exceeding the first and fourth by 23 and

33 millimetres respectively, and being also darker in colouring.

It is much paler in colour, larger, and the legs i. and iii.

are nearer the same length, than in Mr. Pocock’s Z. variatus

(Willey, New Britain, etc., vol. i. 1899, p. 117). It differs also

from Z. @urvillec (Walck.) in the much brighter colouring and

absence of rings on the legs, while the pattern of the epigyne

differs from that drawn by Von Keyserling (Die Arach. Aust.

pl. exx. fig. 4d).

Males. In most respects these agree so closely with the

females described above and are represented by so nearly the

same number of specimens in the present collection, that it is

difficult to avoid the conclusion that they are males of the same

species. The chief difference is that the front pair of legs are

much longer than the others, a fact which does away with one of

the characteristics of the genus, in which the third pair of legs

are said to be much longer than the others. Were it not, how-

ever, for the special enlargement of the first pair, the third would

be the longest.

The clypeus is not quite so broad as the diameter of the front

median eyes, the rear row is as broad as the front row. The

eye-space slopes forward, and from the hinder row the thoracic

part slopes steeply to the rear margin.

The mandibles are flatter than in the female, both the outer

and inner sides being strongly corrugated transversely. The

tooth on the upper margin is of moderate size ; the inner margin

of the falx-sheath is cut away to about one-third of the length of

the outer margin and exists only for a short distance near the

base. The lip is as broad as long, rounded in front, hollowed

out on either side of the base, and transversely corrugated.

The sternum is three-fourths as wide as it is long, truncated in

front, and the coxe are as far apart as the greatest width of

the lip. The male palp has a flagellum in about three spirals at

the anterior end of a plain oval bulb.

The front pair of legs is strongly fimbriated on the under side

of the patellar, tibial, and metatarsal joints. The cephalothorax

and abdomen are black-brown, with a pattern of opalescent pearly

scales.

Under metatarsus i. are two pairs of stout spines, but none at

the side. Three pairs under tibia i. One spine each on patelle

il, and iv., and a bunch at the anterior end of metatarsi iii.

and iy.

FROM DUTCH NEW GUINEA. 517

The measurements (in millimetres) are as follows :—

Long. Broad.

Gils "

Cephalothorax... 5 | 2 TL AO

Abdomen......... 6 34

Mandibles ...... 3

Pat. Metat

Coxa. Tr. &fem. & tib. & tars.

Wess? 00m 1 a A PE, Sey eS ey 2

2 12 4A 4 3d == slay

3 le 5 ay Pies So = Ibs

4 13 4 4 4. = 152

IEE Ol Poe NaRne 3 2 ls 13 = dz

ZENODORUS RHODOPE, sp. n. (Text-fig. 7.)

2 males and 2 females. (Including the types of the species.)

Males with one tooth on inner margin of falx-sheath.

Females with two teeth on inner margin of falx-sheath.

Female. The cephalothorax is dark yellow-brown on the

Text-figure 7.

Zenodorus rhodope, sp. u., 6.

a., female; 6., mandible showing teeth; c., epigyne; d., male palp.

cephalic part; all the eyes on large black tubercles. Small

white lancet-shaped scales and upright brown bristles spread at

518 : MR. H. R. HOGG ON SPIDERS

intervals about the eye-space. The thoracic part is nearly black-

brown to the sides of the posterior margin. On the clypeus are

long white upstanding bristles.

The mandibles are dark red-brown, with a paler patch at the

lower inner margin of the falx. The fangs red-brown. The lip

and maxille dark brown, yellow-brown at the margin with

yellowish-brown fringes. The sternum is yellow-brown. The

coxe bright yellow. Legs and palpi as described for Z. syrinx.

The abdomen on the upper side is dark greyish-brown, covered

round the sides and base thickly, in the central area more

sparsely, with pearly-white squamules and upstanding brown

normal hairs. On the under side it is much more thickly covered

with short, upstanding, white bristly hair; the spinnerets are

the same; the epigyne dark yellow.

Arrangement of spines on the legs ;—

I. Femur: 1 1 1 above and 111 small at anterior end.

Patella: 1 on inner side and long bristle.

Tibia: 3 pairs underneath.

Metatarsus : 2 pairs very long and stout on under side.

IJ. Femur: 1 1 above and 111 at anterior end on inner side,

1 1-on outer.

Patella: 1 and bristle.

Tibia.: 1 at side. 2 fine pairs underneath.

Metatarsus : 2 pairs much more powerful.

Tit. Kemur: 14 and 11 1.

Patella: 1.

Tibia : 1 on inner side and pair at anterior end.

Metatarsus: 2 pairs and bunch at anterior end.

IV. Femur: 1 on outer side, 1 in middle, 1 at anterior end.

Patella: 1 on inner side and bristle at anterior end.

Tibia: 1 1 on inner side and pair at anterior end.

Metatarsus: 1 1 on inner side and thick bunch at anterior

end.

Palp: 11 on femur above.

The measurements (in millimetres) are as follows :—

Long. Broad.

Hoa

Cephalothorax... 4k { 7 a Ee.

Abdomen......... +) 3

Mandibles ...... 2

Pat. Metat.

Coxa. Tr. & fem. & tib. & tars.

JOYS tscer se ] 13 4 34 Sale

2 iz 3 3 7 93

3 11 5) 4 43 = 143

4 11 4 3 4 = 121

Pal pias. ccs 1 Ls 1+ it = 5

FROM DUTCH NEW GUINEA. 519

The females are rather more highly coloured than the males

and the legs more distinctly ringed, but so closely resemble

them in structure and general coloration that I have supposed

them to be of the same species, though unless captured together

the matter is always open to doubt.

In the males the tooth on the inner falx-sheath is raised above

the lower end of a straight base, which in the females has a

point at the other end also, enough to make it really fissidentated ;

but these two and the following Z. syrinzx with no teeth at all

are all so much alike, and so distinctly resemble Zenodorus

@urvillei, the type species, that I am unable to separate them

from that genus.

ZENODORUS SYRINX, sp. n. (Text-fig. 8.)

1 male and 1 female. (Types of the species.)

Male without teeth on falx-sheath.

Female without teeth on inner margin of falx-sheath.

Male. Cephalothorax black-brown on the eye-space, bright

red-brown just behind the same, behind this again dark brown

to the rear margin ; scattered over this at intervals are lanceo-

late pearly scales, more thickly at the sides and on the clypeus,

where some of them are lengthened into flat bristles. On the

front margin of the clypeus is a flat raised rim. The mandibles

are black-brown with transverse corrugations of green opalescence,

and, except for a few flat bristles at the inner edge of the base,

quite bare.

The base of the falx is as thick through as it is broad trans-

versely, but tapers towards the anterior end. A transverse

section at the base would be almost square. The median part of

the inner margin is hollowed out, thus leaving an oval opening

between the two falces.

In the specimen before me there is no tooth on the lower

margin of the falx-sheath, and none visible in the thick fringe

on the outer.

The lip and maxillze are nearly black-brown, with a narrow

yellow-brown edging and dark grey fringes. The sternum and

coxe are dark orange-yellow, with fine upstanding yellow-brown

hair.

The femur and patella of the front pair of legs are dark

orange ; the tibiaand metatarsus black-brown with a thick black

fringe of long bristles on the under side, those on the metatarsus

being the shorter. ‘The tarsi quite pale yellow. The metatarsus

of the second pair is pale, otherwise the same colour as the first ;

the third and fourth pairs are paler yellow-brown.

The abdomen on the upper side is dark greyish-brown, with

short upstanding brown hair over the basal area ; following this

is a broad field of pearly-grey scales, and along the sides patches

of the same as far as the spinnerets; the under side is pale

yellowish-grey.

520 MR. H. R. HOGG ON SPIDERS

Text-figure 8,

Zenodorus syrinx, sp.n., 3d.

a., female; 6., epigyne; c., male palp; d., mandible.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 5 { 1 oat,

Abdomen......... 2 24

Mandibles ...... 2

Pat. Metat.

Coxa. Tr. & fem. & tib. & tars.

esse eens, Me 13 43 5 ee — yeh

2. 1 3 3 2 93

on iW 54 44 5 = 16

4., 1 4.

Female. Cephalothorax dark yellow-brown; all the eyes on

large black tubercles. Long brown bristles and short white

FROM DUTCH NEW GUINEA. 521

lancet-shaped scales spread here and there. The mandibles

black-brown with red-brown fangs and brown fringe. A paler

patch just below the lower corner of the falx-sheath. The lip

and maxille are dark yellow-brown with paler edgings. The

sternum pale yellow-brown, with upstanding brown bristles and

pale yellow-grey hair. The coxe bright yellow. The whole of

the palp the same, with upstanding yellowish-grey bristles. The

basal half of the femora of all the legs is bright yellow-brown,

the anterior half brown; patella and tibia yellow-brown, meta-

tarsus and tarsus orange-yellow.

The abdomen is black, with a fillet of white squamous hairs

round the base reaching along each side to half the length of the

abdomen. From the end of this on each side are three large

white spots continuing the line to the spinnerets ; on the under

side it is plain greyish-yellow; the hairs all ordinary, very fine,

pale yellow. ‘The spinnerets rather long, conical, and darker

brown, on a white chitinous base half their length, have long

straight brown hair and a short cylindrical second joint.

The inner margin of the falx-sheath is cut away and hollowed

out to the outer margin, at the lower end of which are two

moderate-sized teeth. The inner side of the falx-sheath itself is

also hollowed out, most deeply about the middle.

The lip is as broad as long, truncate and slightly hollowed

anteriorly, half the length of the maxille, which are convex,

upright, rounded anteriorly. The front coxe are farther apart

than the lower margin of the lip is wide. The sternum ovate,

truncate, and narrowest in front, is flat at the anterior end for

one-third of its length and thence to the posterior end convex.

The epigyne consists of two oval hollows resting against the

upper part of a broad longitudinal convex septum; below these

hollows and fitting into the lower part of the same central

septum are two chitinous triangular cushions. The whole inside

an oval frame.

Arrangement of spines on the legs :—

On femur i. & ii. above are single spines 1 1, and a row of

three small on the inner side at the anterior end.

On patella i. & ii. a short spine on the inner side and a very

long bristle at the anterior end.

On tibia i. & 11. three pairs on the under side.

On metatarsus i. & 11. two pairs (very stout) on the under

side.

On patella iii. & iv. one small on inner side and a very long

bristle at distal end.

On tibia iii. & iv. one small pair below at the anterior end,

and two single on inner side.

On metatarsus ii. & iv. one small underneath about the

middle, and bunch at anterior end.

On metatarsus iii. two single at the side, but none on iv.

522 MR. H. R. HOGG ON SPIDERS

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 3 | : ai ae

Albdomens.... 22 52 33

Mandibles ...... 2

Pat. Metat.

Coxa. Tr. & fem. & tib. & tars.

LOGS i.. neineees Ik lg 34 33 22 ile== «oor

2. 1 24 24 Ob risen) Dg

3) 1 42 33 Sy Tee ia

4, 1 3+ 3 a5 = 102

12E:]| CMS ene eenos as 3 13 13 ae)

This species differs from Z. rhodope in having no teeth on

the falx-sheath margin, in not having the legs ringed, slight

differences in the male palp and epigyne of the female, and in

the spines as detailed, but otherwise the two species closely

resemble one another.

Group THYENES.

Genus Mopsus Karsch.

Mopsus Karsch, Mittheil. Miinchener Entom. Vereins, vol. 11.

1878, p. 31.

Moprsus mormon Karsch.

Mopsus mormon Karsch, loc. cit.; 'T. Thorell, Ragni Austro-

Malesi, p. 462 (1881).

Ascyltus penicillatus Keyserling, Die Arach. Aust. p. 1319,

pl. exii. (1882).

Mopsus mormon Keyserling, loc. cit. p. 1475 (1883).

1 male and 2 females.

Previously recorded from New Guinea, Cape York, Bowen,

Rockhampton, and Sydney.

Group PLEXIPPES.

Genus Baruiprprus Thor.

Bathippus Thorell, Ragni Indo-Malesi, pt. iv. vol. ii., 1891-2,

p. 401; E. Simon, Hist. Nat. des Ar. vol. ii. 1903, p. 740.

BATHIPPUS MONTROUZIERI, var. PAPUANUS Thor.

Plexippus montrouziert Lucas, Revue et Mag. de Zool. 1869,

p. 209, pl. xi. figs. 8-12; Thorell, Ragni Austro-Malesi, i1i. 1881,

p. 526.

6 males.

Previously recorded from Wokan, Aru Islands, and Fly River,

New Guinea.

FROM DUTCH NEW GUINEA. ByAs}

Section Fissidentati.

Group HaAsAripZ&.

Genus Hasarius Simon.

Hasarius KE. Simon, Hist. Nat. des Ar. vol. ii. 1903, p. 795.

Hasarius GLaucus, sp.n. (Text-fig. 9.)

1 female. (Zype of the species.)

Female. The cephalothorax is orange-yellow, black hetween the

side eyes; mandibles, lip, maxille, sternum, and legs all pale

yellow, with nearly white hairs and grey spines.

The abdomen above is dark grey at the sides with short recum-

bent white hairs, a broad pale area at the base, and a pale yellow-

erey scolloped longitudinal stripe down the middle. On the

under side the median area is the darker, the sides being pale

yellow-grey. The spinnerets are yellow, springing from a long

white chitinous base nearly as long as themselves. The epigyne

is rather dark yellow.

Text-figure 9,

ay WWE yy \\

& has’

Hasarius glaucus, sp.n., 9.

a., front eyes and mandibles ; b., mandibular teeth and fang; c., lip, maxille,

sternum, and coxe; d., epigyne.

The eye-space is spread over two-thirds of the cephalothorax.

The lip is longer than broad, Sternum ovate, truncate, and

narrowest in front; the third coxa is isolated from the second

and fourth.

On the under side of tibia 1. are two pairs of long spines, one

Proc. Zoou. Soc.—1915, No. XXXVI. 36

a24 MR. H. R. HOGG ON SPIDERS

pair of short ones at the anterior end, and two single spines on

the outer side; underneath the metatarsus there are three pairs

(two very long g) and a bunch at the anterior end.

Two single spines above and two pairs on metatarsus li. and

ive8 nenwlere tibia ili. and iv., a pair of fine spines in the middle

and another at the anterior end.

The measurements (in millimetres) are as follows :—

Long. Broad.

5} an

Cephalothorax... 24 | a in front.

IN COITNEIN 4550000 3 )

Mandibles ...... ]

Pat. & Metat.

Coxa. Tr. & fem. tib. & tars.

COS Beene: 1. $ 2 22 oe a On,

Saeed 2 2 Le BS

3. 3 24 ZA eer aes

4, L 2 2 2) — moe

Pali cerevisiae 4 IZ 1 ees

Differs from H. coprea Thor. (Rag. Indo-Mal. pt. iv. vol. 11.

1892, p. 434), which it somewhat resembles in having a longitu-

dinal pale median stripe on the abdomen instead of transverse

black streaks, and the hind pair of legs clearly longer than the

first instead of equal.

Group CytHE&.

Genus Cyrma Keys.

Cytcea Keyserling, Die Arach. Aust. 1882, p. 1380 ; E. Simon,

Hist. Nat. des Ar. vol. 11. p. 816.

CYT#A SYLVIA, sp.n. (Text-fig. 10.)

1 female. (Type of the species.)

Female. Cephalothorax dark red-brown, black between the side

eyes. White and coloured scales on the cephalic part. On a

lighter red ground behind the eye-space is a large patch of white

squamules reaching to the rear margin; on the clypeus is a

bush of long white bristles; the mandibles are dark brown on the

inner margin, pale chestnut-red on the outer sides and brown

underneath. The fangs red, paler at the points. The lip and

maxille are brown on the lower and middle parts, yellow over a

rather wide area in front. The sternum and coxe are pale

yellow with white upstanding hair. The two front pairs of legs

have the femora pale red-brown on the upper side with a dark

brown patch at the anterior end, and dark brown on the under

side, thickly covered with white ‘squamules on the pale portions,

and with coloured squamules on the darker parts. The patella

and tibia are pale and dark brown in alternate rings; the

FROM DUTCH NEW GUINEA. 525

metatarsus and tarsus pale yellow-brown ; the two posterior pairs

of legs are somewhat paler, but with similar brown rings and

squamules intermixed with upstanding brown bristles. The palpi

are yellow-brown, with thick brushes of long white bristles.

The ground-colour of the abdomen is pale yellow, covered with

thick masses of white, black, and red lancet-shaped scales all

mixed up together.

The eye-space and thoracic part each occupies about one-half of

the total length of the cephalothorax, but, the upper flat part

continuing for about halfway on the thoracic, the rear slope is

quite steep at the end.

The front row of eyes is straight along the upper edges, the

rear eyes about the same diameter as the front lateral. The

second row of eyes halfway between these two are quite small ;

the clypeus about one-fourth the diameter of the front median

eyes.

Text-figure 10.

Cyt@a sylvia, sp. n., 2.

a., lip and maxille ; 6., mandibular teeth and fang; c., epigyne.

The mandibles are convex on the outer side. The fissidental

tooth is rather high and its own length distant from the base of

the fang, its lower corner forms the larger of two conical teeth ;

on the outer margin are four teeth, the two upper being largest

and the lower quite small.

The lip is clearly longer than broad, narrowed and rounded in

front, and more than half the length of the maxille. The front

cox are barely separated by the distance of the breadth of the

base of the lip, so that the sternum is narrower in front than at

the posterior end; it is ovate, convex over the posterior two-

thirds, with a broad flat margin between it and the coxe. The

abdomen is oval, but is partially destroyed. The epigyne consists

of a pair of deep oval depressions upright, side by side, separated

by a black chitinous ridge widest in the middle, which runs

36*

526 MR. H. R. HOGG ON SPIDERS

round the upper and lower edges and inner side; the upper part

of each of these depressions is covered over with a light sort of

deck, dark streaks run down from their lower ends to the genital

fold, and the whole stands on a raised area arched anteriorly.

In the middle of the upper side of each femur is a single spine

and a row of four abreast at the anterior end; on each patella

one spine each side; under all the metatarsi are three pairs of

spines, on the inner side two single and on the upper side a pair

near the base followed by a single. Under metatarsus i. and 11.

two pairs of very long spines, and two single on the inner side; on

metatarsus iii. two bunches of spines; on metatarsus iv. one

pair underneath, one above, two at the side, and a bunch at the

anterior end.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 4 | ; aL EROS

Nndonrent scenes: 24

Mandibles ...... ji

Pat. & Metat.

Coxa. Tr. & fem. tib. & tars.

Megs hen ae. Jk ie 34 4 Sees aie

2. 1 Da 23 20 TS)

oe 1 22 3 2. = 9

4. 1} 24 24 Son = AOE

EP alloy ease hae > Ie Le Ms eet et

In the structure of the mouth-parts, proportion of legs, and

pattern of epigyne, this rather closely resembles C. alburna Keys.

It is at least a third larger, and is easily distinguished by the

prominent white streak on the cephalothorax and the bright

coloration of the scales on the abdomen.

CyT#A LAODAMIA, sp.n. (Text-fig. 11.)

1 male and 2 females. (Jncluding the types of the species.)

Female. Cephalothorax black-brown with greyish-white squa-

mules and brown bristles, reddish bristles between the front

eyes, and thick long white bristles on the clypeus. Mandibles

black-brown with red-brown fangs. Lip, maxille, sternum,

and coxe dark brown, the fringes on the former brown and

short upstanding greyish-white hair on the remainder. The

abdomen on the upper side is brown over the median area, with

a white fillet of squamous hairs at the base and down each side as

far as the spinnerets; the brown area has also white squamules in

patches mingled with brown, and a more or less distinct median

line of the same running down the anterior half; on the under

FROM DUTCH NEW GUINEA. 527

side it is brown at the sides and yellow-grey in the central area

with the same pale squamules. The femora, patella, and tibiz

of the legs are brown thickly covered with greyish squamules: the

metatarsi are dark brown at the base and in front, with a yellow

ring in the middle, and the tarsi yellow. The palpi are likewise

brown, with white squamules on the upper surface, and long

white bristly fringes on the sides of the patellar, tibial, and

distal joints, smooth underneath.

The epigyne is greenish-grey in the central area, pale brown

round the sides.

The spinnerets are pale brown, and spring from a conjoined

chitinous base as high as one-half of their length. The bifid

tooth on the inner falx-sheath stands out high, hollowed in the

centre, the two ends form large flat conical teeth. There are

three medium-sized teeth on the outer margin. The lip is

convex, rounded anteriorly, but is hollowed out from nearly

Text-figure 11.

Cytea laodamia, sp. n., °.

a., lip and maxille ; 6., mandibular teeth and fang of male; c., epigyne;

d., male palp; e., inner mandibular teeth of female.

halfway down to the lower corners; this part being continuous

with the rest, leaves it as wide at the base as it is high—it is

rather more than half the length of the maxille. The sternum

is oval, and while it narrows to the width of the convex portion

of the lip, the front coxe are as far apart as the full distance

between the next pair.

Male. Similarly coloured to the female, but the upper side of

the patellar and tibial joints are paler red-brown, and the

fringe on the under side of the palpal distal joint darker brown.

928 ON SPIDERS FROM DUTCH NEW GUINEA.

The bifid mandibular tooth on the inner margin is longer than in

the previously described species, less distinctly shaped at the

corners, and slightly serrated in the middle.

The measurements (in millimetres) are as follows :—

Female. Male.

Long. Broad. Long.. Broad.

Hens eae, : ;

Geplalottors a 23 in front. ; 3 in front.

3 33 34

Abdomen......... 41 24 43 24

Mandibles ...... 13 2

Female.

Pat. Metat.

Coxa. Tr. & fem. & tib. & tars.

JOC aeaP ee Jk 1g 3 3h ole

2 i 3 24 2 oo

3 i 3 3 21) == ou

4 13 3 3 2

Pall piv eeere censors 3 13 1} a 4

Male. _

Gees je cAe.n: il 12 4 5 es Tels

2. 13 3 34 24 = 10}

3. 1} 3 3 2 = OE

4, 13 3 3. 22 ee oe!

Ballipig ink aaccuetes 3 2 13 We es 9 OF

This, although a good deal larger than either von Keyserling’s

or Prof. Kulezynski’s species, agrees in. shape and the arrange-

ment of the spines with the definition of the genus as amplified

by M. Simon.

It will be seen that pat. + tib. 11. and iv. are of equal

length, in which it differs from C. alburna Keys. as well as

in the pattern of the epigyne and general coloration.

The form of the epigyne is the same as that given by

Prof. Kulezynski for his C. swbsiliens, and, as in that species, the

eye-area, although considerably broader than long, is still longer

than the pars thoracica. It differs from the latter in the third

and fourth pairs of legs being shorter than the first, instead of

longer. ie:

THE HOUSE-FLY EXHIBITION, 529

EXHIBITIONS AND NOTICES.

May 11, 1915.

Dr. A. SmrirH 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 April 1915 :—

The number of registered additions to the Society’s Menagerie

during the month of April was 230. Of these 150 were acquired

by presentation, 25 by purchase, 38 were received on deposit, 6 im

exchange, and 11 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 115.

Amongst the additions special attention may be directed to :—

1 Goeldi’s Marmoset (Callimico goeldii), from Bunda River,

Bolivia, new to the Collection, deposited on April 14th.

1 Houbara Bustard (Houbara undulata), from North Africa,

received on deposit on April 27th.

A valuable Collection of Waterfowl containing sixty-eight indi-

viduals representing twenty-five species, presented by Alexander

L. Duncan, F.Z.8., on April 23rd.

The House-Fly Exhibition.

Prof. H. Maxwe.it Lerroy, M.A., F.Z.S., Curator of Insects,

exhibited specimens of various kinds of fly-traps and made the

following remarks upon the House-Fly Exhibition :—

“ A small exhibition of Flies and methods of dealing with them

will be open on Wednesday, May 12th, and continue open while

it is of use and interest. There will not necessarily be anything

original in the exhibition, and it was planned at first simply

to show the American fly-traps, which are not used in this

country and which are likely to be valuable. From this it has

grown to include all the information about flies that a health-

officer might require, and we are endeavouring to arrange it so

that such an officer can obtain in a short time the technical infor-

mation about flies that is available, without himself having to

hunt up the literature. We have also tried to make it useful by

getting samples and prices of the necessary chemicals ete., by

having copies of books and other literature, and by condensing

into a short illustrated pamphlet authoritative information.

Since this was planned the Press have taken it up, somewhat

to our embarrassment, and we have had to make it as interesting

and instructive as we can for the general public: so we are

enlarging the scope, providing tabloid information and posters,

and generally trying to rub in the essential facts.

Flies are likely to be important this year for several reasons

the first is that with a shortage of labour scavenging will not be

530° PROF. H. MAXWELL LEFROY ON

so well done, manure cannot be so quickly disposed of, and there

will be local outbreaks of flies if the weather STL S are

suitable.

The usual vigilance exercised in inspection, and in otto

material capable of breeding flies will be impossible this year ;

there will be accumulations of manure and refuse in towns, on

farms, on market gardens and where cavalry are quartered, and

this will provide bre eeding-material particularly for house-flies.

The second reason is that j in areas affected by the war, flies are

increasing, because sanitation and the disposal of fly-breeding

material will not be possible, and this will accumulate to an

unusual extent. I have evidence of this already from the Con-

tinent: these flies will not come over here, but they will lead to a

greater spread there of the diseases that they carry and we shall

be affected; if cholera breaks out in Serbia, Austria, or other

parts of South Europe, it may spread in fly-invested areas and

get widely diffused. I do not want to bean alarmist, but I think

we must be prepared for such possibilities when a war of this

magnitude is in progress.

If flies are going to be unusually numerous so will the diseases

they carry, and one of these is typhoid: we inoculate the men

who go to the front, but we here are not inoculated.

The third consideration is that there is a likelihood of a very

terrible plague of flies actually where the fighting is in progress,

which will very deeply impress those who are There no one quite

knows where the fighting-line will be, but wherever it is the flies

are likely to be a feature. This isa good time, then, to do what

we can to rub the facts in, to get them known, to arouse interest,

and to help those whose business it will be to fight flies: already

we have been visited by Army Doctors anxious to get quickly all

the facts about flies, and this will become incr reasingly useful.

For these reasons we have made all we can of this little

exhibition, and, though it is not nearly complete, it contains useful

information and will be open at once.

It is not necessary for me to go into the details of the life-

history and habits of flies to-night: you can see it all in the

exhibition to-morrow, and you are probably all aware of the main

facts. We are showing the various stages of the House-fly and

the Blow-fly, the common flies of houses, where they breed, what

they feed on, the diseases they carry, how they carry them, and

soon. The facts are condensed on diagrams, posters, pictures,

and the methods of fighting flies are described in a short

pamphlet.

What can one do against flies? There are three lines on which

one can work, adjusting one’s methods to local circumstances of

course :—

. Removal or treatment of breeding-material.

Traps.

Poisons.

i) i) VS

The materials in which house-flies lay eggs and develop are

THE HOUSE-FLY EXHIBITION. 531

well known; they are horse-manure, excrement, rotting vegetable-

matter such as vegetables, compost heaps, rubbish heaps, and the

like: manure, excrement, garbage, and fermenting rubbish are the

danger sources for house-flies. For blow-flies animal-matter is

the chief source, even in extremely small amounts: a dead mouse,

a hollow bone, a putrifying whelk, a dead sparrow, a scrap of

meat, a fish-head, these will all nourish blow-fly maggots: you

will be astonished when you see what a small amount is needed

and what a number of blow-flies will develop from the scraps in

the dust-bin.

The number of flies this year will depend very largely on the

extent to which this material is removed or destroyed, not only

by the authorities but by the efforts of every one concerned.

Until people at large realize what a danger flies are and what

it is that they breed in, we are certain to have quantities of

flies; it is an unpleasant subject that people prefer to leave

alone, but there may be enough flies this year to make people

want to know about them.

If the breeding-material cannot be disposed of, can it be

treated so that it will not breed flies? Can we treat manure

without impairing its value, and can we show how to treat the

manure-heap in every garden? We have the available infor-

mation collected, and the original papers can be consulted ; so far

as is possible inquirers will be advised.

The second method is to trap, and I show to-night two:-American

traps, with Mr. Seth-Smith’s improvement, a folding- trap made

in the Gardens, and two other patterns of folding-trap that are

being tested. These and any improved ones will be shown, and

if any are available the prices etc. stated. Some patterns are for

home manufacture, some will, we hope, be available at popular

prices.

We show also a trap that is meant for the household dust-bin,

which will catch any flies going to the dust-bin and any that have

developed in it and that seek to escape. If a dust-bin of this

kind is used to contain stable-manure it will probably be a very

valuable trap for house-flies generally, and the stable become a

means of generally reducing flies in the neighbourhood instead of

adding to them.

The third method is to poison flies, and this has yielded

remarkable results abroad. There are harmless liquids for use

indoors and poisons for use by skilled persons outside, particularly

for hospitals and large institutions. These will be of greater

interest to professional men than to the public generally :

have the original accounts of these, and it will be possible for a

health, officer to read quickly what has been done.

It may perhaps sound to you as if we knew all about flies, and

as if we should be able to solve all the difticulties that will be

brought to us—I wish, indeed, it were so. It is astonishing how

little | practical infor mation there i is; what there is, is largely from

America—very little has been done here, and we are very far

indeed from having the information we need.

532 _ PROF. H. MAXWELL LEFROY ON

We have here to-night representatives of the Press, the

educators of the man in the street, and the exponents of his

collective opinion. I hope Fellows of the Society will forgive me

if I take the opportunity to urge the Press to emphasise to the

Great British Publie this fact, that we are actually at a loss to

know how to meet the fly problem here and at the front because the

necessary scientific investigation has not been done. As a nation we

neglect and underrate the value of applied science; it is not the

function of the Zoological Society to deal with the application of

Science to national health, and it is the wide sympathies of its

Council and officers that enables me to make use of its resources

to deal with this problem: but there is no organised body, no

department of Government that stimulates and develops the

study of Apphed Biology. If there had been, we could answer

the questions that come in to us daily from municipal health

authorities, Army Sanitary officials, and the public, as to means

of dealing with flies and other vermin.

I am not talking in a general way, but I have definite specific

problems that are in need of solution at once.

1. Baits for Flies—The recommended baits for fly-traps are

formalin, beer, vinegar, milk, alcohol, and a few others: these are

things arrived at by chance. It seems likely that we might find

a really good bait, something that would draw in every fly for a

quarter of a mile round, if we worked systematically on the line

of testing substances likely to be found in the breeding-materials

that flies seek. What brings the fly to the manure heap? Smell

possibly; if so, can we find a substance of not too intolerable a

smell that could be used to bait a trap in a corner of the garden

and that would really bring in all the flies. I think we can, but

it will require investigation, and it is a bad time to start inves-

tigating when the problem is imminent: we have actually begun

with the assistance of the Organic Chemistry Department of the

Imperial College, and we hope to get something out, but it is

work that should have been all done long ago.

2. Beside baits based on the fly’s breeding-place, can we get a

bait based on his food? Why do flies sometimes take formic

aldehyde? Is this the best or the only one, or are there other

definite substances of far greater attractiveness to flies which

they associate with their food ¢@ It is worth trying, and obviously

if we can get a really potent attractor it will help immensely.

3. Baits based oi Sex.—How does the male find the female ?

By scent, by sight, by what ?

I remember in India a planter sent me some fruit-flies with

the remark that when he put a particular mosquito essence on

his handkerchief these flies followed him continually ; the essence

contained citronella oil, and a drop of this oil will bring the males

of these flies from all around: investigation showed that the

female produces this oil, and it is by its scent that the males find

them : we have now a method of dealing with this fly.

I quote this quite true story as an example; and it might be

THE HOUSE=FLY EXHIBITION, 533

worth while studying the house-fly and the blow-fly from this

point of view.

4. Questions are coming in about treatment of manure-heaps to

keep flies away: is there anything deterrent to flies? It happens

that we have some recent research work on this very point, but

only as regards blow-flies ; it is work done at the Cooper Research

Laboratory at Watford. There is here an important line of work

and one which is going to be of immediate importance.

5. Treatment of Manure-heaps to kill Maggots in them.—Some

work has been done on this point in the United States and

Canada, but I believe that much better methods only need to

be worked out, possibly using the new vapour-poisons that have

been found during the last year. Im America they recommend

borax, but it seems likely that better and cheaper methods will

be found.

I have now shown you investigations which, if done in the

past, would have enabled us to deal with this fly problem ; so it

is, of course, with all branches of science, but I have been able

here to give you definite concrete cases and not simply glittering

generalities.

I have devoted more time to this point perhaps than I ought

because this particular problem will affect people closely, because

the lives of children will continue to be lost till we grapple with

it, and because it is not often one gets a chance of reaching the

Press. Perhaps some wealthy and enlightened person will endow

fly-research, perhaps some organised body will take it up, perhaps

even some day the Government will think of it. I do hope that

in this case the Press will put the issues clearly and definitely.

To return to the flies. We hope to open the exhibition to-

morrow, and to admit the public at certain hours, the health

officials, doctors and technical folk at others. We owe much to

those who have helped to organise, and who have prepared or

lent models, posters, pictures, traps, samples, and appliances.

Dr. C. J. Martin, the Director of the Lister Institute, has very

kindly revised the pamphlet as regards diseases carried by flies,

and we hope to have that out this week.

In these times we all do what we can, and I am privileged to

be able to utilise the facilities of the Gardens for this exhibition.

It is perhaps a new departure for the Society, an unusual feature.

that may be looked on as unseemly by some of the Fellows. I

think that the circumstances justify it, and that if it does good

and anything comes of our work, the Nation will be indebted to

the Society, and this incursion into practical applied entomology

will not be regretted.”

534 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

May 25, 1915.

Prof. E. W. MacBrips, D.Sc., F.R.S., Vice-President,

in the Chair.

Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals, ex-

hibited two pieces of skin cut from the shoulder of a wild boar

and a wild sow (Sus scrofa) to show the difference in thickness

between the two, the skin of that area in the boar being about

four times as thick as in the sow.

Mr. Pocock also exhibited some skins of Asiatic and African

Porcupines, and pointed out the gradation that could be traced

from the Bornean Zrichus through Atherura to Hystria in the

shortening of the tail, the evolution of the rattle, the growth of

the crest on the head, and the elaboration of the spine-armature.

He also showed a piece of the skin of a Javan Porcupine with

some of the quills cut short to illustrate their definite arrange-

ment in short, regular transverse rows.

June 8, 1915.

Dr. 8. F. Harmer, M.A., 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 May, 1915 :—

The number of registered additions to the Society’s Menagerie

during the month of May was 136. Of these 66 were

acquired by presentation, 9 by purchase, 14 were received on

deposit, 35 in exchange, and 12 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 123.

Amongst the additions special attention may be directed to :-—

1 Feline Douroucouli (Aoées felinus), 2 White-browed Hares

(Sylvilagus superciliaris), and 1 Collared Peccary (Tayassu

tajacw), from Banco, Colombia, presented by W. K. Pomeroy,

F.Z.8., on May 3rd.

4 Patagonian Cavies (Dolichotis magellanicus) and 2 Golden

Agoutis (Dasyprocta agouti), from Argentina, received in ex-

‘change on May 4th.

1 Reindeer (Rangifer tarandus), born in the Menagerie on

May 9th.

1 Leopard Cub (Felis pardus), from Kongwe, Nyasaland,

presented by Miss A. Winch on May 3rd.

1 de Winton’s Mouse (Wus sylvaticus wintoni), from Horsham,

new to the Collection, presented by Sir E. G. Loder, Bart.,

V.P:Z.S., on May 21st.

4 Siamese Fighting-Fish (Betta pugnaa), from Siam, new to the

Collection, presented by C. Lamont Groundwater on May 3rd.

ON A ‘“‘NEST ”-MAKING CHIMPANZEE. 535

Mr. E. T. Newron, F.R.S., F.Z.S., exhibited two horns of the

Sabre-horned Antelope (Ory« lewcorya) not attached to the skull,

but supposed to be a pair. The differences between the two

horns, both as regards their curvature and peculiar annulation,

raise a doubt as to their belonging to the same species, unless

these differences may be due to sex. Theyare said to have come

from West Africa. Each of these horns has about five inches of

the basal portion covered in leather, with a large loop of the

same material, which is evidently native work; but for what

purpose this covering was intended is not clear. There is one

horn in the British Museum which has indications of having

been similarly covered.

A‘ nest’’-making Chimpanzee.

Myr. GrorGE JENNISON contributed the following note upon the

“nest? made by a Chimpanzee in the Belle Vue Zoological

Gardens, Manchester :—

A female Chimpanzee (Anthropopithecus calvus) was purchased

for our collection on May 8th, 1913. She was not, in fact is not

yet, adult, but had good health, and was kept until May 1914 in

a rather small cage (12' x12 x10’) having access to the open

air. She was then removed to our new Chimpanzee house and

installed in a large cage, through the middle of which there is a

beam about 3 inches wide. Early in August it was noticed that

she took a small supply of hay, or would even fray out a rope and

lay it carefully along the beam and there lie. .

We therefore nailed a rough branch horizontally from the

beam to the wall,so enclosing a space of about 3 feet, and another

cross-branch to make a very rough base, and provided a supply

of hay, straw, and leafed twigs which were thrown on the floor

ten feet below. Next morning the nest was partly made. Careful

watching by W. Antcliffe, the keeper, showed that she carried up

at first one or two straws and then proceeded to gather a bundle

of twigs, which she tucked between one leg and her thigh,

dragging herself to her nest by her arms and the other leg.

The twigs were carefully arranged with the leaves to the centre

of the nest, and she also gathered up one of her swinging ropes,

which she laid in short paraliel lines on the twigs.

Mindful of Du Chaillu (‘ Exploration in Equatorial Africa’), we

nailed suitable branches over the nest as a basis for a roof, but no

attempt has been made to utilize them to form a shelter, as, of

course, there is no rain in the house.

The animal spends most of her time in the nest, to which she

carries all her food, even a glass of tea, which is taken up like

the nesting material in the hollow of the thigh.

From time to time the nest is either thrown out or falls

through, and is reconstructed with fresh material.

Having succeeded so well with this animal, similar facilities

536 MR. R. E. HOLDING ON

were given to three females (4. troglodytes) in the adjoining cage,

but no attempt was made to use them. Nevertheless, although

they usually sleep on the floor-level, they will carry a sack into

their trees and sleep upon it. A. calvus, the “ nest ”-builder,

will also take up a sack and sleep on it.

The Habits of Chimpanzees in African Forests.

Dr. C. C. Coristy, M.B., C.M., F.Z.S., remarked “ That it was

somewhat misleading to talk of a Chimpanzee’s nest. The little

sleeping-platforms of the Chimpanzee made by bending inwards

the leafy parts of two or three of the smaller branches of some

sapling or larger tree, are quite a feature of the Ituri forests.

“They are generally close to the stem, and often comparatively

low down, sometimes as low as 15 feet from the ground.

“ They are small structures, are occupied by one ora pair of

animals, and are never used a second night. They are made in

a minute by reaching out and pulling in the branches, bending

them or breaking them off.

‘‘Chimpanzees feed largely on the ground, but I am doubtful

if they ever sleep there. They are extremely wary and noisy.

When met with in the daytime they are usually in the trees.

At the first alarm the big males come down from any height in

two swings and a drop and make off, but the females and rest of

the troop swing and climb slowly from branch to branch. They

rarely jump as monkeys do, and being too clumsy to travel

quickly are easily overtaken. For such a big strong animal they

are extraordinarily easy to kill. One serious body wound with a

little -22 bullet is sufficient to bring them down. A slightly

wounded one will make for the top of a big tree,and by breaking

off branches and pushing them beneath him will in less than a

minute construct a big platform, upon which he will sulk or

keep up a furious screeching entirely hidden from beneath.”

Variation in Horns of Cattle.

Mr. R. E. Honprne exhibited and made remarks on several

specimens indicating variation in the horns of certain local

breeds of West-African humped cattle—viz. a single horn of the

Gnami or Botlitli cattle from the neighbourhood of Lake Gnami,

of which a fine skull has recently been added to the British

Museum (Nat. Hist.). The horn exhibited is 4 ft. 5 in. long

over the curve, and if say 11 inches were allowed for width of

skull, the total measurement from tip to tip equals 9 ft. 9 in.

The specimen exhibited was part of a cargo of West-Atrican

horns recently sold in London.

The other West-African specimen, also of the humped breed,

537

VARIATION IN HORNS OF CATTLE.

The frontal bone was

f lyre-shaped black horns.

rounded considerably, thus

Was a palr O

g the horns a posterior direction

giv

oy oyeyug UvIpUy ue Jo preety cligl jo SULME

juepisay opie “y "a “UN Sq (StH FN)

‘ap}qeo peduny Jo Ajortwa youd uBoLIpY “AAN JO susoy padeys-aatryT *—

‘uoryisod [enjor MoYs

Ip 8 wo paovyd ‘ojagng uetIpuy pazworysoutop Jo AJatea .. 1 Jaq] ., JO ULOFT “gq

dof OT SPANO ayy 19A0 dry 0} dy WOLF YJouaT ~puvxypluesyy ‘ozerqsisepy

tmasny, YSyMg ef} oF poyuesatd ApyNaoat “xO wu) JO SULOT, puR [[UYG “Ww

"T aInsy-9xeq,

It is

Length of one horn 36 inches.

thers, the points turning outward.

apparently a common breed.

reaching over the wi

538 MR. ALFRED EZRA ON

The pair of Indian horns shown were those of the “ Delhi”

variety of the domesticated Indian Buffalo (Bos bubalus), of

which there are several local varieties—viz. Deccani, Kathiwar,

and others. The specimen was of unusual form—massive at the

base, with a compressed spiral growing backward.

Mr. Holding also exhibited a print from a German colonial

paper showing another variety of these long-horned cattle from

Ruando, N.W. Africa, where considerable herds are owned by

the Sultan.

Humming-birds in Captivity.

Mr. AurreD Ezra, F.Z.8., exhibited a living specimen of

Ricordi’s Humming-bird (Sporadinus ricordi) and made the

following remarks :—

‘This little bird was brought over from Cuba by a dealer

twelve months ago in almost a dying state. Having been fed

only on sugar and water on the voyage, he was extremely weak

and not able to move from his perch. His feathers were stuck

together with the sticky syrup, and the bird looked miserable,

not being able to make use of his wings. The first thing I did

was to wash him (a difficult matter with such a little mite) in

warm water with a drop of brandy in it, and after drying him

thoroughly he was put back into a cage near a fire. In half an

hour he was buzzing about in the cage and looked much happier.

I fed him as I do my sun-birds, and he took to the food at once.

For the first three nights I kept the light on for him to feed by,

and in a week’s time the bird began to pick up, and grew

stronger every day. About November last he went. through a

partial moult, and again this April he went through a very

heavy moult and got over it perfectly. He is kept in a fair-

sized wire cage, and is given his freedom in a large room every

morning for an hour, when he darts about at a terrific pace

and enjoys it immensely, returning to his cage when he has had

enough. It is astonishing to see how he will never knock himself

against the window as most birds would do, but will always pull

up just in time, no matter what pace he is flying at. Besides the

syrup, which is his chief food, he will eat grapes and aphides, but

I do not think the latter are essential, he having done quite well

without them for all the long winter months. All his food is

taken on the wing, which gives him plenty of exercise, and I am

sure my success 1s due to this. He will not eat the aphides

unless they are flying about, and I have never seen him pick one

up, even if he sees them crawling on his perch. If some are

lying at the bottom of the cage he will fly at a great pace close

to them, and when they fly up he will swallow them one after

another “quickly. The bird is sprayed with tepid water every

morning, no matter what the weather is like, and then he will

wash himself by flying in and out of the damp leaves of a small

HUMMING-BIRDS IN CAPTIVITY. 539

plant placed in the cage. He loves the sun, but seemed quite

happy without it through the winter. For a song he makes a

sound like the sparks of a wireless at work. I think that so long

as the birds have artificial light to feed by in the long winter

nights for two or three hours, they do quite well. They must

be kept warm, for as soon as the temperature drops below 65

degrees they begin to look unhappy. I had a very extraordinary

experience with a Garnet-throated Carib (Hulampis jugularis),

which was sent to me by a friend from Paris last May. It was

brought over by a friend in a small cage which was well wrapped

up. “When I got the bird home I found him lying at the bottom

of the cage, as I thought, dead. He was stone-cold to the touch

and showed a bsolutely no signs of life. J took the bird in my

hand into a very warm room, where, after about half an hour,

I suddenly felt his heart beat ; then he opened one eye and then

the other, and put his long thin tongue out. I put the tongue

into some hot syrup, to which I had added a drop of brandy. ‘He

instantly started to feed, and in another few minutes was flying

about the cage. In ten days this bird was perfectly well, and I

still have him in perfect health, and he is just going through his

second moult with me. It was a very cold day when he was sent

over from Paris, and I think the cold and the want of food were

too much for him. Most humming-birds, I believe, go into a

sort of torpor as soon as the temperature goes down below a

certain point. Both my humming-birds are most pugnacious

and have te be kept in separate cages. To my mind, they are

the most intelligent and fascinating of all birds. My sun-bird

mixture is made up in the following way :—I mix into a paste

one heaped-up tea-spoonful of Mellin’s food, one tea-spoonful of

honey, half a tea-spoonful of Nestlé’s milk, and the inside of

about a dozen meal-worms, and add to this a large breakfast-

cupful of boiling water. All my sun-birds have thriven on this

food, and I have one now that I have had for five years and it is

still in perfect health. I use the same food for the above two

humming-birds—the first and only ones I have ever had—with

great success.”

Proc. Zoou. Soc.—1915, No. XX XVII. 37

No. 147.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON®

June 8th, 1915.

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

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Szcrerary read a Report on the Additions to the Society’s

Menagerie during the month of May, 1915.

Mr. Grorce JENNISON contributed a note upon the “nest”

made by a Chimpanzee in the Belle Vue Zoological Gardens,

Manchester.

Dr. Curnperr C. Curisty, M.B., C.M., F.Z.S., remarked upon

the temporary sleeping-platforms that he had seen made by

Chimpanzees in the Ituri Forest.

Mr. Atrrep Ezra, F.Z.8., exhibited a living specimen of

Ricordi’s Humming-Bird, which had been in his possession for

about twelve months, and described his methods of feeding and

exercising these birds, with which he had been go successful.

Mr. R. E. Hotprne exhibited several specimens showing varia-

tion in the horns of some local breeds of West African Humped

Cattle.

* 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 cn the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Stx Shillings per annum, payable in advance.

Mr, E. T. Newron, F.R.S., F.Z.8., exhibited two horns of

the Sabre-horned Antelope (Oryx lewcoryx), and remarked on

certain peculiarities presented by them.

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

read a paper on the feet, scent-glands, and other external cha-

racters of the Paradoxurine Viverrids, belonging to the genera

Paradoxurus, Arctogalidia, Arctictis, and Nandinia, showing how

these may be distinguished collectively from the Viverrine genera

(Genetta, Viverra, etc.) and also how they may be differentiated

from each other in the characters discussed.

Dr. A. SmirH Woopwarp, F.R.S., V.P.Z.8., read a paper on

the skull of an extinct mammal related to Aelwropus, obtained

from a cave at the ruby mines, Mogok, Upper Burma, which he

described as the type of a new genus and species.

Miss K. M. Parker, B.Sc., communicated a paper on “The

Karly Development of the Heart and Anterior Vessels in

Marsupials, with Special Reference to Perameles.”

In Marsupials, as in Eutheria, the pleuro-pericardial canals

become continuous at an early stage, forming a_horseshoe-

shaped cavity lying round the anterior end of the embryo. The

lateral endothelial tubes first arise in the hind brain-region and

grow forwards. The anterior portion of the pleuro-pericardial

cavity now increases rapidly in extent and its crescentic posterior

wall, which forms the lip of the anterior intestinal portal, moves

backwards as a whole, a process which brings about the length-

ening of the foregut, so that the heart-primordia come to le

ventral to the closed gut. The heart-tubes are brought into

contact with each other by the growth of the pericardium, which

increases rapidly in antero-posterior length without any compen-

satory growth in width.

In the succeeding stages, the heart-primordia increase in

length, and undergo curvature and differentiation into ventri-

cular and auricular limbs. Complete fusion of the endothelial

tubes does not occur till a relatively late stage, and proceeds in

the antero-posterior direction.

The development of the aortic arches is typical, while the

cardinal veins are derived partly from a vessel which lies close

against the neural tube throughout its length and partly from

more laterally situated capillaries.

Lieut. R. Broom, M.D., D.Sc., R.A.M.C., C.M.Z.8.,sent a paper

dealing with certain Triassic Stegocephalians, Restorations are

given ‘of the skulls of Brach yops laticeps Owen and Bothriceps

australis Huxley, which are regarded as forming, with Batracho-

suchus brownt Broom, a distinct family, Brachyopide. Bothriceps

hualeyt Lydekker is shown to differ from Bothriceps australis

in the structure of the occiput, and in having numerous small

teeth on the parasphenoid, pterygoids, and prevomers, and thus

to belong to a very distinct new genus.

This Meeting closes the Session 1914-1915. The next Meeting

of the Society for Scientific Business will be held on Tuesday,

October 26th, 1915, at half-past Frve o’clock p.m.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Socrery or Lonpon,

Recent’s Park, Lonvoy, N.W.

Jume 1ld5th, 1915.

PAPERS.

é Page

. The Head Cavities and Development of the Eye Muscles in Trichosurus vulpecula, with

Notes on some other Marsupials. By Euizanurn A. Frasur, B.Sc.(Lond.), F.Z.S.,

Embryological Laboratory, Department of Zoology, University of London, University

Collese: (Plates I. & IL, and Wext-fioures 1-26.).. 2.2... 2. a eee ee eens ee oe 299

25. On the Organ of Jacobson and its Relations in the “Insectivora.”—Part II. Talpa,

Centetes, and Chrysochloris. By Lieut. R. Broom, M.D., D.S8c., C.M.Z.8., R.A.M.C.

(Pleas MUR ed VEN ol oe Raha eate Ceca bb! Gaul aides Ono ceibeor poms praca mse

2 . On the Anomodont Genera, Pristerodon and Tropidostoma, By Lieut. R. Broow,

Sea aD D2Se,,© MEZ-Ss) RwACM Con (Mext=figures| 14.) ooo ce oe aac Na cides es see alc 300

. On the Triassic Stegocephalians, Brachyops, Bothriceps, and Lydekherina, gen. nov.

By Lieut. R. Broom, M.D., D.Sc., C.M.Z.S., R.A.M.C. (Text-figures 1-3.) ...... 3863

5 zs . A List of the Snakes of Madagascar, Comoro, Mascarenes, and Seychelles. By G. A.

See Bouumnerr, WSR.S., BZS. os. ces cs eee es Soe RTO RES OE CR PREG RO ECPI 369

29. On a Blood-sucking Gamasid Mite (Ichoronyssus serpentium, sp.n.?), parasitic on

Couper’s Snake. By Sraniey Hirst, F.Z.S. (Text-figures 1 & 2.) ........-+.-.. 383.

30. On the Feet and Glands and other External Characters of the Paradoxurine Genera

Paradoxurus, Arctictis, Arctogalidia, and Nandinia. By R.1. Pococn, F.R,S., F.LS.,

F.Z.8., Curator of Mammals. (Text-figures 1-10.) .............+.202seeereeeee 387

. Anatomical Notes on the Gruiform Birds Aramus giganteus Bonap., and Rhinochetus

kagu. By P. Cuaumers Mrrenenn, D.Sv., LL.D., F.R.S., F.Z.8., Secretary to the

Society. (Text-figures 1-5.).. 0... cece ce cece cee eee eee e ee ete c ete e cece ces . 413

. On the Skull of an extinct Mammal related to Hluropus from a Cave in the Ruby

Mines at Mogok, Burma. By A. Smirm Woopwarp, DID EIDE JOLIE AV ley Ash

(Plate I. and Text-figure 1.) 1.0... 1 cece ce cece eee cnet eter e eee e nes sete en ee 425

88. Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.—

: XVII. On Tenia tauricollis of Chapman and on the Genus Chapmania. By

‘Frank E. Bepparp, M.A., D.Sc., F.R.S., F.Z.8., Prosector to the Society. (Text-—

TEU) Re oeion acidbinn omotin ts midh GGb 6) Iori aaiclon Sricc Solr as sexo ror 429

Minchinia: A Haplosporidian. By Hetun L. M. Pixeti-Goopricu, B.So., Beit

Memorial Research Fellow. (Plates I. & II.) ......-.---. 00s see eeeree Gennes 4d.

‘The Early Development of the Heart and Anterior Vessels in Marsupials, with

Special Reference to Perameles. By Karuantne M. Parker, B. Sc.(Lond.), Assistant

in the Department of Zoology, coe ‘of London, University a (Plates -

1. & IL, and Text-figures 1-25.) . Pee TPR A aL hacia) fralel cal eeeiatayerie a stiataV carne fetec Ata) Oh

36. On Spiders of the Family Salticide collected by the British Ornithologists’ Union

___ Expedition and the Wollaston Expedition in Dutch New Guinea. By H. R. Hoge,

_ MLA, F.Z.S. -(Lext-figures 1-11,) ..........- PO A ne gaat Touts teptarctatens Pete (il

LIST OF PLATES,

1915, Parr III. (pp. 299-539).

Page

aS ER vs a Eye-Muscles in Trichosurus ..0+..ecesereee 299

Broom: Pl. III. Organ of Jacobson in Zalpa and Centetes .... } 347

IV. Organ of Jacobson in Chrysochloris ....++....

Woopwarp: Pl. I. 1. Hiuwreidopus baconi. 2. Aluropus melano- \ 495

[eucus erie eo Nee ST RiGE NES Rae rs Ga :

JES alii GOS E EOS et = i WWCDALODIAE Ad SO dow aoo OND oA Cano TON 445

- PARKER ; Pl. I. Heart of Marsupial Embryos ......... sous \ 459

II. Models of Embryonic Marsupial Heart ...... E

NOTICE.

The ‘ Proceédings’ for the year are issued in fowr parts, paged consecutively,

so that the complete reference is now P. Z. 8.1915, p.... The Distribution

is as follows :—

Part I. issued in March.

Penal il einer June.

Pa GIR Says September,

SO eV eae December.

‘ Proceedings,’ 1915, Part II. (pp. 157-298), were published on

June 9th, 1915.

The Abstract of the ‘Proceedings,’ No. 147, is

contained in this Part.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

| OF TITE

ZOVQOLOGICAL SOCIETY

OF LONDON,

1915.

_ — VES — iL

PART IV.

CONTAINING Paces 541 ro 712, witH 2 PuLarzs

DECEMBER 1915.

PRINTED FOR THI SOCIETY,

SOLD AT ITS HOUSE IN REGENT’S PARK

LONDON : |

MESSRS. LONGMANS, GREEN, AND CoO., eS le

| AND 27 Text-ricUREs, TIvLEPAGE, INDEX, ETC.

PATERNOSTER ROW, |

[Price Twelve Shillings.) ae; 2

=e) |

—

LIST Or. CONTENTS

1915, Parr IV. (pp. 541-712).

EXHIBITIONS AND NOTICES.

Page

The Srcrutary. Report on the Additions to the Society’s 1 Menageri ie during the months

of June, July, August, and September 1915 ..........-. +. eee. eens wee e ecb eeeees 705

Mr. R. I. Pocock, F.R.S8., F.Z.S., Curator of Mammals. Exhibition of burrows of

Mrapdoor Spiders, . fate: <\es slates aie ete ove tine welfare efeiercials|= wim aie) sale ptnom eit diate cates seni etOe

‘

Mr. D. Seru-Surrn, F.Z.S., Curator of Birds. Exhibition of a living male hybrid

Siinmalnosieahne, Shien IDNR lao Soee. Puoaewe oben aenbadeooasocas Ra ee ac 707

Mr. D. Seru-Surrn, F.Z.8. Exhibition of an abnormal male Mandarin Duck (4a

gdélericulata). (Text-figure 1.) 20. sci eee ee ee een eee ee etree wees ss oe FOB

‘The Sucretary. Report on the Additions to the Society’s Menagerie during the month

Olt Oolrerse WOLD iad an Gon ooo Gouger Gkoco Teo 6 Obed AG inn ord COG oc a ola sae Sivonee 708

Prof, H. Maxwei.t Leroy, M.A., F.Z.S., Canakor of Insects. Notice of Report on the

House-Bly Investigations... 0... 2.56 Boeke see es ce ee et tee ewe ws ee telcnen nn ne 709

Mr. C. J. C. Poot, Assistant Curator of Insects. Notes from the Caird Insect House .. 709

Mr. R. W. Harotp Row, B.Sc., F.Z.S. Exhibition of photographs of an abnormal Frog.. 712

PAPERS.

37. Some Alcyonaria and a Stylaster from the West Coast of North America. By Sypnny

J. Hickson, M.A., D.Se., F.R.S., F.Z.S8., The University of Manchester. (Plate I.,

and Text-figures 1-5.) «2... 00s eee b eee ee we en eet eee es eUreneae DOUIO Oe 541

38. On Specimens of Cuvier’s Whale (Ziphius cavirostris) from the Irish Coast. By

Sipney F. Harwer, Se.D., F.R.S., F.Z.S., Keeper of Zoology in the British Museum. 559

Contents continued on page 3 0f Wrapper,

ZOOLOGICAL SOCIETY OF LONDON.

Tas Society was founded in 1826 by Sir Sramrorp Rarruzs,

Mr. J. Sasrne, Mr. N. A. Vieors, 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,

Ricwarp H. Burne, Esa., M.A.

Atrrep H. Cocks, Ese., M.A.

Tote Rr. Hon. Tare Eart or

Cromer, G.C.B., F.R.S., Vice-

President.

F. G. Dawrrey Drewitt, KEsa.,

M.A., M.D.

Caartes Drommonp,

Treasurer.

Ksa.,

Tur Kart or Dunmore, V.C.,

M.V.O.

Aurrep Hzra, Esa.

Sipnty Freprric Harmer, Esa.,

MiA., Sc.D., F.RiS:, Vee

President.

Str Epmunp Gites Loner, Br.,

Vice-President.

Pror. Ernest W. MacBripz,

WEA ID ISG, IE IRS, |e

President.

Guy A. K. Marsnatn, Ese.

K. G. B. Meapr-Watpo, Ese.

P. CHatmers Mrrcnert, Esa.,

MET AC DAS crs) LIEe EIR Ses

Secretary. .

W. R. Ocitvin-Grant, Ese.

Aubert Pam, Esa.

Tue Hart or Porrsmoura.

Tue Maravusss or Srieo, F.S.A.,

Vice-President.

AUBYN

M.A.

Antuony H. Wrinertetp, Ese.

Trevor-Barryr, Esa,

Arraur Suitx Woopwarp, Ksa.,

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

The Society consists of Fellows, and Honorary, Foreign, and

Corresponding Members, elected according to the By-Laws. It

carries out the objects of its foundation by means of its 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. BR. I. Pocock, F.RS., 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 HE. Beddard, M.A., D.8c., F.R.S., Prosector,

assisted by Professor H. G. Plimmer, F.R.S., M.R.C.S., Pathologist

to the Society.

TERMS FOR THE ADMISSION OF FELLOWS.

Frriows 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 leu thereof; the whole payment,

including the Admission Fee, being £50.

No person can become a Frrxow until the Admission Fee and

first Annual Subscription have been paid, or the annual payments

have been compounded for.

Fettows elected in November and December are not liable for

the Subscription for the vear in which they are elected.

PRIVILEGES OF FELLOWS.

Fettows have Personal Admission to the Gardens upon signing

their names in the book at the entrance gate, and may introduce

Two Companions daily.

The Wire or Hussanp of a Fetiow can exercise these privileges

in the absence of the Fellow.

Until further notice, Frtrows 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 Week-day Orders, each Order

available for any day during the week except Sunday. Special

children’s tickets are no longer issued, but the Green and White

Cards are perforated, and each half is 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

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Frttows have the privilege of receiving the Society’s ordinary

Publications issued during the year upon payment of the additional

Subscription 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. FxrLtows 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.

Fettows may obtain a Traysrerasite 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 Fettow, 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,

Regent’s Park, London, N.W.

December, 1915.

ZOOLOGICAL SOCIETY OF LONDON

MEETINGS

OF THE

FOR

SCIENTIFIC BUSINESS.

1916.

Turspay, Fepruary .... 8 and 22.

us IMUSR OH: meine 7 aud 21.

We PANP RL tate oto ale 4 and 18.

i Wave cyt hs je 9 and 23.

- Siw, on 2265 6.

BA Ocroper’=/5 2). 24

BS NovEMBER 7 and 21

The Chair will be taken at half-past Five o'clock precisely.

ZOCLUGICAL SOCIETY OF LONDON.

THE ZOOLOGICAL RECORD.

HE Zootogicat Record gives, by means of an annual Volume,

complete lists of the Works and Publications relating to

Zoology that have appeared during the year preceding the issue

of the Volume, together with a Subject and a Systematic Index.

Since 1906 the ZooLoeicaL Rucorp has been amalgamated with

the Zoology volume of the International Catalogue of Scientific

Literature, and has appeared in two forms, different only in title-

page and binding, one in series with the Zoological. Record, the

other forming Volume N, Zoology, of the Annual Issue of the

International Catalogue.

On account of difficulties arising from the War, the Executive

Committee of the International Catalogue is at present unable to

undertake the issue of any volumes of the 14th Annual Issue.

The Zoological Society of London, in these special circumstances,

has undertaken to produce the Zoological Record as usual, pre-

cisely in accordance with the form that the volume has assumed

since the amalgamation.

Fellows of the Zoological Society, and Institutions already on its

subscription-list, and any new subscribers whose subscriptions were

received before August Ist, 1915, will receive the volume dealing

with the literature for 1914 (Zoological Record, Vol. 51: Inter-

national Catalogue, 14th Annual Issue, N, Zoology) as usual at the

end of the year.

Subscribers to the International Catalogue, who wish to assist

the Zoological Society in this heavy burden, and to obtain the

volume as soon as it is ready, may do so by sending 30s. to the

Secretary of the Zoological Society, on or before August Ist 1915.

In the event of the International Catalogue being issued as a

whole for the 14th issue, later on, the sum of 30s. paid in advance

to the Zoological Society will be deducted from the £17 due in

the ordinary way for the complete issue.

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. each.

Vols. 43 to 50 (obtainable separately only from Messrs. Harrison

& Sons), 40s. each.

Vol. 51. 40s. each.

Index Zoologicus, An alphabetical list of names of genera

and subgenera proposed for use in Zoology, as recorded in the

‘Zoological Record,’ 1880-1900; together with other names not

included in the ‘ Nomenclator Zoologicus’ of 8. H. Scudder. Com-

piled (for the Zoological Society of London) by Caries Owen

Warernovuse and edited by Davip 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.

[P T. 0.

Index Zoologicus, No. II. An alphabetical list of names of

genera and subgenera proposed for use in Zoology, as recorded in

the eFoological: Record,’ Vols. 38-47 inclusive (1901-1910), and

the Venleey volumes of the ‘ International Catalogue of Scientific

Literature,’ Annual Issues 1-10. Compiled (for the Zoological

Society of London) by Caartes Owen Wareruovse, I.8.0., and

edited by Davip Swarr, 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.

SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.

Divisions of Vols. 39 to 42 and Vol. 51 of the ‘ Zoological

Record’ can be supplied by the Society, but those of Vols. 43 to 50

can be had only from Messrs. Harrison & Sons, 46 St. Martin’s

Lane, W.C.

List of abbreviations of journals, ete. .. .. 2 O net

Special Records, viz. :—

J. General Subjects .. Oe Ores

II. Mammalia PAD en

IISA Vests) ae OO -

TV. Reptilia and Batr Hehe 2 Ors

V. Pisces phe Lee 220 a

VI. Tunicata a Oe

VII. Mollusca Onur

VIII. Brachiopoda . . Enh as

IX. Bryozoa tO ss

X. Crustacea 2 AO are

XJ. Arachnida Or tel eras

XII. Myriopoda iG aaee

XIII. Insecta TA @ os

XIV. Echinoderma 3 Os

XV. Vermes .. : a) ngs

XVI. Coelenterata .. U Baral ieee ee

XVIT. Spongise PAO

XVIII. Protozoa slug. AON

Index of new names of genera cn sauecnees ZOOM.

P. CHALMERS MITCHELL,

Secretary.

Reaent’s Park, Lonpon, N.W.

December, 1915. {

FOOROGICMiesOClETyY OF LONDON.

LIST OF PUBLICATIONS.

Tue scientific publications of the Zoological Society of London

are of two kinds—“ Proceedings,” published in an octavo

form, and “ Transactions,” in quarto.

According to the present arrangements, the “ Proceedings”

contain not only notices of all business transacted at the scien-

tific meetings, but also all the papers read at such meetings

and recommended to be published in the ‘ Proceedings” by

the Committee of Publication. A large number of coloured

plates and engravings are issued in the volumes of the

“ Proceedings,” to illustrate the new or otherwise remark-

able species of animals described therein. Amongst such

illustrations, figures of the new or rare species acquired in a

living state for the Society’s Gardens are often given.

The “ Proceedings”’ for each year are issued in four parts,

paged consecutively, during the months of March, June,

September, and December. From January 1901 they have

been issued as two half-yearly volumes, indexed separately.

An ‘ Abstract of the Proceedings” is published by the

Society on the Tuesday following the date of the Scientific

Meeting to whichit 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 obtained on the day of publi-

cation at the price of Sixpence, or, if desired, sent post free

for the sum of Six Shillings per annum, payable in advance.

The ‘‘ Transactions” contain such of the communications

made to the Scientific Meetings of the Society as, on account of

the nature of the plates required to illustrate them, are better

adapted for publication in the quarto form. They are issued

at irregular intervals.

Fellows and Corresponding Members, upon payment of

a Subscription of One Guinea before the day of the Anni-

versary Meeting, are entitled to receive the Society’s

Publications for the year. They are likewise entitled to

purchase the Publications of the Society at 25 per cent. less

than the price charged to the Public. A further reduction

of 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 defore the Ist of December following.

The following is a complete list of the publications of ‘the

' Society already issued.

TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 20 vols. and Index. Price to Price to the

Fellows. Public.

Vols. I.-IV. (out of print).

Vol. V., containing 67 Plates.. (1862-66) .... 5 4 3..., 619 0

per Vil. ” 92 ee (LSOG=69)) Gn eo Omer ons OmEO

Ay MiUins 73 moo (lS) Shon LO) 4) O 5555 JB 12 ©

Srey DUR 99 82 A co EAD oor SS Sooo LA JUL &

53 OK, 55 99 oo. Cisne LAIN G555 16 2 ©

. X., 95 SoCs tess JON Sousa lS 7 Y

Index siViolis}SI XS. Sectors teres (esse) Goss OO” Gs OPORTO

Vol. XI., containing 97 Plates .. (1880-85) .... 912 0.... 1216 0O

3 XIL, 3 65 on UIE) Goon. & SB Oi 7-4 0

33 XIII, 0 62 Se ClSO1=95)) eee Om mRo 8 ii <0

bi XIV, <3 47 Ae oo (uch) song 8 1 O , i LOMO)

SGN. me 52 ry oe (CIGSEEIOO) 54 & Na By eh ©

7 NL. ko eo COICO) oe a te) ©. , 74.70

pe ho. 8 Soar ae NE ap brels (1903-1906) Sei Om So. SON: (elsane

55) ICO eee 23) ee we LOO (TOT) er. ton Ole Re} 0)

i a MIDI a 24. od BUSSIGIO) fo IO) 4 Oy. 13 12 0

Pe: OS eat 1, (Pls. I.-V.) .. (Feb. 1912) OAS KOR: 14 0

fp 2 (Celie WAL XV.) (AGMA) .. 2H O . 3 O70

» & (Pls. XVI-XXIII.)(Feb. 1913). I eB TORO

py ae (Pls XX VEO Vie ) -O ¢ 12 0

, 5. (Pls. XX VII.-XXX.)(March1914) 9 0. 12 0

dy (GEIL 2O2ONIL) (Gulle@prelan UNIZ)) 5 BD 5 0

» 1. (Pls. XXXITL., XXXIIL)(Mayl914) 7 6. 10 0

apt RO en hay aL OHA) ere cteeige tere helsene Boo Bc 3.0

sens Q) a(EMlnyon OAL) teres aeestac ora as By 6 3 0

sp AORN (itine WOMAN evens poewenee DIS. 3 0

» Ll. (PILXXXIV.) (Nov. 1914) ¢ @ . OPO

,, 12. (Pls. XXXV.—-XXXVII.)(Nov.1914)9 0. 12 0

» 13. (Pl XXXVITI.) (June 1915) . SemOKe 12 0

periZe Ginnie USMS) eee ea ced oan cede D210): CRO

ls ulna ONS) ue ens epee 2.2. 3 0

» 16. (Pl. XXXIX.) (Oct. 1915) ) Os 2 0

, 17. (Pls. XL.-LIII. Title & Index.)

(iG UO ease oo ncs5 SOOM aa be) (0)

now able to offer for sale, at the reduced price of £30, sets of Vols. V._X VI. 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). Price to Price to the

Fellows. Public.

Part I. 1830-31. 1 vol. 8vo., out of print.

5 I s52! Fy eae Leaitttass tater: canta As Gis one OSs

PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.

First Series.

Parts I-XV. (1833-1847). 8vo. 15 vols. (Letterpress only.)

Fellows :

Index 1830-1847. Price to Fellows: 4s. 6d. ; to the Public, 6s.

4s. 6d. each part ; to the Public, 6s.

Parts I., VII.-IX., XI., XIV., XV., out of print.

SEcoNP SERIES.

Price to

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

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PSAMMOGORGIA TERES. 2. CALIGORGIA FRASERI.

8 SINIGLANS THAR NORWEGICUS,

4. CLAVULARIA MORESBII.

ON NORTH AMBRICAN ALCYONARIA. 541

PAPERS.

37. Some Aleyonaria and a Stylaster from the West Coast

of North America. By Sypney J. Hickson, M.A.,

D.Se., F.R.S., F.Z.S., The University of Manchester.

[Received July 14, 1915; Read October 26, 1915. |

(Plate I.* and Text-figures 1-5.)

INDEX.

MorpHonoey: Page

Note on the Gonophores of the Stylasteride ......... 545

GEOGRAPHICAL DISTRIBUTION ................00.0000.. OAL

SYSTEMATIC :

Stylaster norvegicus Gunn. .......1.....---.-..-..-.-.-.. O44

Clavularia moresbii, Sp. Vi. ... 1.006. 2022-0 eee eee sees s- (046

Tai pangponaape) CAA: NUVI, oes san sop nancedannncacnaccsces Bek

JETPOLUIOG UDANCO Oy Do Wo sas geeseansgecdase tun wnncendeneseece Os!

CUA/DG CIPI J fFREISEIPO, Ds We wat Linctlodsenagnee sesenonsd sedan! | B33

LEST OGOR TOG) CEES OMS gihcesceacss coussenceccaunedee 2!

Although there have been several contributions to our know-

ledge of the marine fauna of the north-west coast of the

American continent in recent years, the Aleyonaria are almost

unknown, <A long while ago specimens of a very large and

remarkable Sea- pen (Osteocella septentrionalis), from the British

Columbian fishing-grounds, were examined and described, but

apart from this solitary example there is no record in the

literature of Zoology, so far as I have been able to discover, of

any other species of the Alcyonaria from this region.

The division of sea-areas into zoo-geographical regions is always

a matter of great difficulty and controversy, and particularly so

along an uninterrupted coast-line extending from the Arctic

Circle to the Equator. Nevertheless, the study of the marime

fauna of the western coast of the North American continent

shows such changes in character as we pass from north to south,

as to justify an attempt to name and define regions of dis-

tribution. .

Such an attempt will not be made in this paper, but there is

just one point bearing upon this division into regions upon

which the study of this very small collection of Aleyonarians may

throw some light.

In his papers on the Mollusca of the west coast of N. America,

Dall (1898) has given the name Oregonian region to the shore-

waters extending from Point Conception, near the south boundary

of Upper California, to, and including, the Aleutian Islands.

Subsequent authors have noticed a remarkable change in the

character of the fauna in the neighbourhood of the Straits of

Fuca, the exit of Puget Sound, which would justify the sub-

division of Dall’s Oregonian region into two nearly equal

* Wor explanation of the Plate see p. 557.

Proc. Zoo. Soc.—1915, No. XX XVITT. 38

542 DR. S. J. HICKSON ON

subregions, one north and the other south of the British-

American frontier.

There is, of course, no zoo-geographical barrier between these

two subregions, and we should expect to find considerable over-

lapping, some of the characteristic southern genera appearing

in the northern subregion and vice versa.

One of the characteristic features of the northern subregion is

the occurrence of genera and even species that are familiar to us

on our North Atlantic coasts, suggesting that they are the

representatives of a circumpolar fauna. Thus Professor Herdman

(1898, p. 249), writing about some simple Ascidians collected in

Puget Sound, says, “I think it may with truth be said that

all the Ascidians I collected in this arm of the N. Pacific are

closely related to familiar species on our own North Atlantic

coasts. ‘This, taken with the similarity between the two faunas

shown in other groups, suggests the possibility that there is a

common northern cireumpolar marine fauna which extends south-

wards on the western coasts of Kurope and America.” This

view is supported by Walker (1898, p. 269), who says, in writing

on the Crustacea collected by Herdman in the same locality,

‘< Besides the species in the collection that are absolutely identical

with the British species, the resemblance between others is

remarkable.”

Tn an account of the Hydroids of the Alaskan expedition

(1910, p. 179), Nutting gives reasons for believing that Puget

Sound is a natural region of demarcation between faune, but

Fraser (1911), in his account of the Hydroids of the Vancouver

Island region, considers that there is no justification for a state-

ment that there is a distinct break at any point along the coast.

“At the present time,’ he says (p. 7), “out of a total of

196 species there is a record of 155 species from the Vancouver

Island region and north of it, and 88 south of that region. No

less than 47, or 24 per cent. of the whole number, are common

to the two. Furthermore, 22 species that are found north of

Vancouver Island are found in the Vancouver Island region as

well as in the region south of it.”

As regards the Aleyonarian fauna of the Oregonian region, we

possess some knowledge of the genera and species found on the

coast of California, 7. e. the southern subregion, thanks to the

researches of Nutting and others, recently summarised and

revised by Kiikenthal (1913), and it 1s therefore of no little

interest to compare them with the few species collected off

Vancouver Island and in the Gulf of Alaska that are described

in this paper.

The following is a list of species of Aleyonaria now known to

occur in the region of Puget Sound and north of it :—

Clavularia moresbi. Caligorgia frasert.

Paragorgia arborea. Psammogorgia teres (sp. ?).

Primnoa willeyt. Osteocella septentrionalis.

NORTH AMERICAN ALCYONARIA, 543

Two of these six species belong to genera (Paragorgia aud

Primnoa) that do not occur in Kiikenthal’s list. As regards the

Sea-pen, Osteocella septentrionalis (Hickson, 1911), it may be a

matter of controversy whether we are justified in separating the

species from the genus Pavonaria, of which two species (P. calt-

fornica and P. willemoesi) have been described from Californian

waters, but there seems to be little doubt that the species is quite

distinet.

Clavularia moresbii is closely related to, but quite distinct

from, the C. pacifica of Californian waters. The genus Clavularia,

however, being cosmopolitan in distribution and having many

very variable species, does not afford much assistance in the

determination of marine zoological regions.

The genus Psammegorgia, on the other hand, appears to have

principally a tropical and temperate distribution, and the occur-

rence of one species north of the Straits of Fuca may be regarded

as an example of the fauna of the southern subregion over-

lapping the boundary-line. Three species of this genus have

been described by Nutting from Californian waters, but I have

found the determination of species of Psammogorgia, without the

examination of type-specimens, so extremely difficult that I feel

great hesitation in my identification of the Vancouver specimen

as Ps. teres and can make no further comments upon it.

The genus Caligorgia has many species in the Pacific Ocean,

but according to Versluys (1906, p. 169) it is unknown in the

North Atlantic. The species found off the coast of California

(C. kinoshite), however, is quite distinct from the species

described in this paper from the Gulf of Alaska.

The occurrence of a specimen of Stylaster in the Vancouver

seas is of interest, because it belongs to the same form or sub-

genus (Allopora) that occurs in the Norwegian fjords, and not to

the form or subgenus (Séylaster) which is so common in tropical

and subtropical waters. It has been previously described by

Verrill from the coast of California, but is probably a migrant

from the north.

To summarise the results, it may be said that in this small

collection three species at least (Stylaster norvegicus, Parayorgia

arborea, and Primnoa willeyt) are representatives of a cireumpolar

fauna, one (Psammogorgia teres) is a representative of the south

coast fauna, and the other three (Clavularia moresbii, Caligorgia

fraser, and Osteocella septentrionalis) may represent a common

Pacific element which extends both north and south of the line

between the two subregions.

A comparison of the list of species described in this paper with

Kiikenthal’s list of Californian species shows that not a single

species of Aleyonaria (except possibly the Psanmogorgia) has

been found both north and south of the Straits of Fuca, and

seems therefore to justify a division of the Oregonian region at

that point into two subregions. The specimens I have been

able to collect together may represent only a small fraction of the

38*

544 DR. S. J. HICKSON ON

Aleyonarian fauna of the Columbian and Alaskan waters, and

subsequent researches may modify any deductions that may be

drawn from them, but so far as our knowledge extends at present

the facts are significant.

I wish to acknowledge my indebtedness to Professor A. Willey,

F.R.S., and to Mr. McLean Fraser for the specimens described

in this paper; to Miss Constance M. Lightbown, B.Sc., for much

valuable assistance in making preparations and drawings, and to

Mr. J. T. Wadsworth for taking the photographs (PI. I.) and

for the drawing of text-figure 3.

Order STYLASTERINA.

StyLasTeR (ALLOPORA) NoRVEGICUS Gunnerus. (PI. I. fig. 3.)

Millepora norvegica Gunnerus, 1768.

Allopora californica Verrill, 1868, Essex Inst. vol. v.

Allopora oculina Moseley, 1881, ‘ Challenger’ Reports, vol. ii.

p. 89. '

Stylaster norvegicus Broch, 1914, Danish ‘ Ingolf’ Expedition.

Swiftsure Shoal, off Barkley Sound, W. coast Vancouver

Island.

Local name. Roseate stag’s horn coral.

A single dried specimen of this Stylaster was taken by

Professor MeMurrich from the Swiftsure Shoal.

It is 45 mm. in height, with seven short blunt branches

arranged in a single plane. One of the branches shows a barnacle-

gall. As there is no base of attachment, the specimen may be a

branch of a much larger colony.

The main stem is 10 mm.in diameter, and the branches about

5 mm. in diameter. :

The cyclosystems are evenly distributed on all sides of the

branches, and they are not more numerous on one side of the

flabellum than on the other. Hach cyclosystem projects slightly

from the surface of the ccenosteum and is about 0°75 mm. in

‘diameter.

The number of dactylopores in each cyclosystem varies, but in

the majority of cases there are 6 or 7. There is a large brush-

like style in the gasteropore and a very small style in each of the

dactylopores. No ampulle can be seen on the surface of the

ceenosteum, but at the broken base a few small cavities (0°5 mm.

in diam.) may be seen which are probably young ampulle.

The colour is salmon-pink.

The difficulty of separating the two Stylasterid genera Allopora

and Stylaster was pointed out ten years ago by myself and

Miss England (1905, p. 6). Broech (1914), agreeing with our

view on this matter, has included Allopora in the genus Stylaster,

retaining the name Allopora as a subgeneric name for species of

Stylaster included in our group of species D. “ For group D the

old generic name Allopora should be retained.” This is clearly

NORTH AMERICAN ALCYONARTA. HAD

an error. According to our definition of the groups, group D

has cyclosystems on the anterior surface of the branches only. It

is group C which has the cyclosystems evenly distributed over

the surfaces of the branches. In Allopora nor vegica, according

to Broch’s own description and figures of the species, as well as in

the Cape species (Allopora nobilis), the cyclosystems are dis-

tributed on all sides of the branches, they are certainly not

confined to the anterior surface of the flabellum, as they are in

the species of our group D.

The subgenus Allopora, however, is usually distinct from the

other subgenus Hustylaster of Broch, not only in the character

given to our group C—that the cyelosystems are distributed on

all surfaces of the branches,—but also in two ill-defined but

still mutually dependent chamaitens. namely, that the terminal

branches are relatively thick, and that the ampullee do not project

or project slightly from the surface of the ceenenchym.

Returning now to our species from Vancouver Island. It

clearly belongs to our group O, as the cyclosystems are more or

less evenly but irregularly distributed on all sides of the branches.

It may therefore be placed in the subgenus Allopora of the genus

Stylaster. The determination of the species is a much more

difficult matter in the absence of any information about the

ampulle or gonophores. The characters of the ccenosteum, as seen

without fracture, are similar to those of the species from the

Norwegian coast except in respect of colour, which is salmon-red

instead of white or faintly rose. The colour-difference by itself

does not seem to me to be a character upon which it is wise to

establish a distinct species, and therefore | am disposed to regard

the species as identical with the Norwegian species.

The proper name of this species has recently been discussed

by Broch (1914, p. 17), and I am in agreement with him that

it should stand as Stylaster (Allopor i“) norvegicus Gunnerus.

Whether this species is identical or not with the Allopora oculina

of Ehrenberg cannot be determined with any degree of certainty,

but there can be little doubt that it is the same as the Allopora

oculina of Moseley’s ‘ Challenger’ Report (1881, p. 85).

It seems very probable that the species is also identical with

Verrill’s Allopora californica trom deep water off the coast of

California. Verrill’s specimen was 3 inches (75 mm. ) or more in

height, had cyclosystems ‘02 inch (0°5 mm.) in diameter, with,

usually, six dactylopores, and was light “* minium” red in enon.

From the description there are no points of distinction of this

form from Allopora norvegica, except colour.

Note on the Gonophores of the Stylasteride.

In his recent work, Broch (p. 20) attributes to Moseley the

view that the gonophore of the Stylasteridz is a special formation

in the group, and is not homologous with the adelocodonie

gonophore of other Hydrozoa. This was certainly not Moseley’s

546 DR. S. J. HICKSON ON

view, and when I suggested it to him he expressed to me per-

sonally his disagreement with it. The term ‘“ trophodise” was

suggested by myself and was never used by Moseley at all.

As regards the view itself. It was expressed as a result of a

long investigation of the development of the gonophores of

Stylaster (Allopora) from the coast of Norway, and every fact

described was confirmed by the examination of many comparable

preparations. Knowing now, after twenty-five years’ experience

of this group, better than I did then, the difficulty of the

investigation, I realise the probability that some stages in the

development may have been missed. Moreover, the study of

Kuhn’s excellent memoir on the development of the gonophores

of Hydrozoa has to some extent shaken my faith in my own

view; but the homology of the gonophores of Stylasteride with

the adelocodonic gonophores of other Hydrozoa has not yet been

proved, and wili not be established by scattered observations on a

few stages of the development of the male gonophore alone. The

principal difficulty mm accepting the older view held by Moseley

arose from my observations on the development of the female

gonophore, and until this investigation has been repeated with

more modern methods of study than I had at my disposal in 1890,

the true homologies of these organs must remain undetermined.

In the meantime the discovery that Millepora, notwithstanding

its calcareous skeletal structures, does give rise to free-swimming

meduse, has removed one of the principal initial difficulties I felt

im believing that the gonophores of Stylasteride could represent

reduced meduse, and I am quite prepared, when the time comes,

to abandon my own view in favour of the more conventional and

older one of Moseley.

Order ALCYONARIA.

Family CLAVULARIIDS.

CLAVULARIA MORESBII, sp. n. (PI. I. fig. 4; Text-fig. 1.)

W.S.W. off Moresby Island, 100 fathoms.

The widely distributed and very variable genus Clavularia is

badly in need of revision. It is probable that such a revision

would lead to a considerable reduction in the number of the

species, many of which have been founded on very inadequate

characters. Nevertheless, the specimens from Moresby Island

show very distinet specific characters and must be regarded as

the type of a new species.

The genus is well represented both in the Norwegian waters

and in the North Pacific Ocean. ‘The occurrence of a species in

British Columbian waters might have been anticipated, and in

itself is not a fact of any zoo-geographical importance.

The specimens were obtained by Prof. Willey in 1914, who

writes that these whitish ‘ rose-headed” Clavularias were growing

NORTH AMERICAN ALCYONARIA, 547

on the stem of the Gorgonid (Primnoa willeyi). They were

preserved and forwarded to me in formalin.

The stolon is in the form of flat anastomosing bands, spreading

out in places and fusing to form membranes. From the stolon

the zooids arise in a very irregular manner, in some places at

considerable intervals, in others close together. The stolon

follows the support in a quite irregular manner, and, so far as can

be determined from the material at my disposal, is never thickened

to form a sympodium. The zooids are never retracted into the

stolon, but they all show the tentacles contracted tightly over the

oval dise. They vary very much in size, large and small ones

being irregularly distributed on the stolon.

Text-figure 1.

Spicules of Clavularia moresbii. X 350 diam.

The larger zooids are from 7-10 mm. in length, with a diameter

of 2mm. ‘The crown of tentacles is never retracted into a calyx,

and the aboral surfaces of the eight tentacles formiug the apex of

the zooid, in the preserved state, therefore can always be seen.

The body-wall is smooth and cylindrical. It does not shew, as

preserved, the eight longitudinal furrows that have been described

in many other species of the genus.

The spicules (text-fig. 1) are tuberculated spindles 0-15 mm.—

0°2 mm. in length by about 0:05 mm. in diameter, and do not

548: DR. 8S. J. HICKSON ON

shew a tendency to become club-shaped. They are densely

crowded both in the tentacles and body-wall, and are nearly all

arranged in a direction parallel with the long axis of the zooid.

The species seems to be most closely related to Clavularia

pacifica (Kukenthal, 1913, p. 237) from the coast of California,

previously described by Nutting (1909, p. 686) as Sympodium

armatum. This species, however, differs from Clavularia moresbir

in having stouter and more retractile zooids (5 mm. x 2 mm.),

with eight deep longitudinal grooves and larger ,spicules (0°25—

0-3 mm. in length). The spicules, moreover, are much more

crowded together in Clavularia moresbii, and do not show in such

marked degree as in C. pacifica a transverse disposition at the

base of the tentacles, and I cannot find in my preparations any

spicules that by becoming thickened at one end show a tendency

to be club-shaped.

In Clavularia eburnea (Kiikenthal, 1906, p. 14) from Japanese

waters the zooids are larger (12 mm.), but in the spicular

armature and in other characters C. ebuwrnea is more closely

related to C. pacifica than it is to C. moresbit.

But, although the relationship of our new species with the

@alifornian species C. pacifica 18 pronounced, it must be pointed

out that its relationship with some of the Norwegian species, such

as C. borealis of Koren and Danielssen (1883, ple 1.), may be as

close, and it affords therefore no special reason for regarding

the fauna of British Columbia as being more closely “related

to the Pacific than to the N. Atlantic fauna.

Family BRIAREIDA,

PARAGORGIA ARBOREA Linn. (Text-fig. 2.)

Alcyonium arboreum Linneus, Syst. Nat. 10th edit. 1758,

p- 803.

Alcyoniwm arboreum Pallas, Klenchus Zooph., Edit. Wilkens,

1787, pt. 2, p. 164.

Paragorgia arborea Milne-Kdwards, Hist. Nat. Cor. 1857, t. 1,

jon UO.

Paragorgia nodosa Koren & Danielssen, Nye Alcyonider, ete.

1883, p. 19.

? Paragorgia nodosa Nutting, Pacific Aleyonaria, 1912, p. 99.

2 Paragorgia regalis Nutting, |. c. p. 100.

Off Kodiak Island, Gulf of Alaska. Depth? 1 specimen.

Local name. Fiiable brick-red coral.

In Wilkens’ edition of Pallas’ ‘ Elenchus Zoophytorum ’ there

is a long account of this species, probably copied in great measure

from the writings of Koelreuter (1761). There can be no doubt

from this that the Aleyoniwm arborewm of Linneus and Pallas

is the same species as the common species of the Norwegian

fjords, now known as Paragorgia arborea.

Although the external features of the species were fully

NORTH, AMERICAN ALCYONARTA. 5AYQ

described by Koelreuter, and measurements and illustrations of

the spicules of Paragorgia nodosa—which does not seem to me a

distinct species—are given by Koren and Danielssen, there is no

good modern description either of the genus or species.

The genus, however, seems to be quite well distinguished from

others of the same family by the very well-marked dimorphism

of the zooids and by the characters of the spicules. The occur-

rence of dimorphism in the genus, first recorded by myself in

1883, is of importance because it is of only exceptional occurrence

in the Pseudaxonia, and can be clearly determined, not only in

fresh and spirit specimens, but also in specimens that have been

dried for many years.

The geographical distribution of Paragorgia arborea cannot be

very accurately determined from the literature. It is probable

that some of the specimens from the Mediterranean Sea and

Atlantic Ocean, refered. to Aleyoniwm arboreum by the older

naturalists, belonged to a different genus or species. Of recent

years there is no record of any specimens being found outside

the area of the Norwegian coasts. It was not found by the

‘Challenger’ Expedition in the Atlantic, nor is it recorded from

the deep water off the west coast of Ireland by the Irish

Fisheries Investigations.

It was therefore with some surprise that I found, in the collec-

tion sent to me by Mr. Fraser from the Gulf of Alaska, a

specimen that was clearly a species of Paragorgia. For com-

parison with the Alaskan specimen I have examined a specimen

of P. arborea from the Trondhjem Fjord, and I have no hesitation,

after making this comparison, in placing the two specimens in

the same species.

The specimen is probably only a fragment of a large colony,

but it shows a simple bifurcation at the distal end. It is

170 mm. in length. The stem at the base is oval in section

(14x18 mm.). As in the specimen from Norway, the stellate

pores of the autozooids are scattered irregularly in clusters all

round the stem, many of the clusters being mounted on dome-

shaped prominences from the surface. The pores of the siphono-

zooids are numerous, quite irregularly distributed, and easily seen

with ahand-lens. The axial part of the stem is creamy-white in

colour, and is penetrated by canals corresponding in arrangement

with those in my specimen of the type-species. The crust, or

outer layer, of the stem is about 1 mm. in thickness, and is

distinguished by its brick-red colour.

The spicules of the autozooids are irregularly tuberculated

spindles, of which the larger ones are about 0°25 mm. in length.

These spicules are probably distributed in the tentacles or body-

wall of the anthocodiz of the autozooids, but I cannot, determine

this with certainty.

The spicules of the cceenenchym and axial region are double

stars of the type shown in Kéolliker’s ‘ Icones,’ pl. xviii. fig. 45.

They are of nearly constant size, 0°07 mm. in length.

0) DR. S. J. HICKSON ON

Text-figure 2.

A. Spicules of Paragorgia arborea from Alaskan coast. 375 diam.

B. Spicules of Paragorgia arborea from coast of Norway, for comparison

with A. X 375 diam.

NORTH AMERICAN ALCYONARIA. ill

A comparison of the preparations of spicules, made from the

Alaskan specimen and from the Norwegian specimen, shows that

there is very little difference either in size or shape between the

spicules of the two specimens. In fact, the preparations can

hardly be distinguished without the assistance of the labels

(text-fig. 2, A, B).

In a recent paper, Nutting (1912, p. 99) has described two

species of the genus from the Japanese seas, one he attributes

to the species P. nodosa of Koren and Danielssen and the other

to a new species, P. regalis. It does not appear to me that

either of these species is very well defined from the type-species,

but without examination of type-specimens from Japan it is

impossible to determine with certainty whether they are identical

with P. arborea or not.

Family PRIMNOID 4.

Subfamily Prrmnoin®.

PRIMNOA WILLEYI, sp. n. (Text-fig. 3.)

Locality. W.S.W. off Moresby Island, British Columbia,

100 fathoms.

Concerning this species Prof. Willey writes: ‘In fishing for

halibut a magnificent scarlet Gorgonid was brought up on one

of the hooks. It was four feet in height, with a diameter at the

broken off base of 1:5 inches. The branches anastomose and

the axis is black and horny.”

Specimens of the Clavularia described above were growing on

the base of the horny stem of this Primnoa.

The only specimens sent to me were a number of fragments

well preserved in formalin. I am unable therefore to give an

account of the colony as a whole or its method. of branching.

The method of branching, so far as I can judge, is dichotomous,

but I have no evidence of the anastomoses referred to by

Prof. Willey.

The structure and arrangement of the zooids, however, afford

sufficient evidence to show that the species is not identical with

any that has hitherto been described.

At the time of the publication of Versluys’ memoir on the

Primnoide (1906), there was only one well-established species of

the genus, the well-known Gorgonia reseda of Pallas, subse-

quently called Primnoa lepadifera by Lamouroux.

Since that date Kinoshita (1908, p. 42) has described a new

species, Primnoa pacifica, from the Sagami Sea.

The genus Primnoa is distinguished from other Primnoine by

the irregular distribution of the zooids on the branches—or, in

other words, the zooids are not arranged in definite whorls nor

in definite spirals. Moreover, it seems to be a character of the

two known species that the zooids are bent downwards away

552 DR. 8. J. HICKSON ON

from the apex of the branches, instead of upwards towards the

apex of the branch, as they are in most of the Primnoine.

In the specimen from British Columbia, the terminal branches

including the zooids are about 6 mm. in diameter and excluding

the zooids about 3 mm. in diameter. The zooids are quite

irregularly distributed on the branches, larger and smaller ones

being mixed, and they are all bent downwards towards the base

of the branch. The larger zooids are about 5 mm. in length by

about 1°5 mm. in diameter.

The opercular scales are triangular in shape, with a very well-

marked keel passing along the adoral side from the apex towards

the base. These scales are 1-5 mm. in length (from apex to base)

and 0°6 mm. in breadth. Behind and partly overlapping the

Text-figure 3.

A single zooid of Primnoa willeyi. X 15 diam.

opercular scales on the abaxial side there is a half circle of large

oblong scales 1°05 x 0-75 mm., and arranged very irregularly over

the rest of the abaxial side there are long narrow scales of various

sizes up to 1-8 mm. in length by 0-2 mm. in breadth (text-fig. 3).

Some of these elongated scales occur on the adaxial side of the

margin of the zooid overlapping the opercular scales, but the

greater part of this side of the zooid is naked.

In Primnoa reseda and in Primnoa pacifica there is less

difference between the marginal scales and the other seales of the

abaxial side of the zooids, and in both these species the zooid is

more completely covered and protected by square or oblong

scales.

The new species differs from both the other two species, not

NORTH AMERICAN ALGYONARIA. 553

only in the shape and arrangement of the scales as described

above, but also in the actual size of the larger zooids.

I have compared them with the zooids of a specimen of

P. reseda from Norway, and found that, whereas the measure-

ments of spirit-specimens of P. willeyi are 5 mm. by 1:5 mm.,

in dry specimens of P. reseda they are 7mm. by 3mm. In

P. pacifica the zooids are said to be 5-7 mm. in length, and from

the figure are evidently broader and stouter than in P. willeyi.

CALIGORGIA FRASERI, sp. n. (Pl. I. fig. 2; Text-fig. 4.)

Gulf of Alaska, 50-100 fathoms.

Local name. “ Verticillate fan-coral.”

This new species 1s represented by two dried specimens 220

and 130 mm. in length respectively. The base of attachment is

Text-figure 4.

Scales of Caligorgia fraseri. X 280 diam.

missing from both specimens, and consequently they may repre-

sent portions of a larger colony. The branches arise alternately,

but irregularly, from the main stem (or branch) in one plane.

The diameter of the largest stem, including the whorl, is 3-5 mm.,

and of the axis 2 mim.

d54 DR. S. J. HICKSON ON

The zooids are arranged in closely-set but not overlapping

whorls. In the thickest branches there are 11 or 12 zooids in

each whorl, but they diminish to 5 or 6 in the more slender

distal branches. The zooids are about 1 mm. in length by

0-5 mm. in diameter, and are closely adpressed to the side of the

branch.

The zooids are protected on the abaxial side by three or four

longitudinal rows of overlapping scales, but the axial side is free

from scales except at its distal extremity. The apex of each

zooid 1s protected by a complete circle of triangular opercular

scales.

The large abaxial scales are round or oval in shape, and the

outer surface is ornamented with numerous long spiny tubercles .

which radiate outwards from a common centre.

It is the presence of these remarkably long tubercles on

the scales that constitutes one of the most important characters

of the species. ‘These scales attain a size of 0°3 mm. x 0°23 mm.

The triangular opercular scales are also covered with long

tubercles, and their size may be 0°28 mm. in height by 0:2 mm.

at the base.

In addition to the abaxial plates and opercular scales, a pre-

paration of the spicules of a zooid reveals a number of smaller

seales and irregular tubercular calcareous nodules. The exact

position of the latter cannot be determined owing to the density

of the plates which cover them, but they probably correspond

with the deep-seated warty sclerites described by Versluys (1906,

p- 76) in Caligorgia ventilabrum, but far more commonly found

in the genera Primnoella and Prinnoides. The presence of these

sclerites constitutes a second important character of the species.

The new species appears to be most closely related to Caligorgia

aspera (Kinoshita, 1908, p. 39) from the west coast of Satsuma,

Japan, from which it differs in the less profuse branching, in the

smaller number of zooids in a whorl, as well as in the larger and

more profuse tuberculation of the scales.

In the method of branching it is more like C. granulosa of the

same author (p. 37), but in this species the scales are much

larger. In C. elegans (Gray), also described by Kinoshita from

the coast of Japan (p. 40), the number of zooids in a whorl

corresponds more closely with that of the new species, but the

tubercles on the scales are much smaller.

The specimens were taken on the Halibut lines on Albatross

and Portlock banks, in the Gulf of Alaska, and are said to be

pink when fresh and to be “common.” They were collected by

Professor A. Willey, F.R.S.

Family PLEXAURID4.

PsamMocoreia TERES Verrill. (PI. I. fig. 1; Text-fig. 5.)

Psammogorgia teres Verrill, Trans. Conn. Acad. vol. i. 1868,

p. 416.

NORTH AMERICAN ALCYONARIA. 555

W. coast of Vancouver Island.

Local name. Coral-pink candelabrum coral.

The genus Psammogorgia is so badly in need of revision and

full description that I refer this specimen to Verrill’s species

with the greatest hesitation. The type of the species was taken

off Pearl Island, which I believe to be off the west coast of

tropical America, in 6-8 fathoms, and is said to be rare.

The specimen from Vancouver Island is dry, and, in the

presence of a flattened base of attachment, is evidently complete.

Tt is 115 mm. in height and has six branches. The diameter of

Text-figure 5,

Spicules of Psammogorgia teres. X 500 diam.

the thickest branch is about 4 mm., and the branches are almost

cylindrical in shape. The diameter of the axis just below the

point where ramification begins is 4 mm., and the disk of attach-

ment is thin and about 10 mm. in diameter. The crust varies in

thickness from about 0:25 mm. below to | mm. near the terminal

ends of the branches.

The positions of the zooids are marked by flat or slightly

convex prominences, quite irregularly distributed over the sur-

face of the conenchym, and in the centre of each prominence

there is a stellate aperture.

556 DR. S. J. HICKSON ON

The spicules are tuberculated spindles, very variable in size,

but with an average of about 0°12 x 0°04 mm., and double stars

rather smaller in size (text-fig. 5).

The genus Psammogorgia is represented by four species from

the tropical region of the west coast of America described by

Verrill (1868), and by three species from the coast of California

described by Nutting (1909, p. 719). In a recent paper

Kiikenthal (1918, p. 268) expresses the opinion that one of

Nutting’s species should be referred to the genus Huplexauwra

of Verrill, as amended by him.

It appears to me very doubtful whether the genus Psamme-

gorgia will stand as an independent genus, but I am not disposed,

until a further study is made of the species attributed to it from

the west coast of America, to merge it into the more widely

distributed genus Huplexaura. The diagnostic characters of the

species are all most unsatisfactory, and it is almost certain that

if the genus stands it will stand as a single-species genus. In the

meantime, I refer the specimens from Alaska to the species to

which they appear to be most closely related.

LITERATURE.

1914. Broca, H.— Danish Ingolf-Expedition. Vol. v. 5.

Stylasteridee.

1898. Datu, W. H.—The Fur Seals and Fur Seal Islands of the

Northern Pacific. Edited by D. H. Jordan. Part ii.

pp. 9389-546.

- 1911. Fraser, C. McL.—‘‘ West Coast Hydroids.” Bull. Univ.

Towa, No. 28, New Series.

1768. Gunnerus, J. E—‘“Om nogle norske Coraller.” Norske

Selsk. Skr. 4 deel. (/%de Brocn.)

1898. Herpman, W. A.—‘ Description of some simple Ascidians

collected in Puget Sound.” Trans. Liverpool Biol. Soc.

vol. xii. p. 248. ,

1883. Hickson, 8. J.—‘“‘ On the Ciliated Groove in the Stomo-

deeum of the Alcyonarians.” Phil. Trans. R. 8. 111.

1905. Hickson, 8. J., and Eneranp, H. M.—Stylasterina.

Siboga-Expeditie, viii.

1911. Hickson, 8. J.—‘‘On a Specimen of Osteocella septen-

trionalis.” Manch. Mem. vol. lv. No. 23.

1847. Jounston, G.—Zoophytes. 2nd edition, p. 171.

1908. Kinosuira, K.—‘‘ Primnoide von Japan.” J. Coll. Sci.

Univ. Japan, xxi. Art. 12.

1761. Kortreuter, J. F.—‘ Zoophyti marini.” Nov. Comment.

Acad. Sci. Imp. Petropolitanee, vol. vil. pro Ann. 1758-

1759.

1883. Koren and Danietssen.—Nye Alcyonider, Gorgonider og

Pennatulider. Bergens Museum.

1906. KiikentrHat, W.—“ Japanische Aleyonaceen.” Abhi. der

K.-Bayer. Akad. Wiss. Suppl.-Bd. I. Abt. 1.

NORTH AMERICAN ALCYONARTA, 557

1913. KtxenruaLt, W.—‘‘ Ueber die Alcyonarienfauna Cali-

forniens.” Zool. Jahrbiicher, Syst. xxxv. p. 219.

1857. Mitne-Epwarps, H.—Hist. nat. des Coralliaires, t. 1,

p. 190.

1909. Nurrine, C. C.—* Aleyonaria of the Californian Coast.”

Proce. Nat. Mus. Washington, vol. xxxv. p. 719.

1910. Nurrine, C. C.—Hydroids of the Harriman Alaskan

Expedition. Vol. xiii.

1912. Nurrine, C. C.—‘‘ Aleyonaria from Japanese Waters.”

Proc. U.S. Nat. Mus. vol. xlii. p. 1.

1787. Pauuas, P. S.—Characteristik der Thierpflanzen. Trans-

lated into German by C. F. Wilkens. Niirnberg,

Part 2.

1868. Verrint, A. E.—‘ Synopsis of Polyps and Corals.” Com-

munications of the Essex Institute, 1868, p. 37.

Usually bound with Proc. Essex Inst. vol. v.

1869. Verritt, A. E.—‘‘ Notes on Radiata.” Trans. Conn.

Acad. vol. 1.

1906. Verstuys, J.—Die Primnoide. Siboga-Expeditie, xiii a.

1898. Wanker, A. O.—‘“ Crustacea collected in Puget Sound.”

Trans. Liverpool Biol. Soc. vol. xii. p. 268.

EXPLANATION OF THE PLATE.

Fig. 1. Psammogorgia teres (dry). Nat. size.

2. Caligorgia fraseri, sp. n. (dry). Nat. size.

3. Stylaster norvegicus (dry). Nat. size.

4. Clavularia moresbii, sp. n. Two zooids preserved in spirit. X 2 diam.

[NOTE. z=

My attention has been called to a paper by W. H. Dall, “On some Hydro-

eoralline from Alaska and California,” in the Proc. Biol. Soc. Washington, vol. 1.

1885, p- 111.

In this paper three new species of Allopora are described from the Aleutian and

Shumagin islands, namely, A. verrillii, A. moseleyi, aud A. papillosa.

They differ in some respects from the specimen I have identified as Stylaster

(A.) norvegicus from the Swiftsure shoal off Vancouver Island, but I do not

consider, without reference to the type specimens, that these differences are

sufficient to justify a specific distinction from the older species.

In this paper there is also a record of a specimen identified as Calligorgia

compressa Verrill from the Aleutian islands, but as there is no figure or description

of it, comparison with the specimen described by me as Caligorgia fraseri cannot

be made. The type specimen of V7. compressa is (fide Versluys, p. 81) only a

naked axis without polyps or spicules.

December 13, 1915. Se epeley

Proc. Zoot. Soc.—1915, No. XX XIX. 39

ON CUVIER'S WHALE. 559

38. On Specimens of Cuvier’s Whale (Ziphius cavirostris)

from the Irish Coast. By Stpney F. Harmer, S8c.D.,

F.R.S., F.Z.8., Keeper of Zoology in the British

Museum *.

[Received October 15, 1915: Read October 26, 1915.]

INDEX.

GEOGRAPHICAL: Page

New records of the occurrence of Ziphius cavirostris in British

TW ELEOT:Sy erates ctet nen aioe ara Pee RS tee RE cco eA aL 559, 561

Dis tiilloubromtet a cease eoeatey ee cies eae tonal ocasce see aus tes 560

STRUCTURE, ETC.;7

Coloratnontand¥skin-rnankameseeee eer eeeeee a peers sere -ceeesesee 562

Uy e9 0 Nes AR SIERO Sere ecl bis Dane Geib ae ae Bar aaa Sooo c nocc HAAEES DECOM M PRO LOU NS Or-Aa 0741

Sexualiditterences, auaitesnccans sate saeichreceateasenee adncspements 563

SYSTEMATIC :,

The Ivish specimens belong to Z. cavir0stris..........cccce cee eee ees 565

Among the Cetacea included in lists of the British species, the

subject of this notice is one on which further information is

specially desirable. Its claim to be regarded as British was

established by Professor (now Sir William) Turner (1872, 1912),

on the evidence of a skull obtained off Hamna Voe, Northmaven,

Shetland, and now in the Anatomical Museum of the University

of Edinburgh. So far as I have been able to ascertain, this is

the only authenticated record of the occurrence of Ziphius cavi-

rostris in the British area. Van Beneden indeed states (1888,

pp. 87, 91) that a male specimen of this species was stranded on

the Irish Coast (place and date not indicated), and that its

skeleton is in a Dublin Museum. In order to obtain information

with regard to this record I wrote to Dr. R. F. Scharff, Keeper

of the Natural History Collections in the National Museum,

Dublin, who has been good enough to inform me that he knows

nothing whatever of the supposed Ziphius, and suggests that

Van Beneden may have mistaken a record of an Irish J/esoplodon

bidens for one of Ziphius cavirostris. He assures me that the

National Museum at Dublin possesses no skeleton of Ziphius or

any part of one; and that he has satisfied himself that no such

skeleton exists in the Museum of the Royal College of Surgeons,

Dublin, or in that of Trinity College, Dublin.

Although I am thus unable toascertain what was the evidence

on which Van Beneden’s statement was made, I am in a position

to record the oecurrence of two undoubted specimens of Z. cavi-

rosiris on the Southern Coast of Iveland. The circumstances

under which these have been obtained by the British Museum

lead me to suspect that the species is not so uncommon a visitor

* Published by permission of the Trustees of the British Museum.

39"

560 DR. S. F. HARMER ON

to our shores as has hitherto been supposed*. It seems not

improbable that a certain proportion of the Cetacea recorded

as ‘ Bottle-nosed Whales” belong either to this species or

to the allied genus JMJesoplodon, of which two species appear

to occur in our seas, namely Sowerby’s Whale (JZ. bidens)

and M. ewropeus Gervais, the validity of which has been the

subject of much dispute, but which, from the evidence recently

given by the late Mr. F. W. True (1910, p. 11), has considerable

claims to be regarded as distinct from J. bidens.

The adult males of Ziphioid Whales usually possess one or two

pairs of large and conspicuous teeth, either at the extreme

anterior end of the lower jaw or further back in the same jaw ;

and in most of the species these teeth form a conspicuous feature

of the animal in the flesh. In young specimens of either sex

and even in adult females, the corresponding teeth do not cut

the gum, in most of the species; and a living specimen thus

appears to be completely edentulous. While the adult male of

the Common Bottle-nosed Whale (Hyperoodon rostratus) is

characterised by its remarkably swollen forehead, this feature

is absent from the females at all ages, as is shown in the figure

given by Capt. David Gray (1882, p. 728). When it is remem-

bered that the coloration of the skin is very variable, both in

Hyperoodon and in Ziphius, it will be realised that the external

differences between apparently edentulous specimens of these

two genera are not so striking as to preclude mistakes in deter-

mination by observers who have not had considerable experience.

In order to obtain any certainty with regard to the determination

of Ziphioid Whales it is thus desirable to scrutinise most care-

fully the evidence relating to reputed ‘ Bottle-nosed Whales.”

The considerable number of records of the occurrence of

Ziphius in the most widely separated localities, taken in conjunc-

tion with the evidence supplied by some of the best authorities

that but a single recent species has been proved to exist, leads to

the conclusion that 7. eavirostris is a cosmopolitan species which

inhabits the open oceans of the world and is oceasionally stranded.

It does not follow that it is in reality a rare animal. If the

conelusion that but one living species occurs be correct, the

species has an almost world-wide distribution, since it has been

recorded not only on both sides of the Atlantic and in the

Mediterranean, but also in such widely separated localities as

South Africa, New Zealand, and Bering Sea.

In 1912 the Board of Trade issued instructions to Receivers of

Wreck to inform the British Museum of the stranding of Cetacea

* Tt is not impossible that two living Whales which were observed from the cliff

at Great Saltee Island, Co. Wexford, June’ 15, 1913, by Mr. W. P. Pyeraft, in

company with the late Mr. R. M. Barrington and Mr. W. W. Grant, may have

belonged to Z. cavirostris. Mr. Pycraft has kindly given mea copy of the notes

which he made at the time, according to which all three observers were struck by

the remarkably white appearance of the fore part of the body, and were agreed that

this was not due to the effect of brilliant sunshine reflected from a black surface.

The hinder part of the body was evidently darker than the front part. It will be

noticed’ that this account is in complete agreement with the observations recorded

below on the coloration of the Wextord Ziphius stranded on July 19, 1915.

CUVIER'S WITALE. dbl

along the British Coasts; and as the result of this action a

number of telegraphic intimations of such occurrences have been

sent to the Museum from time to time. The telegrams thus

despatched have in many cases been supplemented by written

Reports, sketches, or photographs; and often by the transmission

of lower jaws or other parts of the specimens stranded. From

the information thus obtained two Reports have already been

published by the Trustees of the British Museum (Harmer,

1914, 1915). For the assistance given to this enquiry by

Receivers of Wreck and Coastguard officers I wish to express

my most cordial thanks; and it is hardly necessary to add that

a similar expression of gratitude is also due to the Board of

Trade, by whose action the collection of this information has

been rendered possible.

In receiving these telegraphic Reports the possibility of adding

to our knowledge of the rarer Ziphioid Whales has throughout

been borne in mind; and in particular it was hoped that oppor-

tunities would be aftorded of obtaining specimens of Ziphius

cavirostris. The inauguration of the scheme was more successful

than was suspected at the time; since the very first specimen

received after it came into full working order, namely the Whale

recorded in my 1914 Report as 1913, No, 1 (Unionhall, Co. Cork),

ultimately proved to bea specimen of the wished for Ziphius.

This discovery was only made a few weeks ago, on removing the

skeleton from the sand-pit in which it had been cleaned * ; the

specimen having at first been determined, on the evidence of its

lower jaw, as a /Typeroodon.

On July 19 of the present year a telegram was received from

the Coastguard officer at Fethard, Co. Wexford, announcing the

stranding of a Whale, said to be 19 feet long and to have two

teeth at the extremity of the jaw. It was supposed that this

animal would prove to be a Common Bottle-nosed Whale

(Hyperoodon rostratus) ; but the lower jaw was asked for in order

to render its determination certain. On the arrival of the jaw,

the pair of large and massive teeth at its anterior end showed

at once that the animal was not a Common Bottle-nosed Whale,

and it was more than suspected that it would prove to be a

Ziphius cavirostris. Mr. A. H. Bishop, one of the preparators

of the Museum, was accordingly sent to Fethard; and he was

fortunately in time to secure the remainder of the skeleton, as

well as to make observations on its external characters and to

prove that it wasa male. By a happy coincidence the skeleton

of the 1913 specimen already alluded to was at this time removed

from the sand; and the characters of its skull proved beyond

doubt that, like the Fethard specimen, it was a Ziphius cavirostris.

The British Museum is thus in possession of two Irish skeletons

(the Unionhall specimen not quite complete) of this interesting

* For this method of cleaning skeletons, specially to be recommended for Cetacea,

since it enables very Jarge specimens to be dealt with without trouble, and moreover

removes the oil from the bones, see R. F. Scharff, ‘The Museums Journal,’ x. 1911,

p. 196.

562 DR. S. F. HARMER ON

Cetacean. When the skeleton of the Wexford specimen has

been cleaned I hope to be able to publish a further account of

these two animals, in conjunction with my friend Mr. W. P.

Pycraft; but the records are interesting enough to justify the

publication of a preliminary notice on the subject.

The Wexford specimen was stranded on July 18, 1915, at the

entrance to Bannow Bay, on the east side of the point of land

separating that Bay from Waterford Harbour; and it was alive

when first observed. Jam indebted to Mr. Dennis McCarthy,

of H.M. Coastguard, for valuable information with regard to its

appearance, as well as for the trouble he took in facilitating the

acquisition of the specimen by the British Museum. In a

written description sent after the despatch of the original tele-

gram, Mr. McCarthy states that the lower jaw ‘was narrow

and projected beyond the upper.” The head was ‘“ mostly white

in colour and along the upper part of the back as far as the fin

with numerous whitish streaks running downwards in all direc-

tions; the remaindera darkish colour.” Thisaccount is confirmed

by Mr. Bishop, who made an excellent drawing of the animal,

based on his photographs, measurements and notes, on his return

to the Museum. This drawing represents the head, including

the whole of the lower jaw, and part of the back as white in

colour, the rest of the skin being black, or at least dark. The

white colour extends over the whole of the head and part of the

body, im front of an oblique line drawn from the anterior end of

the dorsal fin, and passing in front’ of the pectoral fin, to the

posterior end of the lower jaw. The colouring of this specimen

is thus extremely similar to that of the New Zealand animal

figured by von Haast (1880, pl. xxii.). Much of the skin was

covered by long, linear streaks, similar to those usually seen on

the skin of Granpus griseus, According to the observations of

Mr, Bishop, who did not see the speeimen until July 23, when

much of the epidermis had been lost, the streaks were arranged

singly. In the specimen figured by von Haast the skin is said

to have been marked with parallel, elongated streaks, in pairs,

as well as with oval scars, each with two dots in the centre.

These dots, and the two members of each pair of streaks were a

constant distance apart, and were “ evidently made by the teeth

of other individuals of the same species.’

One of the most striking features of the Wextord specimen

was its possession of a pair ‘of lar ‘ge and massive teeth, situated

at the extreme end of the lower jaw, the left tooth being appre-

ciably larger than its fellow. The exposed parts of these teeth

are roughly conical, the teeth diverging from one another and

being 20 mm. apart at their base. Other measurements of the

exposed parts of the teeth are :—

ee Right. Left.

Antero-posterior diameter, at base... 32 mm, 36 mm.

Transverse diameter, at base ......... 29 32

Te Gl ears rete me PE Niants 365 ca UR ai! 42

CUVIER’S WHALE. 563

- The larger tooth thus has a diameter of nearly one and a half

inches, while the part which projects beyond the gum exceeds

one and three-fifths inches.

The teeth just described have a close resemblance to those of

the specimen from the Chatham Islands, described and figured

by Hector (1872, pl. v.) under the name of Hpiodon chathamensis,

now usually regarded as a synonym of Z. cavirostris. They are

also precisely similar to those of a specimen, No. 21248, from

Bering Island, described and figured by True (1910, p. 52,

pl. xxii. fig. 4) and regarded by him (pp. 31, 35) as probably

belonging to an adult male. It is important to notice that the

Wexford specimen was definitely ascertained to be a male, from

its characters in the flesh; and it thus confirms the accepted

conclusion that the teeth of Ziphius are larger and more massive

in the male than in the female. In the paper just cited (p. 54)

True states his conclusion that adult males have ‘“ fusiform teeth

with closed roots and a diameter of from 25 to 30 mm.,” while

in females they are “quite slender, with a diameter of from 10

to 14 mm.”

The specimen* from Unionhall, Co. Cork, was stranded on

Feb. 13, 1913; and it was described as much injured, being in

an advanced state of decomposition, with most of its skin worn

away. There are thus no observations to record with regard to

its colour or sex. Its length is said to have been 20 feet. The

lower jaw had a considerable resemblance to that of Hyperoodon ;

and, as there were no teeth piercing the gum, it was supposed

that it belonged to that genus. The examination of the skull,

two years later, at once corrected this mistake. The absence of

the large maxillary crests which are so characteristic of Hyper-

codon is alone sufficient to show that it cannot belong to that

genus; while as features specially distinctive of Ziphiws and

possessed by this specimen, mention may be made of the peculiar

. form of the premaxille and the great difference between those of

the two sides, as well as of the elongated nasals, separated from

one another by a long suture. The length of the skull is about

3 feet 3 inches; the Shetland skull described by Turner (1912,

p- 77) being 364 inches long.

In his recent elaborate account of the cranial characters of

Z. cavirostris, True (1910, p. 54) states his belief that the two

sexes of this species are distinguishable from one another by

certain well-marked cranial characters, in addition to those

afforded by the teeth, as noted above. He admits, however,

that the sex of some of the specimens from which his conclusions

were drawn was not definitely known. The adult male was

believed to be distinguishable by the great development of the

“‘mesorostral ossification ” (Turner) and by the presence of a

deep “prenarial basin.” In the adult female the mesorostral

ossification is less developed, while the premaxille are narrow,

* For assistance in obtaining the skeleton of this specimen, the Museum is in

debted to Mr. J. Phelan, of H.M. Coastguard, Unionhall.

564 DR. S. F. HARMER ON

and flat proximally (pp. 35, 36), and the prenarial basin is un-

developed. It appears to me that these conclusions are well

founded; and applying them to the Unionhall specimen, there

seems every reason to believe that it was a female. ‘his con-

clusion is strengthened by the fact that the teeth of the apex of

the lower jaw do not cut the gum, although the dissection made

by Mr. Pyeraft proved that they are present and of considerable

size, projecting for about 25 mm. beyond the bone of the jaw

and nearly reaching the surface of the gum, and having a basal

diameter of about 13°5.mm. It should be added, however, that

the Unionhall specimen was not fully adult, since the sutures of

the skull are still extremely distinct, while the epiphyses of the

vertebre are not yet ankylosed to the centra.

It would be extremely desirable to be able to state characters

by which a Ziphius in the flesh can be distinguished certainly

from the other Ziphioid Whales; but Lam not prepared to do

this without further study of the subject. In the case of the

males, in their adult condition and probably at earlier stages of

their life, the occurrence of a pair of large teeth at the extreme

front end of the lower jaw and cutting the gum is probably

amply sufficient for recognition, particularly when taken in con-

junction with the absence of the enormously swollen forehead

so characteristic of the adult male Hyperoodon. But a young

Ziphioid Whale of either sex, or an adult female which has no

teeth visible during life, is less easily referred to its proper

genus.

It has already been pointed out that the Wexford specimen of

Z. cavirosiris and von Haast’s New Zealand specimen (1880,

pl. xxi.) were white above and dark below—a type of coloration

which is by no means common in Cetacea. Other specimens

which have been referred to this species were, however, dark

above, even on the head, and light below. Making full allowance

foy the uncertainty which so often prevails with regard to the

real colour of Whales, owing to post mortem changes and to the

fact that the examination frequently has to be made some time

atter death and under unfavourable conditions, it must be con-

cluded, in the present state of the evidence, that 7. cavirostris is

a species of very variable coloration (cf. True, 1910, p. 35). It

may further be noted that according to Van Beneden (1888,

p- 60) the rostrum and forehead of Hypereodon become white in

colour with age, The projection of the lower jaw beyond the tip

of the snout is apparently a positive character of Ziphius ;

and the snout is probably less distinctly marked than in either

Mesoplodon or Hyperoodon.

A further difficulty in defining the external characters of

4. cavirostyis arises from the uncertainty which prevails with

regard to the two mandibular teeth in the female. In some of

the published accounts of this sex, these teeth are described as

visible during life ; while in the Unionhall specimen (probably a

female) they were completely concealed beneath the gum.

CUVIER’S WHALE. 569.

It is hardly necessary to attempt to give a full account of the

literature of the subject in this preliminary paper, but it may be

convenient to refer to the lists of recorded occurrences given by

Turner (1872, p. 770), Flower (1872, p. 207), Van Beneden

(1888, p. 86), and True (1910, p. 30); the Memoir last cited

giving far the fullest account of the general characters of the

species ati present published, and being accompanied by excellent

figures of skulls and other parts of the skeleton.

With regard to published figures of the entire animal it may

be noted that some of the earlier representations are so different

from one another that, in the absence of other evidence, it would

be hard to believe that they all represented the same species.

Attention may, however, be directed specially to the following

published illustrations :—

A Spanish specimen figured by Cabrera (1914, p. 380), who

records three specimens from Santander, Bay of Biscay ;

Specimens respectively from Alaska and Newport, Rhode

Island, of which photographs are reproduced by True (1910,

Pip exlia ties dau4))s

It may be noted finally that whether all recent specimens of

Ziphius ave referable to a single species or not, there is every

reason to believe that the Irish specimens belong to 4. cavirostris ;

a conclusion reached with regard to the Shetland skull by

Sir William Turner. This species was named by Cuvier (1823,

p. 352) on the evidence of a skull obtained from the Bouches-du-

Rhone, at first regarded as a fossil specimen, but shown by later

writers to have been really that of a recent animal.

Memorrs Crrep.

1888. BenEeDEN, P. J, Van.—“ Les Ziphioides des Mers d’Kurope.”

Mem. couronnés Acad. Roy. Belgique, Coll. in 8vo, xh.

Mem. No. 2.

1914, Caprera, A.— Fauna Ibérica, ‘‘ Mamiferos,” Madrid,

p: 376.

1823. Cuvirr, G.—Recherches Ossemens Fossiles, Nouv. Ed. v.

1r¢ Partie, p. 352.

1872. Frowrr, W. H.—‘‘On the Recent Ziphioid Whales.”

Trans. Zool, Soc, vii. 1874, p. 207.

1882. Gray, Davip.—‘t Notes on the Characters and Habits of

the Bottlenose Whale (Hyperoodon rostratus).” Proc.

Zool. Soc. p. 726.

1880. Haast, J. von.—‘‘On Ziphius nove-zealandie.” Proc.

Zool. Soc. p. 232.

1914. Harmer, S. F.—‘“‘ Report on Cetacea stranded on the

British Coasts during 1913.” Published by the Trustees

of the British Museum.

1915. Harmer, S. F.—‘ Report on Cetacea stranded .... during

1914.” Lhid.

566

1872

1910

ON CUVIER’S WHALE.

. Hecror, J.—‘* On the Whales and Dolphins of the*New

Zealand Seas.” Trans. Proc. N. Zealand Inst. v. 1873, —

p. 164.

. True, F. W.—* An Account of the Beaked Whales of the

Family Ziphiide in the Collection of the U.S Nat. Mus.,

with Remarks on other specimens in other American

Museums.” Smithsonian Inst., U.S. Nat. Mus., Bull. 73

(Ato), pp. 1, 30.

1872. Turner, W.—‘‘ On the Occurrence of Ziphius cavirostris

WN,

in the Shetland Seas, and a Comparison of its Skull with

that of Sowerby’s Whale (Wesoplodon sowerbyt).” Trans.

Roy. Soc. Edinb. xxvi. p. 759.

TuRNER, Sin W.—‘‘ The Marine Mammals in the Anato-

mical Museum of the University of Edinburgh,” p. 77.

ON A NEW NEMERTEAN, 567

39. Fauna of West Australia.—IIT. A new Nemertean,

Geonemertes dendyi, sp. n., being the first recorded

Land Nemertean from Western Australia. By W. J.

Dakin, D.Se., F.Z.S., Professor of Biology in the

University of W. Australia.

[Received August 5, 1915: Read November 9, 1914. }

(Text-figure 1.)

INDEX.

SYSTEMATIC ; Page

Geonementes dendiya, Sp. seesueseseaccescacteeeeceees | DOT

SDRUCDUR Beas ascot eae tae etic cae eae enema reer OOO.

Land nemerteans are notoriously rare animals, and it is there-

fore particularly interesting to record a new species from the

Western State of Australia. The record is interesting too, because

the other known species in Australia comes from Victoria and New

South Wales, about two thousand miles distant from this western

locality. The country between can scarcely be called suitable tox

the distribution of such an organism, ‘The animal was discovered

by the author, whilst searching for Peripatus, in a valley in the

Darling Range not far from Perth. Land nemerteans are cryp-

tozoic in habit and occur in the same situations as Peripatus and

Jand planarians, yet no specimens have previously been discovered

in West Australia, although many scientists have made collections

of these latter Cryptozoa. I, myself, have looked for Peripatus and

land planarians in the Darling Ranges, and other parts of West

Australia, on very many occasions without ever meeting with a

specimen of Geonemertes. ‘his first record does not indicate any

greater abundance, for only one isolated individual—a mature

female—was found. It is probable, however, that in the keen

search for Peripatus (when the attention is concentrated on

distinguishing this animal from its background) specimens of the

nemertean have been passed over as land planarians. Such, in

fact, would have been the case this time, if the animal had not

protruded a long proboscis on being disturbed.

The example belongs to the genus G'eonemertes, and I have

much pleasure in naming the species after Professor Dendy, who

was not only the first to discover land nemerteans in Australia

and New Zealand, but who elucidated many points in the anatomy

of the genus. The previously recorded species from Australia

and New Zealand are Geonemertes australiensis Dendy, occurring

in Victoria, New South Wales, and Tasmania; and Geonemertes

nove-zealandiw Dendy, a very rare species occurring in South

Island, New Zealand. The new form G'. dendyi is more like

G. australiensis than the New Zealand species.

Habitat.—The specimen was found under a small log in a

rather damp situation, about two yards from a small stream, in

568 : PROF. W. J. DAKIN ON

one of the valleys near Armadale. Under the same log were

two specimens of Peripatus gilesii, a slug, and some of the usual

small cryptozoic arthropoda. Scutigera was also common in the

neighbourhood.

External characters.—Geonemertes dendyi is apparently much

smaller than the Hast Australian species, for its total length when

living and uncontracted was only 15 mm. (proboscis retracted).

The greatest breadth occurred at a point about one third of the

length from the posterior end. Just in front of this was a

shght constriction, as if the animal had been nipped. It is

probable that this feature is only some temporary or individual

character of the specimen captured, but as no others are to hand

for purposes of comparison, it is worth mentioning.

Text-figure 1.

Geonemertes dendy?.

A. Dorsal view. B. Anterior end considerably enlarged.

The colour of the living animal is brown-pink, but the shade

is not uniform over the entire surface. The lateral parts of the

body are more of a light flesh-colour and signs of the large ova

were visible, showing through the skin. The under surface is

pale. On the dorsal surface, and most prominent on the posterior

third of the animal, are two dark stripes of chocolate-brown.

They are not sharply defined, and the dissecting-microscope

indicated that they are collections of little brown spots. There

is just a faint indication of the continuance of the stripes forward

over the anterior part of the dorsal surface.

Close to the anterior end of the body, and on the dorsal surface,

are the eye-spots. According to Dendy, Geonemertes australiensis

differs from other known species of Geonemertes in the possession

of a large number of eyes. The New Zealand species has only

four eyes, and four or six are the usual numbers. In Dendy’s

specimens from the eastern states the eyes numbered as many

as thirty or forty, and they were arranged in two groups. It is

A NEW NEMERTEAN. 569

interesting to note that the author describes each group (con-

taining about 20 eyes of various sizes) as sometimes showing

indications of a division into an anterior and a_ posterior

group, and he says that “it suggests that the numerous eyes

of G. australiensis may have been derived by subdivision of four

eyes, two large anterior and two small posterior, such as we find

in G. chalicophora.’

In Geonemertes dendyi there are four quite distinct groups of

eyes—two anterior groups of large eyes, and two posterior groups

of smaller eye-spots. There are five or six spots in each of the

anterior groups and three in each of the posterior, making a total

of sixteen.

ANATOMY.

The anatomy of the animal as made out from serial sections

does not differ in any points of importance from that of @. aus-

traliensis. Unfortunately, the animal was fixed in an acid fixative

and consequently it is impossible to see anything of the calcareous

stylets.

The epidermis and basement-membrane are both of the usual

type. There are no rod-like bodies, and calcareous bodies like

those deseribed by Dendy in G@. austr aliensis are not to be seen.

This, however, is no proof of their absence, for Dendy was never

able to find them in his sections although no acid fixative was

employed.

Within the basement-membrane are two layers of muscle- fibres,

but judging from the figures the thickness of the layers is not

so great as in G. australiensis. The outer sheath is of circular

fibres, the inner of longitudinal muscle-fibres. If a layer of

diagonally disposed muscle-fibres exists between these two sheaths,

it is not evident in the transverse sections. The muscular dia-

phragm, described by Dendy asa development in the cephalic

region, 1s well seen in the present species.

The alimentary canal exhibits no new points. In the specimen

sectioned the lumen of the canal is almost obliterated by masses

of protoplasin such as have been described by von Graff as

occurring under certain conditions in G. chalicophora. The mouth

opens into the rhynchodeum as in G. a@ustraliensis and the New

Zealand species. The position of the opening is in front of the

cerebral ganglia.

The Late ciliated pits are

to be found on the ventral surface near the anterior end of the

body. The ducts pass almost vertically upwards toward the

cerebral ganglia and then turn rather abruptly and run toward

the sides of the body. Cilia can be traced in these ducts for a

considerable distance—they are to be seen where the ducts are

quite close to the ganglia. There is little to add further in con-

nection with these, except that the ducts come into rather intimate

connection with a curious mass of tissue lying ventrally and

slightly posteriorly to the ventral lobes of the brain. This is

probably what Dendy calls the ‘‘ esophageal organ.” The tissue

570 ON A NEW NEMERTEAN.

is most certainly non-nervous. It stains an intense blue with

hematoxylin and has all the characters of glandular tissue.

Cephalic Gland.—One of the most curious differences between

G. dendyi and G@. australiensis is the apparent lack of a well-

developed cephalic gland. I must confess that I cannot recognise

any structures in my sections which seem to fit in with the deserip-

tion given by Dendy. The sections were stained with hematox-

ylin, and glandular structures are well brought out. The dorsal

glands are well developed. and agree in position with those

described as occurring in G*. australiensis, but there are no other

distinctly ‘“ large, irregular, glandular masses, closely packed

together—staining deeply with hematoxylin” overlying the

dorgal lobes of the cephalic ganglia. The only well-developed

glandular structures in this position are the anterior glandular

masses of the dorsal glands.

Reproductive Organs.—The single specimen so far known is a

mature female, and there is no trace whatever of male organs.

The sexes are also separate in G. australiensis. In the classifi-

cation offered in Benham’s treatise (Treatise on Zoology, Ed. by

KE. R. Lankester, Part iv.) the genus Geonemertes is placed in the

family Prosorhoemide : ‘‘ With four eyes, cerebral organs are

rudimentary. Cephalic gland large. Mouth and rhynchoccel

coincident. Usually hermaphrodite.” If G. dendy? is consicered

as one member of this family, the diagnosis of the latter requires

to be made a little more general.

In @. dendyi the ova are found throughout almost the entire

length of the body—the most anterior ones occurring just pos-

terior to the brain-masses. They are present in different stages of

growth, but most of them are very large and apparently mature.

Curiously enough, I cannot detect the genital ducts which are

figured so distinctly by Dendy. In one or two places there are

signs that might be interpreted as remains of these ducts. If

one did not know, however, that such structures did exist in the

genus, they would never be suspected from my sections.

SUMMARY

The characteristic features of Geonemertes dendyi, sp. n., are as

follows. The length of the mature female is about 15 nm. when

crawling. The colour is brown-pink, with two dorsal and some-

what posterior darker longitudinal bands of chocolate-brown.,

The eyes are arranged in four groups —two anterior, each of five

or six larger spots, and two posterior groups, each of three smaller

eyes, making about sixteen or seventeen altogether. Lateral

organs are well developed, opening by characteristic ciliated

cephalic pits on the ventral surface at the anterior end. The

mouth opens into the rhynchodeum. The sexes are distinct.

Cephalic gland apparently not well developed. Other structures

agreeing well with the description of similar parts in G. austral-

iensis. The specimen was found under a small log, together

with Peripatus gilesii, in the Darling Hills, Western Australia,

not far from Perth.

IAS. WSIS, VAIN, Il 1.

W.J.D.del. Cambridge University Press.

PALAMONETES AUSTRALIS.

ON A NEW CRUSTACEAN. All

40, Fauna of West Australia —IV. Palemonetes australis,

sp. n., being the first record of the genus in Australia.

By W. J. Daxin, D.Se., F.Z.S., Professor of Biology

in the University of W. Australia.

[Received August 5, 1915: Read November 9, 1915. ]

(Plate I.*)

INDEX,

SYSTEMATIC: Page

(PalEinOneteEs @USEFALISyiSpe iv seurereeeedee os ees OL

With the exception of the Phyllopoda and the crayfishes, few

aquatic animals appear to have been collected and recorded from

the fresh waters of Western Australia.

Among the so far unrecorded species is a prawn-like crustacean,

which is extremely cominon in many of the rivers near Perth.

What its actual range in the continent may be, remains to be

discovered. The species was first found by the author in a tow-

net which had been thrown at random froma river-bank and

pulled back ; the net just scraped the bottom on the way. About

fifteen specimens were caught in this very short, careless haul,

and as the animals are pretty active in their movements they

must have been present in large numbers in the water.

Since the first discovery, specimens have been obtained from

several other districts. All proved on investigation to belong to

the genus Palemonetes. ‘This new record marks a considerable

increase in the known geographical range of this genus. So far

as I can determine, only one species of Palemonetes is known to

occur in Kurope, but that is found in Sweden, Denmark, Belgium,

France, Italy, Spain, and the Black Sea. It has also been collected

in the ‘British Islands and even so far south as Egypt. This

species, Palemonetes varians, lives in water that is more or less:

brackish and close to the sea, as well as in the perfectly ores

water of lakes and rivers.

Most of the known species of the genus Palemonetes appear

to be American—the following having being recorded. Palaemo-

netes vulgaris (sea-water—bays and estuaries of U.S.A. coast),

Palemonetes exilipes Stimpson (fresh-water—U.S.A.), P. carolinus

Stimpson (marine—U.S.A. coast), P. argentinus Nobili (South

America), P. kadiakensis Rathbun (North America), P. calcis

Rathbun (blind species found in caves in Cuba), P. antrorum

Benedict (blind species from an artesian well in Texas), P. eigen-

manni Hay (blind species from caves in Cuba). The Rev. T. R.

R. Stebbing writes me that he has described and figured a species

* For explanation of the Plate see p. 574,

572 PROF. W. J. DAKIN ON

from Natal, sothat South Africa has also a representative of the

genus. It is surprising to find that, on the whole, there are only

slight differences between the Australian species and P. varians

from Europe or the species known from the American continent.

Habitat in West Australia.—Specimens of P. australis were

first met with at Northam, a small town inland on the plateau,

about 70 miles from the coast and at an elevation of roughly

500 feet. They were captured in the River Avon, where it breaks

up into small channels below the weir. The water is of course

quite fresh. The largest specimens obtained have been caught

at the above place, and the animal is quite plentiful at all seasons

of the year. Large-sized specimens have also been captured in

Gin Gin Brook, about 50 miles north of Perth and at an elevation

of roughly 300 feet. Mr. W. B. Alexander, M.A., of the West

Australian museum, has placed in my hands collections captured

in Bibra Lake and the Serpentine River. The lake is a small

expanse of water only a foot or so deep, if that in the dry season,

and is situated on the coastal plain not so very far from the sea.

The habitat of these lake specimens is very different from that of

the Northam examples, and there is a characteristic difference

in the size of the individuals, those from the Northam River

being the larger. The average length of the individuals caught

at Northam is 32 mm., against 20 mm. for the Bibra Lake type.

The Serpentine River is about 34 miles south of Perth, and

the specimens were caught where the river leaves the hills and

enters the coastal plain. The specimens from Northam are

described and figured as the type specimens of Palcwmonetes

australis.

DESCRIPTION,

Body stout. Length of largest specimens 39 mm. from end of

telson to tip of rostrum, and 72 mm. to tip of outstretched antenne.

The living animals are transparent, with a prevailing amber-green

tint. Brown pigment flecks occur laterally on the posterior

margins of the abdominal terga, and there are sometimes two

delicate longitudinal lines on the carapace somewhat laterally

situated. The carapace is not quite so long as the last three

abdominal segments. ‘The sixth abdominal segment is almost

twice the length of the fifth.

Rostrum.—The rostrum is long and about equal in length to the

carapace, but may be very slightly longer or shorter. The tip of

the rostrum exceeds the distal margin of the antennal scales.

The rostrum is laterally compressed and has a _ pronounced

curvature trending upwards towards the apex. A few small

chromatophores are present. The dorsal armament consists of

usually five or six teeth which are almost equally spaced along

the entire length of the rostrum. This is without counting a

distal tooth which forms the upper portion of the bifid apex.

The most posterior dorsal tooth is well behind the orbital notch,

and the next one is immediately over it. There are three or four

A NEW CRUSTACEAN. 573

ventral teeth. The dorso-ventral thickness of the rostrum is not

so great as that of P. varians.

Hyes.—The eyes are well developed and on fairly long stalks.

Antennules.—The antennules are as long as the abdomen with

telson. ‘The peduncle when extended does not attain the length

of the rostrum by about one-third of the latter, and falls short of

the distal margins of the antennal scales by about one-fifth the

length of the scales. The shorter ramus of the outer antennule

is fused to the longer for about half its length.

Antenne.—The antennal scales are large, long, and broad, and

more than twice the length of the antennal peduncle minus the

first segment. ‘They are not quite so wide distally as proximally.

The antennal peduncle falls short of the first segment of the

peduncle of the antennule.

Mandible.—The mandibles are without palps. Incisor and

molar processes are well developed and tipped as indicated in the

figure.

Maxillule and Maxiile.—These appendages are as figured.

They possess no features of systematic importance marking them

off from the similar appendages of P. varians.

Mazxillipedes.—These are similar in structure to those of P.

varians. The third maxillipedes when extended reach approxi-

mately to the end of the antennary peduncle.

Percopods.—The second peropods are the longest, and when

extended they overlap the antennary scale by palms and fingers.

The third perzopods are about as long as the first, the fourth a

little longer, and the fifth pair are the longest of the posterior

three. The first pair of perzeopods attain almost the apex of the

antennary scales when extended forwards. The chele of the

second pair are shorter than the carpus, and the dactylus slightly

more than two-fifths length of palm.

Telson.—The telson is longer than the preceding segment. It

terminates in the mid-line with a spine. Thearmament consists

of two pairs of stout spines and one pair of sete. The latter are

situated one on either side of the median spine. The posterior

margin of the telson differs distinctly in shape from that of

P. varians. Two pairs of spines are present on the dorsal surface

of the telson.

CONCLUSION.

Palemonetes australis occurs in the fresh-water of rivers some

distance from the coast in West Australia, and is also found in

shallow lakes on the coastal plain.

Females bearing eggs which have been just extruded have

been captured in September (Bibra Lake), and others bearing

embryos not far from the hatching stage have been collected

on January lst from Gin Gin Brook. This would indicate that

the breeding season coincides with the early months of the

summer—the dry season. Most of our West Australian fresh-

water animals breed in the winter or spring, and many possess

Proc. Zoou. Soc.—1915, No. XL. 40

574 ON A NEW CRUSTACEAN.

drought-vesisting eggs. Unfortuately, newly hatched larve

have not yet been captured, so that we are unable to figure this

stage. :

The species of Palemonetes known at present differ but slightly

from one another, and the characters of most importance

systematically which mark the Australian species are (a) rostrum,

(6) rami of antennule, (c) length of pereopods and_ their

segments, (d) telson.

EXPLANATION OF THE PLATE.

Palemonetes australis.

Fig. 1. Rostrum (form with seven dorsal spines). X 9.

2. Outer antennular flagellum (basal portion).

3. Antennal scale and base of antenna. 8.

4. Mandible. X 20.

5. First maxilla. X 12.

6. Second maxilla. X 12.

7-9, First, second, and third maxillipedes. 12.

10. Second perwopod. X 7.

11-18. Third, fourth, and fifth pereopods, x 3.

14. Telson, 12,

ON THE TASMANIAN DEVIL. 575

41. The Keeping and Breeding of Tasmanian Devils

(Sarcophilus harrist). By Mrs. Mary G. Ropsrts,

CMEZ_ Sa MAReAROE IU):

[Received June 21, 1915: Read October 26, 1915.)

(Text-figure 1.)

JeNieae dl

Until I was asked by Mr. A. 8. Le Souéf, Director of the

Zoological Gardens, Moore Park, New South Wales, early in 1910

to obtain, if possible, Tasmanian Tigers (7hylacinus cynocephalus)

and Devils (Sarcophilus harrist) for the London Zoological Society,

I had never thought of keeping either of these animals in my

collection ; in fact, they were quite unknown to me except as

museum specimens, although I had frequently visited remote

parts of our island. I have vivid recollections, however, of how,

when a young child at boarding-school in the late forties, some

of the girls trom Bothwell, near the Lake District, used to give

graphic and terrifying accounts of the Tasmanian Devils with

their double row of teeth. This belief is not yet exploded, as it

was impressed upon me lately with the utmost confidence by a

country visitor that such was the case; he not only believed, but

said “he had seen.” ‘The teeth have been described to me by a

scientist as truncated.

Shortly after hearing from Mr. Le Souéf, by means of adver-

tising, writing, etc. I obtained three for the London Society, and

having then become thoroughly interested I determined to keep

some myself. Since that time a large number have passed

through my hands, and more than once I have been “a woman

possessed of seven devils.”

In April 1911 I received a family (a mother and four young),

and again in September of the same year a similar lot arrived.

The former were very young, and | had the opportunity of

watching their growth almost from their first appearance when

partly protruding from the pouch. When sending them, the

trapper wrote that *‘the mother was so quiet, I need not be afraid

to pick her upin myarms.” The little ones hung from her pouch

(heads hidden in it), and she lay still and motionless as if afraid

of hurting them by moving, and allowed me to stroke her head

with my hand. However timid they may be, and undoubtedly

they are extremely so, growling and showing their teeth when

frightened, they always evince this gentleness and stillness when

nursing little ones.

The skin of the young, on arrival, had the appearance of a

slate-coloured kid glove, the tail darker towards the tip. The

hair could be seen growing black and velvety from the head

downwavds, the latter being hidden in the pouch for some days,

40*

576 MRS. M. G. ROBERTS ON

and it was interesting to note the progress of the growth of the

hair from day to day. The shoulders were covered while the

hind-quarters were almost, or quite, bare, although a faint streak

of white was discernible where the white markings were to come

later on. At this early stage, should the mother get up to

move about, which she rarely does in the daytime, the young

somehow scramble into the pouch again.

This family went later to the London Society, but the second,

which came on the 16th of September, I kept for my own pleasure,

with the exception of the mother; as she had lost a foot when

being trapped, I thought it best to have her destroyed later on.

Unfortunately, when they were about half grown one escaped into

the garden, and the next morning her mutilated remains were

found—she had fallen a victim to our two fox-terriers. The three

survivors have been ever since an unfailing source of interest

and amusement to my family, to visitors, and myself. When a

bone or piece of meat was thrown to them a tug-of-war was

always the result, and sometimes a chase into one door and out

of the other of the little cave. At other times, while one has

been holding on to a bone held in my hand, I have lifted it

completely off the ground, while another would cling on round

the waist and try to pull it down.

Many visitors from the Commonwealth have heard such exag-

gerated accounts of the ferocity and ugliness of the Tasmanian

Devil (others, again, have believed it to be a myth), that they

sometimes express surprise when they see them so lively, sprightly

and excited, running out to my call; they then remark, “the

devil is not so black as he is painted.”

Two of these Devils were latterly kept together as a pair, and

for the purposes of this article I will call them Billy and Truganini,

after the last two survivors of our lost Tasmanian race. These

showed no disposition to breed until April 1913, and my obser-

vation of them and of many others that | have had in my keeping

is, that the disinclination to take up maternal duties is always on

the part of thefemale. I then noticed suddenly a decided change—

that Billy would not allow her to come out of their little den ; if

she did venture when called to be fed, or at other times, he imme-

diately attacked her and would drag her back by the ear, or any

other part, but although otherwise cruel, he would carry food in to

her. When I called her, it was pitiable to hear her whining; but

it was of no avail, for Billy was a relentless tyrant and kept singe in

strict seclusion for quite ten or twelve days; then early in May he

allowed her to be free once more. From thence onward, although

they were sometimes peaceable and affectionate, the balance “of

power was completely on Truganini’s side; she constantly resented

his approach by biting and snarling at him: it seemed as if

coming events cast their shadows before, and she instinctively

felt that he would do the young some injury. From now her

pouch was anxiously scanned day by day, but it was some time

before I could be sure that it was gradually enlarging. I had been

THE TASMANIAN DEVIL. yall

advised by Dr. Hornaday, of the New York Zoological Park, that

if ever the Tigers or Devils were likely to have young, to remove

the male, and as soon as I was certain, I had Billy taken away

Text-figure 1,

Tasmanian Devils (Sarcophilus harrisi).

and placed with the other member of the family. This made

Truganini most unhappy, as he was near enough for her to hear

him, besides which, the two males fought; so, being cautioned by

78 MRS. M. G. ROBERTS ON

my family that perhaps my interference might cause a disaster,

I yielded and replaced him, doing so with many misgivings.

Matters went on much the same el late in September, when to

my delight a tail, and at other times part of a small body, could

be seen sticking out of the pouch, more especially when the

mother sat up to wash her face, or rolled upon her back; unlike

domestic cats, the devils use both paws for washing, placing them

together and thus making a cup-like depression which, when

thoroughly licked, is rubbed well over the face. Everything looked

very promising on the Sunday before Michaelmas Day, when I

noticed Truganini carrying large bunches of straw about in her

mouth, evidently seeking for a , retired place to make a bed, and

we had already placed some fern logs in a corner of their yard.

As Billy would follow her about and interfere, I had a box put

down with a hole cut in the side that she might hide under; but

it was of no use, as where she went he would also go, and a scrim-

mage was the inevitable result. Early next morning, with many

misgivings I left home for ten days, only to find on my return

that her pouch was empty and that the young had disappeared,

and as no remains whatever had been found, I could only conclude

that they had been eaten by Billy.

Thus ended ail my hopes and anticipations for 1913. I have

not so far related an incident that took place just before the

breeding-season. Being hopeful that Truganini might have

young in her pouch, and my assistant being as usual very busy,

Pr Basson ibe Se Flynn, of the Tasmanian University, who is

always interested in our marsupials, kindly offered to examine

her pouch. As soon as an attempt was made to catch her, Billy

grasped the position of affairs and fought to defend her with all

his might, even getting behind her in the little cave, putting a

paw on each shoulder and holding her tightly, lest she might get

into what appeared to him to be the danger zone. By dint of

perseverance and a little strategy he was outwitted at last, but

our hopes were doomed to disappointment.

Truganini has now passed through another period of retirement,

and Iam hoping to record shortly a greater measure of success

for 1914.

I cannot close this article without afew words in defence of the

Tasmanian Devil,as I am sure that it is more or less ‘‘ misunder-

stood,” and the article with photograph published in the ‘ Royal

Magazine’ for October 1913 under the name of L. R. Brightwell,

F.Z.8., is, I consider, greatly exaggerated both as regards their

appearance and character, viz., “‘'They are well named, for they

tear everything, even sheep, to pieces if they get the chance!”

On several occasions when one of mine has escaped, the only

mischief done has been the destruction of a fowl or a duck or two.

Tt would have been just as easy for a wallaby to have been killed

if they had had the inclination, about which our fox-terriers would

not have hesitated for a minute if a chance had occurred. When

in transit to London last year one escaped, and I have been told

THE TASMANIAN DEVIL. 579

by the chief officer of the vessel that ‘‘the passengers were much

alarmed as there were children on board, and someone went about

with a revolver.” Later I came across the butcher who was in

charge at the time, and he appeared to have been rather amused

than otherwise, and told me the missing one was discovered at

last quietly sleeping under the berth of one of the sailors! I

don’t wonder, with the reputation that the devils have, that the

passengers were alarmed.

Part II.

Having written so much in the first part about the keeping

and breeding of Tasmanian Devils, I fear I have not many fresh

facts to relate for 1914. The season arrived a month earlier, and

Billy released his little mate from retirement on the 2Uth of

March. She was just as disagreeable to him afterwards as she

had been on the former occasion, biting and snarling whenever

he approached her, and on the 8th of July I removed him from

the enclosure. There was nothing of importance to note until the

30th of the month, when a little tail was seen sticking out of the

pouch, and on the following day a foot and thigh were visible.

I will now give my observations on certain days following.

August 3rd—Little ones partly hanging out of pouch. J must

not forget to state that about this time, or a little later, Truga was

observed carrying bundles of straw about in her mouth, with

which to make her bed, and finally took them behind the fern logs

that we had thrown down, but unfortunately I omitted to note the

exact date. 4th, 5th, and 6th—Sometimes saw three tails only. .

7th—A little one lying on its back, feet in air and head in pouch.

On the 9th, for the first time, the man saw one standing alone on

a fern log, when it immediately scrambled down to the mother.

10th, 11th, and 12th—Mother frequently seen, sometimes with

only three tails observable, at other times little ones exposed,

bodies resting on the ground with the heads hidden. From

these observations, I may point out how difficult it is to know

exactly when the young are able to leave the pouch, it being

coincident, I think, with the making of the bed by the mother.

On the morning of the 13th Truga ran out to meet me, jumping

over fern logs, and left a little one whining behind, having the

tails of the other two and a foot showing outside the pouch ;

she went back at once to the young one, when it immediately got

on her back. 14th—Mother came out with two dangling from

her, leaving a little one behind crying; she at once ran back and

returned with all three in her pouch. Next day when she came

forward to meet me, only two tails anda foot could be seen.

16th—All three were hanging out, and instead of jumping over

ferns as usual, she had scraped away the straw and earth and

came out from an opening underneath them. By this time they

were getting quite covered with hair, white markings distinct, and

sometimes when disturbed they would make a faint attempt at

580. MRS. M. G. ROBERTS ON

a bark. From the 19th to the 23rd inclusive they were occasion-

ally seen all together out, yet on the three days following I

saw her about with two dangling from her. On the evening of

the 27th, upon her running out to meet me, I threw her some

meat, which she carried in to the young, afterwards returning for

more for them, and eventually lay down contentedly in front of

the opening. 29th—All three playing like puppies, biting each

other and pulling one another about by the ears. 30th—Whole

family hanging from the mother as she ran out, and one hardly

knows which to admire most, her patience and endurance, or

the hardihood of the young in holding on and submitting to so

much knocking about. The whole process seems very casual and

most remarkable, when compared with the breeding and rearing

of other marsupials. With the Kangaroo and allied types the

head is seen first, looking out of the pouch, and in the early stages

is quite bare. Sept. lst—Young ones playing in their little

corner. 9th—Not been seen this month in their mother’s pouch.

30th—Coming out all alone for pieces of meat and evidently able

to look after themselves. At the beginning of the New Year

IT removed them to other quarters and replaced Billy, much to the

annoyance of Truga; probably she resented the loss of her little

ones, and showed her anger by biting him severely about th

body and leaving various tooth-marks.

From observations made during the two seasons, I have come

to the conclusion that about four months and a half elapse

between the breeding-season and the time the young are able to

leave the pouch.

The baby devils had the sense of smell very strongly developed ;

immediately I approached, their nostrils would begin to work and

a vigorous sniffing would go on. ‘They were also expert climbers,

and although I had some specially constructed yards made, they

would get up the wire-netting and walk along the top rail quite

easily ; at other times they would climb a pear-tree growing in

their enclosure and sit in the branches like cats.

GENERAL REMARKS.

IT have always found devils rather fond of a bath; quite

recently, going down to their yard after an illness and finding

only a drinking vessel, I ordered a larger one to be put in, and

they showed their pleasure by going in at once, sometimes two

at a time. I have occasionally poured water from a can over

them, when they would run to and fro under it with much

enjoyment.

Their sight in daylight is rather defective; they seem to pick

up their food more readily by smelling than by seeing, and I

think they can see objects better at a distance.

At the present time I have six running together, my own three

and three that I bought when in their mother’s pouch. All are

tame, frolicsome, and lively. I can go in and havea bit of fun

THE TASMANIAN DEVIL. 581

with them, and when I am outside their enclosure they frequently

climb the wire-netting to the height of nearly six feet, and get

their little black faces close to mine with evident delight. We

have tried more than once to get them photographed, but it is

impossible to keep them quiet, they are on for a scamper all the

time. Recently an adult escaped, and it was discovered by a

passing school-boy sitting on a high fence bordering the street,

- under the shade of some elm-trees, many people passing on the

foot-path without observing it. They are, however, always very

timid when coming down.

They are fond of the sun, and look well when basking in it, the

rays shining through make their ears appear a bright red, fore

feet parallel with the head, hind-quarters quite flat on the ground

and turned out at right angles, somewhat as a frog.

My sympathy with my little black “ brothers and sisters ”’

intense, probably evoked by having suffered much mentally owing

to the gross cruelties which have come under my notice, the

result of capturing them in traps. Frequently three or four have

been sent to me in a crate, only to find later on one with a foot

shot off or a broken leg. In a consignment received some time

ago, a dead one was found; it bore unmistakable signs of a snare

previously, round the neck, one foot was gone (an old injury),

and finally a recently smashed leg much swollen, the cause of

death. I communicated with the 8. P. C. A., and since then

have had none from that district.

I have derived much pleasure from study ie the habits and

disposition of the Tasmanian Devils, and have found that they

respond to kindness, and certainly show affection and pleasure

when I approach them. I have been led to believe that no case

of their breeding in captivity has been recorded, and certainly

not in ‘Tasmania.

Others who do not know or understand them may think of them

as they like, but I, who love them, and have had considerable

experience in keeping most of our marsupials, from the Thylacine

down to the Opossum Mouse (Dromica nana), will always regard

them as first favourites, my little black playmates.

ON SNAKE-FEEDING. 583

42, Notes on the Feeding of Snakes in Captivity.

By E. G. BoutrnceEr, F.Z.S., Curator of Reptiles.

[Received October 8, 1915 : Read October 26, 1916. |

Some years ago, in a paper contributed to the ‘ Proceedings ’

of the Society by Dr. Chalmers Mitchell and Mr. R. I. Pocock,

entitled “The Feeding of Reptiles in Captivity” (P. ZS. 1907,

p. 785), a general account was given of the feeding habits of the

Snakes then living in the Gardens, together with records of a large

number of individual specimens, showing the amount consumed

between the months of May and October. My own general

observations tally with those of Dr. Mitchell and Mr. Pocock,

and my object in presenting this paper is to lay before the

Society some additional facts, to show how unnecessary it is to

feed the snakes on live creatures, and to give a detailed record of

the amount taken and of the regularity of the meals in a number

of specimens over a period of a year’.

Previous to my appointment at the Gardens I had fed the few

snakes kept by me on live animals, being under the influence of

the popular belief either that many snakes would not take dead

food at all, or that, at any rate, in most cases much time had

to be spent in inducing them to do so. On taking over the

charge of the reptiles here, I confess I was surprised to find

how readily they accepted dead prey. I was nevertheless of

opinion that some individuals would not accept dead food under

any circumstances, and I therefore asked and obtained the

Secretary’s permission to offer live food in certain cases where

dead had been persistently refused. From the summer of 1911

up to that of 1915, living prey was offered to nine snakes that

had refused the dead as food. The results given below are, I

think, convincing, showing that, with possible rare exceptions,

a snake that refuses to feed on dead animals is not more likely

to accept these if alive.

1. AnaconpAa (Hunectes murinus).

This snake, a large specimen, 16 feet in length, refused dead

food for the first eight months of its captivity. A live duck was

offered on two occasions in the course of the first fortnight of the

9th month, but was refused on both. The third week it was

onee more given dead food, and this was accepted. It has since

fed with the greatest regularity on dead chickens and ducks,

consuming on an average one a fortnight.

2. RericutarEep Pytuon (Python reticulatus).

This, a very large snake measuring 24 ft., had fed since its

arrival in 1898 with some regularity on dead kids and ducks.

In August 1911 it broke its jaw in the process of swallowing a

584 MR. E. G. BOULENGER ON

kid. The jaw was set in plaster of Paris, and the setting was

removed a month later. After the accident it refused to feed,

and in November it was decided to tempt it with something

alive. Live ducks were offered on two occasions in November,

but were not taken. In the first week of December it was once

more given dead food—a duck, which was immediately accepted.

The snake unfortunately died a few weeks later.

3. Common Boa (Loa constrictor).

This snake, measuring 7 ft., was presented to the Society in

July 1912. It had been kept in captivity some time before its

arrival here, and been fed on live food only. For the first two

months of its captivity at the Gardens it was offered dead rats,

rabbits, and pigeons, all of which were refused. As it had pre-

viously fed well on live animals it was, in September and October,

offered such, but they were not taken. In November dead food

was again offered and this time accepted, the snake feeding

henceforth with some regularity up to June 1913, when pneu-

monia caused its death.

4. Soutn-Arrican Hovuse-Snake (Boodon infernalis).

This snake had been kept by me for two years previous to my

taking over the charge of the reptiles here. It had been fed

on ie mice. On re transference to our Gardens it at once

took dead food.

5. Purr ADDER (Ditis arietans).

Received in June 1911, this snake fed with some regularity on

dead rats up to the end of December. For the first six months

of 1912, however, it refused food, and as it was getting thin we

decided in June of that year to let it have live food. Live rats

were accepted during part of June, July, and August, and part

of September. Towards the end of the latter month, dead food

was substituted and taken, the snake feeding on dead rats to

within a week of its death in January 1914.

6. BusuMastEeR (Lachesis mutus).

This snake was presented to the Society in December 1912.

As I had been informed by Mr. Mole, of Trinidad, who had

experience of this species, that it had never been induced to take

dead prey, after offering it dead rats for three consecutive weeks,

I obtained the permission to give it live food. The live rats

were, however, likewise rejected, and the snake died in March of

starvation.

BusHMASteR (Lachesis mutus).

This specimen, received in April 1913, was likewise offered

living animals after refusing dead food for some weeks. It also

enced to feed at all, and edn in September.

SNAKE-FEEDING. 58

8. Crossep Viper (Lachesis alternatus),

This snake, received in April 1914, was offered living food

after having refused dead prey for fifteen months. The live mice

were accepted, but unfortunately the snake died shortly after its

meal.

9. Texas RatrLesnakeE (Crotalus atrox).

The snake, acquired in June 1909, fed from time to time upon

dead animals up to June 1911. Having refused food from then

up to September, and as it was becoming emaciated, in the first

week of September it was offered live rats, which were refused.

The following week it fed on three dead mice. From that day

to its death in January 1912 it refused all food whether live

or dead.

10, Texas RarrLesNAKE (Crotalus atrox).

As this snake, which was received in May 1912, refused dead

food for the first six weeks of its captivity, it was decided to

give it living animals, and it was offered live rats during the

month of July. It, however, refused both the live and the dead

food, and died in September.

Although a number of persons with much experience in keeping

these reptiles in captivity, have been unsuccessful in inducing

their specimens to accept dead prey, the above records, I think,

prove that, in our Gardens, at least, it is quite unnecessary to

give any snakes live food, as out of about 300 snakes kept here

during the period covering these observations, with the exception

of the Crossed Viper which died shortly after its meal, not a

single snake would feed on live animals only, and in four cases

dead food was accepted after the live prey had been refused.

A point greatly in favour of giving dead animals to the snakes

lies in the possibility of examining the former for tubercle. Up

to the month of June 1910 the animals intended for the snakes

were not examined. At the suggestion, however, of Professor

Plimmer, the Society’s Pathologist, from that date onwards the

food has been carefully inspected and about 5 per cent. condemned,

with the result that tubercle in snakes which prior to June 1910,

accounted for 14 per cent. of the deaths, has been reduced to just

over 3 percent. In the years 1908, 1909, and the first half

of 1910, before the inauguration of the new system, 33 snakes

in all died of tubercle, while since the examination of the food,

i.e. the second half of 1910, and the years 1911, 1912, 1913,

1914, and the first half of 1915, there have been 23 cases only.

Tt is generally believed that if snakes will take dead animals

these have to be quite freshly killed and warm. Such is,

however, not the case, for, as has been pointed out by Dr.

Chalmers Mitchell and Mr. Pocock, the prey is frequently not

taken until long after it has been introduced into the cages.

586 MR. E. G. BOULENGER ON

As an instance of this, it is, I think, worth recording that one

of our large Indian Pythons on one occasion did not take a

rabbit which had been given it at 4 o’clock on a winter’s after-

noon, until 9 o’clock next morning, when, aS was only to be

expected after 17 hours in a temperature of nearly 80°, it was

in an almost putrid condition.

Another point of peculiar interest is that while tame rats are

acceptable to a large number of the snakes, wild rats are seldom

taken, and even when accepted are not digested but brought up

again a few days later. That this should be the case with rats

caught outside the Gardens is understandable, but the wild rats

I refer to are those caught in the Gardens and are therefore,

living as they do on the remnants of the food provided for the

exhibits, comparatively clean feeders.

The experience of Mr. H. N. Ridley who, writing of the

pythons in the Botanical Gardens in Singapore, stated that small

specimens fed about once a month, large ones once in six to

nine months, did not coincide with that of Dr. Mitchell and

Mr. Pocock, who recorded the fact that the majority of specimens

fed with the greatest regularity during the summer months,

some only refusing food when about to shed their skins. As

may be seen below, my experience with the large snakes likewise

differs from that of Mr. Ridley: two large specimens feeding on

as many as thirty occasions during the year, the longest period

of fasting amounting to just over a month ; while of the smaller

specimens it will be noted that a Boa which did not fast fora

longer period than three weeks, fed on thirty-five occasions in

the course of a year.

The table on p. 587 gives a detailed record of the feeding of a

number of healthy specimens over a period of one year.

587

SNAKE-FEEDING,

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SnIUDADE sNPLOYIOLIF ) OYRUG-19zV MA APCs.

(puna endsypnes.t) ang aon

(UyOC ehin) Cog SULMOLING WeIpUy

(45 F) beneee Pee tree ee ree ners

(ay g) oo onddn ChOMcrOND.OC GGaroOD ‘cc ce (4 (14

(936) (oC (.20792498U09 DOT) BOT WOULULOD

(33.91) 7 (snuLunu seqoauniy) epuoowuy

(44.9) “" (agas woyghT) woyydg ree lV

(ht 8) ce ce it3

(45 G1) * (snunjow noyyhg) noyuysg uwerpuy

Gra) tbe ale ta seee vee ‘c u: “c c

(ay LI) Cae wee eee weeny ce ce ce ce

(4h. BS) oo (Sngmpnargas uoyghg) uoyysg peyenoyary

OUD AT

bask eerie eS

RE TOA

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5 ee Se SS)

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fa reon

ON AVIAN CESTODES. 589

43. Contributions to the Anatomy and Systematic Arrange-

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

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

[Received October 1, 1915; Read October 26, 1915. |

(Text-figures 1-6.)

XVIII. On 72NIA STRUTHIONIS (PARONA) AND

ALLIED Forms.

INDEX.

Page

TEM ROC NDEI? os ocsee soe eRe Penh en teats oe OSO

Meveninnn Gt? Doonan te PAae ct cnaee ae nce nenesk MROOO

I have in my possession a considerable number of examples of

a Cestode from the Ostrich Struthio masaicus, which are either

identical with Tenia struthionis of Parona*, or belong to 2

closely allied species. The description given by Parona is not

quite sufficient to enable the identity of his species and mine

to be established beyond doubt. But it is at least clear, as I

shall point out presently, by comparing the facts of structure

one by one, that the species described by Parona and that to be

described here by myself are not to be referred to the species

described under the same specific name by v. Linstow ‘*.

Although the details given by Parona are scanty, they are

qtute sufficient in my opinion to forbid ay confusion between

his species and that more fully dealt with by v. Linstow. My

chief reasons for regarding them as two distinct species are the

following. In the first place, v. Linstow’s Cestode was obtained

from Struthio molybdophanes ; I infer that Parona obtained his

worms from Struthio camelus. The scolex of Tenia struthionis

of v. Linstow is only 1-18 mm. broad, while the species described

by Parona has a stouter scolex of 2 mm. diameter =.

“ Ein eigentliche Rostellum ist nicht vorhanden ”—says

v. Linstow of his species, while that described by Parona

las, according to his figure, a quite strong rostellum. Corre-

jated with this would appear to be the feebler character of

the rostellar hooks in the worm from Séruthio molybdophanes.

The width of the proglottids in the two forms also appears to

differ greatly ; in the Tzeniid described by v. Linstow, the diameter

is but 4mm.; while in Parona’s specimens the same measure-

ment was from 8 to 9 mm., 7. e. quite double that of the first-

named variety. This seems, like the other feature mentioned in

* Ann. Mus. Civ. Genova, (2a) ii. 1885, p. 425.

+ Arch. Mikr. Anat. xl. 1893, p. 447.

* But see the observations of Zilluff quoted later (on p. 591) which tend to

reduce the importance of this apparent difference, but do not affect what follows

in the above réswié.

Proc. Zoou. Soc.—1915, No. X LI. 4]

DY90 DR. F. E. BEDDARD ON

this brief account of differences, to be hard to reconcile with

specific identity. The account given by Parona of internal

structure is so slight that the comparison cannot be pursued

further.

A question of nomenclature thus arises. The name Tenia

struthionis first occurs in Rudolphi'’s “ Synopsis” *, it is there a

nomen nudum, but given on the authority of Houttuyn in

Miiller’s edition of Linneus?. In the earlier work of Rudolphiz

the same worm (I presume) is named Jeenia struthiocameli, and

is also a nomen nudum, and again referred to Houttuyn in

Miiller’s Linneeus§. I am indebted to Mr. C. Davies Sherborn

for kindly informing me that Houttuyn himself || does not refer

to the ostrich at all in his work, though Tzenias are mentioned.

It is thus erroneous to term the species Zenia struthionis or

Tenia struthiocameli Houttuyn.

In Miller’s work there is no name given at all; the occurrence

of a Zenia in the ostrich being merely mentioned. Thus if a

nomen nudum has any claim at all to be admitted, the species is

to be’ referred to Rudolphi and is to be cailed Tenia struthio-

cameli, since the earlier of the two works by that author which

mention the species calls it by that name. Diesing 4], how-

ever, quoting both Muller and Rudolphi’s two works, terms the

species Tenia struthionis, but again as a nomen nudum. The

earliest actual description therefore of a Tenia from Struthio is

that of Parona already referred to. We may perhaps safely

accept his name, since it is accompanied by a description though

not a conelusive one. I shall have to return again to this matter

in considering the species to which it seems necessary to refer

the worms which [I now describe. :

The scolex of the worm which forms the subject of the present

communication is a little over 1 mm. in breadth in the two or

three examples in which I measured it. The region of greatest

breadth is opposite to the suckers; but the breadth was not

increased by the extrusion of the latter. The suckers lay within

the contour of the scolex. It is clear therefore that this species

has a less robust scolex than Parona’s Tenia struthionis. But

while the actual measurements of the scolex of my species agree

more with those of the worms described by v. Linstow as Tenia

struthionis, my species shows a scolex with a well-developed

rostellum, thus disagreeing with v. Linstew’s worms and so far

agreeing with that described by Parona. This is very evident

from the figure given by Parona **, where the hardly extruded

rostellum is “plainly exhibited. Par Onn does not state the number

* Hntoz. Syn. Mant. 1819, p. 175.

ii Linné s Naturgeschichte von P. L. 8. Miller, Th. vi. Bd. ii. p. 904.

* Entoz. Hist. Nat. 1810, p. 209.

§ It is to be noted that the initials of Miller are as st ated here. He is referred

o as “St. Miller” by Rudolphi, and “H. Miller” by Diesing.

2) Sannto ye Historie, vol. i. pt. 14, 1770.

| Systema Helminthum, 1 deat, p. 555.

i Parona, loc. cit. pl. vi. fig. 2.

AVIAN CESTODES. 591

of hooks present on the rostellum; v. Linstow gives the number

found by himself as 180. I find in my species something

between 120 and 130. These hooks are, as in other Davainea, of

the well-known hammer-shape so characteristic of the family

Davaineide.

The hooks really form two concentric rows, which arrange-

ment is only clear in sections which pass through the “ handle

part of the hook; that they are of different sizes is only shown

in the * head” a the hammer, where one series is much shorter

than the other; I could find no such difference in thickness in

the “handle” region of the hooks. An alternation between

larger and smaller hooks is stated by Parona to occur in his

species. The hooks are of course implanted upon the edge of the

circular rostellum. They are of the usual golden-brown colour.

Von Linstow has represented the hooks of his examples, called

by him Tenia struthionis, as being weak and frayed out at the

point of implantation. I have found nothing of the kind in the

robust (though small) hooks of the examples examined by

myself.

My own observations are in fact more in accord with those of a

later investigator than those referred to. Dr. Zilluff*, referring

only to v. Linstow’s paper and not to that of Parona, naturally

finds differences to record (“naturally” if I am correct in

thinking that v. Linstow’s specimens are of another species than

that which Parona and I describe). He emphasizes the rostellum

and gives the diameter of the scolex as 1-33 mm., the dimensions

agreeing with mine rather than with Parona’s. But this author

does not mention from what species of Struthio he obtained the

material.

The suckers are not armed, as is the case in certain other

members of the genus, a great part of the species of which have

armed suckers. 7 believe that I can state this fact positively.

Excepting where the retractor muscles are attached to the

suckers, the latter lie for the most part free within the cavity of

the scolex to which they are fitted. A space is generally visible

between sucker and body-wall. Although there is no apparent

difference that I could detect between the individual suckers, I

have noted in this worm a means of distinguishing the dorsal

from the ventral couple. The two dorsal vessels, instead of

ending in the medullary region like the ventral ae of the

water-vascular system, bend dorsally, each of them perforating

the Jayer of longitudinal muscles of the cortex ends in the

neighbourhood of one of the suckers. The exact mode of ending

T did not ascertain. It is therefore possible to distinguish two

of the suckers as belonging to the dorsal surface. The characters

of the musculature of the scolex I shall deal with later in

* “ Veroleichende Studien tiber die Muskulatur des Skolex der Cestoden.”

Jnane.-Diss. Univ. Ztirich, 1912. (Published also in Arch. f. Nature. of the same

year.) See also Lithe in Zool. Anz. xvii. 1894, p. 280.

| 41*

592, DR. F. E. BEDDARD ON

connection with the general arrangement of the muscles of the

body.

Von Linstow particularly mentions that in the species studied

by himself, the anterior part of the body is devoid of calcareous

bodies. In the specimens which I have examined by sections,

the caleareous bodies are peculiarly numerous anteriorly, and

especially in the scolex, where they form in parts closely aggre-

gated masses as is shown in the accompanying sketch (text-fig. 1).

Text-figure 1.

Mm.

Longitudinal section through scolex.

ea. Caleareous bodies. h. Hooks seen in transverse section through “root.”

m. Muscles of rostellum ending above in rostellum. gs. Sucker.

w.v. Water-vascular tubes.

I need not describe their distribution in the scolex exactly, for

they occur everywhere between the outer skin and the suckers

and rostellum, except, however, among the muscle-fibres of the

longitudinal muscular layer. Further back in the neck region

the medulla is largely occupied by masses of calcareous bodies

AVIAN CESTODES. 593

which are generally speaking very abundant in this species, so

much so that I should be inclined to add the abundance of these

bodies to any definition of the species. This is another reason

for refusing to accept the identity of the worms from Struthio

masaicus With those from Struthio molybdophanes.

The general shape of the body of this worm is as figured by

Parona, The anterior region of the body is slender; it is much

wider posteriorly, but not by any means so wide as in the species

Text-figure 2.

Part of a transverse section through a proglottid in anterior region of body.

C. Cuticle. I & Lz. Layers of longitudinal muscles. T. Transverse

muscles.

described by Parona. I found 5 mm. to be the greatest diameter

of the posterior proglottids. They are overlapping, and as a

rule so contracted as to be much wider than long. In a few cases

the proglottids were, however, more expanded, but were never

actually longer than broad. The worms reacha length of perhaps

nine or ten inches.

The cortical layer is deep, the diameter being greater than

594 DR. F, E. BEDDARD ON

that of the medullary layer. This is particularly marked in the

anterior segments, where the reproductive organs are only just

beginning to appear. The longitudinal muscle-layer presents

definite characters in the arrangement of its fibres, as is general

among Cestodes. It is not usual to find accurate figures of the

course of these fibres, which are constantly of systematic im-

portance. I therefore &ttempt to reproduce here such accurate

drawings.

At the base of the rostellum the longitudinal muscles lie in a

continuous circular layer, in which form they are implanted upon

the rostellum. A little further back, at the level of the suckers,

the layer of muscles is markedly divided up into separate bundles

which are of different sizes. There are 12 or 13 of these

separate bundles which are more or less completely separated.

In the neck, which immediately follows upon the scolex, the

bundles cease to exist as separate structures except at the two

sides opposite to the water-vascular tubes. The wnsegmented or

neck region in this worm is very short and, as in the other

examples ascribed to the species Tenia struthionis vel 7’. struthio-

cameli, it may fairly be remarked that a neck can hardly be said

to be present. Further back—but still in the anterior region

of the body, where the gonads and their ducts are still only

recognizable as a mass of condensed nuclei—the longitudinal

muscular layer has more or less acquired its definitive arrange-

ment. It is here (text-fig. 2) divisible into two quite distinct

sheets. That nearest to the medulla consists of a row of

bundles each consisting of a good number of individual fibres

which are packed close together and separated by vertical fibres

forming a dividing palisade. Above this is a very distinct space

dividing the lower layer from the upper. This space is formed

of ground-tissue, and there is no trace therein that I could

discover of transverse muscle-fibres. On the outer side of this

space 1s a layer of smaller bundles, 7. e. each bundle consisting

of comparatively few fibres, and above this again, without any

marked interval, a certain‘number of single muscle-fibres, which

complex reaches some way towards the subcuticular layer. In-

side the whole longitudinal layer of muscles is a thin layer of

transverse fibres separating these in the usual way from the

medulla. Further back in the body the same arrangement

exists, but it is not so clear cut as anteriorly. That is to say,

the two layers of the longitudinal sheet are quite recognizable,

but they are. not so markedly divided from each other. This is

shown in text-fig. 3.

Besides the sheets of muscle mentioned so far, the worm has,

like most other Cestodes, a dorso-ventral system. I have already

spoken of dorso-ventral fibres running between the bundles of

the longitudinal coat. In addition to these the medulla is

traversed by single fibres which cross it at right angles to its

long diameter, and are numerous, dividing the medulla into quite

narrow segments when seen in transverse sections.

AVIAN CESTODES. 595

The water-vascular tubes of this worm present no remarkable

characters. The much larger ventral vessel is alone present. in

the posterior segments. Anteriorly both tubes are visible and

superposed. The transverse trunks unite the ventrals in each

segment. The usual valvular flaps in the ventral vessel are

obvious and attached, as is usual (but not universal), to the inner

wall of that tube.

Text-figure 3.

Part of a transverse section through a proglottid in the posterior region of

the body.

Lettering as in text-fig. 2.

The male and female efferent ducts open into a cloaca genitalis

which is not specially deep. The genital pores are completely

unilateral in this species, and somewhat anterior in position, at

any rate in front of the middle line of the lateral border. I have

found so many orifices in succession opening on to one side

of the body, that I cannot believe that the conditions are for

instance as in Chapmania tauricollis, where the orifices really

596 DR, F. EB. BEDDARD ON

are alternate, though many open successively on to one side.

All the pores that I found were on the same side of the body.

The curus-sae is comparatively short *, as in many species of

Davainea, but not in all. It only just reaches the nerve-cord.

It opens into the cloaca genitalis in front of and to one side of

the vagina. ‘The exact shape of the cirrus-sac has been carefully

described in many Cestodes by many writers, and thus specific

distinetions have been partly based upon its characters. The

cirrus-sac of the present species shows that care must be taken

in such descriptions. For I find considerable differences between

the cirrus-sac in different segments, a state of affairs to be

accounted for no doubt by varying contraction of its muscular

walls.

Text-figure 4.

Longitudinal section through cirrus-sac (e.).

n. Nerve-cord. o. Orifice of genital cloaca. v. Vagina. v.d. Vas deferens.

More usually perhaps the cirrus-sac has the appearance repre-

sented in text-figure 4, which is drawn from a horizontal

section through a more anterior segment. ‘The cirrus-sac opens

directly into the cloaca genitalis, and is of the same character

and of pretty well the same diameter throughout. The walls are

muscular but not thick; nor are they thicker in one region than

in another. Thevas deferens perforates the muscular coat at the

extremity of the sac fairly exactly in the middle line, and is

* In extended proglottids the eirrus-sac les obliquely, being directed forwards.

AVIAN CESTODES. 597

coiled within the sac. The cirrus, with which it is continuous,

appears to run a straight course and not to be coiled, since it is

shorter than the cirrus-sac. In some posterior segments the

cirrus-sac presented a different appearance. The peripheral and

greater part of the cirrus-sac is thicker-walled than a terminal

rather spherical and wider region into which opens the vas

deferens.

The cirrus-sae is ensheathed externally by a layer of rather

large nucleated hyaline cells, a not unusual character.

Did these two forms of the cirrus-sac occur in ,different

individuals, one would be tempted to see in them a specific

difference.

The vas deferens presents an extensive coil after it issues

from the cirrus-sac. This occupies quite one-third of the dia-

meter of the segment when the latter is stretched laterally.

The coils are at least mainly dorso-ventral in direction, since 1n

horizontal sections the sperm-duct appears as a series of circular

transversely cut areas.

The vagina has a straight or at most slightly sinuous course

back to rather beyond the water-vascular tube—this section

being thick-walled with a narrow lumen as in so many other

Cestodes. <A. little way to the inside of the water-vascular tube

the vagina narrows into an excessively fine bore, though with

equally thick muscular walls at first. This slender region 1s

coiled on the horizontal plane. It opens into the receptaculum

seminis, which is rather pear-shaped. This and the succeeding

portion of the vagina is not thick-walled but has a wider lumen,

less of course in the case of the vagina. Although the proglottids,

in which the vagina and its subdivisions had the characters that

have just been mentioned, were not fully mature, at any rate as

far as concerns the testes and sperm-duct, the receptaculum

contained spermatozoa. It is necessary to point out that there

is nothing to be specially remarked upon in the structure of the

female efferent duct, which is constructed upon the plan usual

in tapeworms. It is important, however, to be accurate, since

there are minor differences to be noted which affect even the

different species of Davainea.

Without attempting any general réswmé for comparative

purposes, [ may direct attention to one or two species which

differ from that now under consideration in these matters. In

D. sphecotheridis of Johnston * there is apparently no distinct

receptaculum seminis at all. In YD. corvinad Fuhrmann 7 the

position of the receptaculum is different, beginning as it does

to the outside of the water-vascular tube. In D. polycaleeola =

the small receptaculum is close to the ovary. It is of further

importance to note the age of the proglottid when giving the

* T. Harvey Johnston, “Second Report on the Cestoda and Acanthocephala

collected in Greenland.” Ann. Trop. Med. Parasit. i. 1914, p. 107.

+ Abh. Senck. Nat. Ges. xxxiv. 1911, p. 252, fig. 3.

tv. Janicki, “Ueber zwei neue Arten.... Davainea,’ Arch. de Parasit. vi.

1902, p. 265, fig. 5.

598 DR. F. E. BEDDARD ON

characters of the vaginal complex. The above description of

that of “ Davainea struthionis” relates to not fully mature pro-

glottids. In fully mature proglottids the conditions observable

are a little changed.

The female duct (see text-fig. 5) from the receptaculum seminis

Text-figure 5.

Part of transverse section through nearly ripe proglottid.

7.s. Receptaculum seminis. v., v;. Proximal and distal ends of vagina.

v.d. Vas deferens.

to its median end is gorged with sperm, and thus presents the

appearance of an elongated receptaculum such as that referred

to above in Davainea corvina. And, moreover, there is this

further resemblance, that the vagina is pressed by its increasing

AVIAN CESTODES. 599

contents up to the margin of the water-vascular tube, occasionally

crumpling up the latter beforeit. Nevertheless, the more dilated

region is still distinguishable as the true receptaculum seminis.

It is evident, therefore, that the differences apparently shown

between species in the vagina must be handled with care. I

may add that wn fully mature proglottids the vagina appears

to be continued onwards beyond its junction with the other

tubes of the female system. This may be merely a_ burst,

though in some cases it has a tubular character. It is here,

I assume, that fertilization occurs.

Text-figure 6.

Ripe ova enclosed in capsules.

o. Capsule containing ovum. sp. Part of vagina gorged with sperm. ¢. Remains

of a testes closely adpressed to an egg-capsule, the nuclei in the walls of which

are represeted.

The uterus in the genus Davainea is never a conspicuous

structure and never, when it exists, does it persist long. It is,

however, too much to say—as does Ransom *—that “a definite

functional uterus is not developed.” For in D. aruensis Fuhr-

mann Y has described a uterus with a lining of cells and con-

taining ripe ova, which uterus, however, rapidly disappears. The

same appears to be the case with D. nicroscolecina and D. corvina,

* <The Tenoid Cestodes of North American Birds.”’. Bull. U.S. Nat. Mus.

No. 69, 1909, p. 14.

+ Nova Guinea, vol. ix. Zoologie, Livr. 3, p. 469.

600 DR. F. . BEDDARD ON

where the same author * remarks upon the rarity of observations

upon the uterus of this genus. I find in the species with which

{tam here concerned very definite beginnings of a uterus, in

which, however, I have not seen a large and continuous cavity.

This consists in horizontal sections through proglottids, which

ave not fully mature but in which nevertheless the receptacula

seminis are full of sperm, of a wide stretch of condensed medul-

lary tissue. This structure appears to me to be exactly like

the commencing uterus of some other Cestodes?. It lies in

front of the ovary and shell-gland, but behind the receptaculum

seminis and vas deferens, occupying thus about the width of

the segment. It extends to a considerable distance right and

left. The string of tissue representing the uterus is mainly to

be differentiated from the surrounding medulla by its crowded

nuclei. It is not solid but contains numerous cavities of various

sizes. Some of these were filled with cells which may well be

egg-cells. These cavities are at least frequently of the same size

and shape as the oval interstices of the medullary meshwork.

Later the proglottids (see text-fig. 6) are full of embryos each

in its own separate cavity.

The following assemblage of characters are perhaps suflicient

to define this species, to which I shall be unable to give a name

with absolute certainty that it requires a new one. It will be

better therefore to leave this matter unsettled for the present.

Definition of DAVATNEA SP. parasitic in Struthio masaicus.

Length 10-14 inches ; greatest diameter of proglottids 5 mm.

Scolex 1-2 mm. diameter, with double row of 130 hooks in all ;

suckers unarmed. Scolex and anterior part of the body abound

unth calcareous corpuscles, which also occur posteriorly. No neck

present. Segments of body not longer than broad ; ripe segments

not moniliform. Genital pores unilateral. Cirrus-sac reaching

to nerve-cord. Dorsal water-vessel absent from posterior reyion of

body. Ova imbedded singly in parenchyma extending into cortex.

The above will be sufficient pending a revision of the genus to

place the species approximately.

GENERAL REMARKS.

Tt is pretty clear from the foregoing observations upon the

external characters and internal structure of this Cestode from

Struthio masaicus, that it is certainly not to be confounded with

the species named by v. Linstow Davainea struthionis, and which

& fig. 4, p. 252. ;

y+ Cf. e. g. Beddard in the instance of Ohapmania tauricollis, P.Z.S. 1915

p- 434, text-fig, 3.

AVIAN GESTODES. 601

is a parasite of another subspecies of Struthio, viz. S. molybdo-

phanes. While the general dimensions and the relative size of

the scolex seem to be much the same in v. Linstow’s species and

in my own, there are several salient features in which they

disagree markedly. The scolex of v. Linstow’s worm has no

rostellum, in the species examined by myself the rostellum is

strong: v. Linstow’s species has no calcareous bodies in the

scolex, while my species is peculiarly well provided with these

bodies : whatever may be the interpretation of the “ovaries” of

v. Linstow in the posterior segments of his species, whether they

are really a divided uterus or paruterine bodies, that Cestode

clearly differs from mine where the embryos are scattered each

oneina cavity of its own: finally, if v. Linstow’s representation

of the cirrus-sac and the vagina opening quite separately prove

correct, there is here a great difference from my species, where

the relations between these ducts is quite normal. These facts

are, as | think, sufficient to show that there can be no identity

between the two Cestodes of Struthio masaicus and Struthio

molybdophanes *.

On the other hand, an exact comparison of my species with

that termed Tenia struthionis by Parona is more difficult. If we

can trust as differential characters the diameter of the proglottids

and the size of the scolex, then the two forms are different.

There are no other data that seem to permit of a more definite

expression of opinion.

* T have suggested (P. Z.S. 1915, p. 430) that v. Linstow’s species may be

actually referable to the genus Chapmania,

ON THE ANATOMY OF RANA TIGRINA. 603

44. Some Notes upon the Anatomy of Rana tigrina. By

Gro. E. NicHouns, D.Se., late Professor of Biology,

Agra College, India *

[| Received October 9, 1915: Read November 9, 1915. |

(Text-figures 1—3.)

_In several skeletal and other characters, Rana. tigrina—the

so-called Bull-frog of ee markedly from our common

European grass or water frogs (2. temporaria, R. esculenta).

These two” frogs, which eee ae one another fairly closely,

appear to be the only species of which a detailed description has

been given, and figures of one or the other alone appear in text-

books all the world over. For the Indian form these figures are

in some respects quite misleading, and since this frog is now

generally used throughout India as a laboratory type, it has

seemed desirable that attention should be called to those features

in which &. tigrina differs from its European congeners.

1. The Vertebral Column.

In correspondence, doubtless, with the much larger size of this

frog, the vertebrz are distinctly more stoutly built than is the

case In &. temporaria. The neural arches are, relatively, greatly

developed antero-posteriorly. Thus, when viewed fiom above,

the vertebral column of this species does. not show series of gaps

between the arches such as is so clearly seen in 2. temporaria

(cf. Howes, 02, fig. 35). On the contrary, there is, in R. tigrina,

a very marked overlap of each arch dorsally upon that immediately

posterior to it, and accordingly, when the vertebre are in position

(text-fig. 1), the centra are not visible from above.

Such a condition as this is said to be imbricate, and to

characterize the Discoglosside and Pelobatide (Boulenger, 797,

p. 38). Concerning the European species of Ranide, Boulenger

points out that precisely the opposite condition prevails. His

statement may be quoted :—‘‘ The neural arch is either closed

above ... or notched between the zygapophyses so as to expose

the spinal cord between every two vertebree; the latter type is

most marked in ana, in which, the lateral openings for the exit

of the spinal nerves being also of large size, the vertebral column

forms an open-work above and on the sides.”

While this ‘ open-work ” vertebral column is seen typically in

the European Ranide, it is also found in most of the other

Anura, so that Boulenger notes this as one of the characters

which separate the Bufonide and Hylide from the more

generalized Arcifera. In this imbricate condition of the

* Communicated by Prof. ARTHUR DrENpy, D.Sc., F.R.S., F.Z.S.

604 DR. G. E. NICHOLLS ON THE

vertebral column it would appear, then, that Rana tigrina has

retained (or reverted to) a somewhat primitive condition.

Text-figure 1.

The vertebral column of Rana tigrina, as seen from above. x 3.

J , 5)

af-t., aperture for filum terminale ; c., coceyx; ep., epiphysis; f2, flange upon

the transverse process of the second vertebra, IT.t.p.

It does not seem, however, that the occurrence of gaps

between the neural arches in R. temporaria is to be attributed

simply to a notching of the arches between the zy gapophyses.

ANATOMY OF RANA TIGRINA. 605

A comparison of the neural arch of a typical (e.g. sixth)

vertebra of the grass-frog with that of the corresponding

vertebra of A. tigrina or of Pelobates fuscus, will, 1 think, bear

out this statement (text-fig. 2, A-C).

In all three cases it will be seen that there is an incisure upon

the anterior face of the neural arch between the zygapophyses.

In the Bull-frog and in Pelobates this incisure is a deep one,

whereas in 2. temporaria it is broad but comparatively shallow.

Upon its posterior border the arch is practically not incised at all

in &. temporaria, aud is most deeply notched in Pelobates, the

condition of 7. tigrina being intermediate in this respect.

Text-figure 2.

The sixth vertebra of (A) Rana tigrina (x 2), (B) R. temporaria, and (C) Pelo-

bates fuscus (both X 3), to show the relative dezree of incisure of the neural

arches and the development of the neural spines in the three species. (a) Dorsal,

(6) ventral, and (c) posterior view.

In Pelobates and in R&. tigrina, however, the centrum has

practically the same length as the neural arch, whereas in Rf. tem-

poraria the centrum is, approximately, half as long again as the

neural arch. When the vertebre are articulated in the normal

manner, therefore, the neural arches do not come into contact in

this species, excepting at the zygapophyses, notwithstanding that

some of the surplus length of the centrum has been absorbed in

the concavity of the following centrum. In the case of FR. tigrina

(and Pelobates) the neural arch is sufficiently long to allow of

considerable overlap upon the succeeding neural arch.

Neural spines, too, are well developed (text-figs. 1,2). This

Proc. Zoou, Soc.—1915, No. X LIT, A2

606 DR. G. E. NICHOLLS ON THE

is especially marked in the anterior vertebre of the column,

where the hinder end of a neural spine may extend backwards

upon the ensuing vertebra almost to the level of the notch

between its post-zygapophyses (cf. text-fig. 1). Posteriorly the

spines become shorter, though still well developed. Thus, upon

the seventh vertebra the neural spine extends but little beyond

the posterior margin of the neural arch ; in the eighth it is

nearly vertical, and in the ninth the spine is slightly forwardly

directed, so that its apex approaches very closely to that of the

preceding vertebra. In this development of neural spines

f. tigrina 1s remarkable, for of Kuropean Anura Boulenger has

remarked : ‘‘ Neural spines are absent or represented by a low

keel, which is much prolonged posteriorly in Discoglossus and

Pelobates” (op. cit. p. 38).

The intervertebral foramina in the Indian Bull-frog are,

relatively, considerably smaller than are the corresponding

nerve-exits in /. temporaria, and the column has, therefore, not

at all the open-work structure which is so characteristic of our

Huropean frogs.

Moreover, the cartilaginous epiphyses found, upon the distal

ends of the transverse processes are particularly noticeable in

R. tigrina. Upon the third vertebra (cf. text-fig. 1) these are

very large indeed, and backwardly directed, recalling strongly

the condition figured by Boulenger for Pelobates fuscus (op. cit.

fig. 75).

The transverse processes also are, relatively, much longer than

are the corresponding structures in /. temporaria, but in this

respect the condition of &. esculenta is closer to that of the

Indian species.

Apart from these generalities, there are notable differences to

be observed in the second, eighth, and ninth vertebrz of the two

species.

In the second vertebra of 2. tigrina there is developed a very

pronounced flange-like projection upon the anterior border of the

transverse process (text-fig. 1, #). This varies somewhat in size,

but becomes very marked indeed in some specimens. It serves,

apparently, for the attachment of the Mdm. intertransversarit

capitis, which have their insertion upon the base of the skull,

slightly lateral to the condyles. The complete absence of this

flange from the European Ranide is doubtless to be attributed to

the much slighter development of these muscles in these more

slightly built frogs.

In the eighth vertebra, the transverse processes are as long as,

and rather stouter than, the diapophyses of the three preceding

vertebre. They are peculiar, in the normal vertebral column of

R. tigrina, in that alone of all the transverse processes they are

sloped slightly forwardly (conf. Howes, ’02, fig. 35, with my text-

fig. 1). The well-developed neural spine rises almost vertically.

In the ninth vertebra, the transverse processes should be

described as slightly conical (with the base of the cone distal)

rather than as cylindrical, which latter shape is said to be

ANATOMY OF RANA TIGRINA. 607

characteristic of these structures in the Ranide. In some

specimens J have observed a flattening even of the distal

extremity of the diapophysis, which, too, is sometimes to be

noticed in the immature Rana temporaria. ‘The neural spine is

somewhat variably developed, but is always represented by at

least a slight elevation in the mid-dorsal line. From this a pair

of distinct ridges diverge. ‘These pass outwards and backwards

onto the dorso-posterior face of the transverse processes, but fade

out before reaching the distal end.

The coccyx, too, in F&. tigrina differs from that of R. tempo-

raria or of FR. esculenta in that the paired foramina which, in

these European species, permit of the exit of the tenth pair of

spinal nerves are very frequently absent from the Indian species.

When, however, they do occur, they are generally very minute

indeed, and not uncommonly, upon one side or both, this

exter nal opening leads only into a blindly ending canal. Asan

infrequent variation, specimens are seen in which there may be

two apertures upon one side. In such cases only the upper

canal appears to have open communication with the vertebral

canal. Such a condition is figured in my account of the Anuran

coccyx (15, fig. 1}, x., x1.).

It is probable that we have here the last vestige of an aperture

for the lost eleventh spinal nerve, such as is still found occasionally

in the more primitive Anura.

From the foramen of the tenth nerve, when present, there

runs backwards and upwards a slight groove which is often

several millimetres in length.

Of a total of 32 coccyges examined, only four (123 per cent.)

showed a pair of canals for spinal x. ,and, of these, the openings

in three were very small. In five other Specimens a pair of

extremely minute external apertures were found, but on neither

side of three of these was there a clear passage for the finest

hair. The remaining to permitted the passage of a very fine

hair on one side only.

In seven other specimens a single aperture only was found, but

in only two cases was I able to pass a hair into the vertebral

canal. In the remaining five specimens the canal apparently

ended blindly internally.

Thirteen coccyges (40 per cent.), including four specimens in

which the vertebral column was abnormal in other ways, showed

a complete absence of the foramina on both sides.

Three specimens exhibited two minute apertures on one side

and a single aperture only upon the other. Of these, in one

specimen all the canals ended blindly, and of the remaining two

each had a single canal on that side upon which there were two

apertures.

In the twelve asymmetrical cases the canal (or aperture)

occurred in eight cases upon the right side and in four upon

the left.

It is a little difficult to decide what is to be regarded as the

normal condition of the coceyx in &. tigrina. It is probably

42*

608 DR. G. E. NICHOLLS ON THE

correct to say that in the great majority of the individuals of

this species, the paired foramina for the tenth spinal nerves are

absent or imperfect.

The Shoulder-Girdle and Sternwm.

The shoulder-girdle of F. tigrina (text-fig. 3) is very stoutly

built and, in general, resembles the condition of the corresponding

structure in 2. esculenta.

Text-figure 3.

The Shoulder-zirdle and Sternum of Rana tigrina (nat. size).

(A) Ventral view, and (B) dorsal view. In the latter the supra-scapulais removed

upon the right side.

cl., clavicle ; m., metasternum ; 0., omosternum ; 1.co., J.co., right and left

coracoids ; sc., scapula.

In one particular, however, viz., in the arrangement of the

coracoids, it presents a condition which has not, I believe, been

recorded in any of the Firmisternia.

ANATOMY OF RAWA TIGRLVA. 609

The coracoids, while having the normal shape and transverse

position, do not meet in a median epicoracoidal cartilage but

overlap each other in the middle line. At first sight 1t appears

that there is merely an uneven suture, such as is seen between

the epicoracoid cartilages of an immature Rana temporaria, but

a closer examination reveals the existence of a definite overlap

(cf. text-fig. 3).

It differs from the overlap of the arciferous condition in that

the epicoracoidal cartilage is completely calcified in the adult

and that the coracoid comes, at its antero-mesial border, into

contact with the clavicle upon the ventral surface. There is, I

believe, synostosis between the two coracoids, for there appears

to be no freedom of movement.

The direction of the overlap, in the specimen figured, resembles

that prevailing in the Arcifera, the right coracoid lying ventral to

the left, but the opposite condition is met with not infrequently *.

Dorsally the pre-coracoid cartilage is seen. It appears to be

calcified, and separates, somewhat widely, the clavicle from the

coracoid.

The bony style of the omosternum is also peculiar (amongst

Ranide) in being bifid posteriorly. The diverging processes

meet corresponding elevations upon the clavicles, and the small

space between the three bones is filled, in life, by a delicate

membrane.

The Tenth Spinal Nerve.

Correlated with the minute size, or the absence, of the foramina

in the coccyx, the tenth pair of spinal nerves are, in Rana tigrina,

either extremely delicate or, more often, altogether absent.

When present, they seem invariably to pass dorsally from their

exits, lying in that small groove in the coccyx, to which reference

has been made. In this disposition, therefore, this nerve differs

considerably from the corresponding structure in 2. temporaria,

in which it passes ventrally after leaving its.foramen. In the

Bull-frog the nerve, even if present, is hidden from view in a

dissection made, in the usual manner, from the ventral surface.

It never, in this species, I believe, makes any contribution to

the sciatic plexus, nor have I been able to demonstrate any

sympathetic ganglion related to it.

Trterature.

"11. Bepparp, F. E.—‘ Contributions to the Anatomy of the

Anura.” Proc. Zool. Soc. 1911, p. 393.

97. BouLencer, G. A.—The Tailless Batrachians of Europe.

Vol. I. Ray Society. 1897.

02. Howes, G. B.—An Atlas of Zootomy. London, 1902.

15. Nicuoutis, G. E.—“ A Note on the Urostyle (Os Coccy-

gewm) of the Anurous Amphibia.” Proc. Zool. Soc.

1915, p. 239.

* Beddard (11, p. 396) has pointed out that variation in this arrangement is also

occasionally encountered in Megalophrys fee.

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ON THE SNAKES OF EAST AFRICA. 611

45. A List of the Snakes of East Africa, North of the

Zambesi and South of the Soudan and Somaliland,

and of Nyassaland. By G. A. BouLencer, F.R.S.,

E.Z.8.*

[ Received Oetober 4, 1915: Read November 23, 1915.]

(Text-figures 1—3.)

INDEX.

GEOGRAPHICAL: Page

JOR SE EMBER, IUNSty OSES, Jy nck doaeieansnennnbenbedk teeegGepenae abe 611

SYSTEMATIC :

Keys to the identification of the Snakes of East Africa...... 611

This list is the third of a series published in these ‘ Pro-

ceedings’ t, and the reader is referred to the first for an

explanation of the method and scope of this aid to the identi-

fication of African Snakes. It is convenient to reproduce here

text-figures 1 & 2 from the first paper, as a glossary to the terms

used in the keys to the genera and species.

Synopsis of the Families.

I. Worm-like, with small inferior mouth, eyes hidden or visible under the head-

shields, and body covered with uniform imbricate scales above and beneath.

18 scales or more round middle of body ; ocular not bordering the

mouth ; tail not-or but little longer than broad................... TyPHLOPIDm.

14kccalesiroundemiddletotybodiyaess-s-eneeetas a aseeece eee eee eee AU CO ONTID a

II. Mouth large, eyes exposed; body with enlarged shields beneath (except in the

marine genus Hydrus).

Ventral shields much narrower than the body; supraocular, if

distinct, broken up into-two or more shields .......... .. Bors.

Ventral shields at least nearly as broad as the body ; supraocular

single; poison-fangs, if present, not in a very large sheath ... CoLUBRID®.

Ventral shields at least nearly as broad as the body; large polson-

tangs in a very large sheath below the eye ......+.....-.....-0605- VIPERID&.

Family TyPHLOPID4.

A single genus.

1. TypHuops.

Schneid, Hist. Amph. 11. p. 339; Bouleng. Cat. Sn. i. p. 7.

* Published by permission of the Trustees of the British Museum.

+ 1. “A List of the Snakes of the Belgian and Portuguese Congo, Northern

Rhodesia, and Angola,” P. Z.S. 1915, p. 1938. 2. “A List of the Snakes of

Madagascar, Comoro, Mascarenes, and Seychelles,”’ t. c. p. 369.

612 MR. G. A. BOULENGER ON THE

Text-figure 1.

(From P. ZS. 1915, p. 194.)

supraocular _ rostral ocular rostral nasal

nasal, * preocular : ‘ preeocular

pracfrontal ; rostral upper

iy nasal i y “labial

przocular EN: ISH

| cs ocular

ocular = SSSS8

: sesseosee

upper labial

TYPHLOPS PUNCTATUS.

rostral _praefrontal

rostral _nasal

supraccular g

nasal

Qc upper

nasal... ocular PA, fabial

ocular aa! \-ocular

j -upper

frontal-- x Tel

GLAUCONIA EMINI.

loreal postocular

post.nasal =; preocular .-” teraial

ant.nasal i Jee TES sees AEERS

eS g a=

rostral ---..- HOO ~~~ post.temporal

upper labial “SCT ESL labial

poison-fang" ect oe ESS ate

symphysial :

subocular 4

rostral ---.... aware |alia)| _-----~" rostral

STEM 5 Nee ne ay OE symphysial

internasal i® lower labial

frontal ~ post chin-shield

supraocular” lower labial

parietal ~~

CAUSUS RHOMBEATUS.

J3.GREEN DEL

SNAKES OF EAST AFRICA. 613

Text-figure 2.

(From P. Z.S. 1915, p. 195.)

WIXI

Hs eine,

aX x AKA

ventrals scales (15 rows) ventrals scales (21 raws)

Scaling of thickest part of body.

A. Gastropyxis smaragdina, with keeled scales and bicarimate ventral shields.

B. Dipsadomorphus blandingit, with oblique scales and enlarged vertebrals.

The following map (text-fig. 3) shows the boundaries of the

divisions adopted in this series of papers. Divisions IT and III

have been dealt with, the present paper treating of Division

IV :—

Text-figure 3.

Division of Africa into seven districts.

614 MR. G. A. BOULENGER ON THE

Synopsis of the Species.

I. No preocular; no subocular; ocular in contact with two

upper labials ; ; snout with sharp horizontal edge; eyes

hidden ; 22 scales round middle of ae the diameter of

which is 80 times in total length.. ee exopncoose — Lab GiRaxedISs

II. A preocular; no subocular; ocular in contaen) t with two or three upper labials.

A. Nostrils lateral; snout rounded; eyes distinct.

20 scales round middle of body, the diameter of which is 35 to

50 times in total length .. .. TL. braminus.

24 scales round middle of body. 5 ‘the diameter of which is 50 to

60 times in total length.. SenbbaGReHooS cus adecnouGenaeaancnosca LAS VDC TOO LUIS:

B. Nostrils inferior; snout Sethe or with ances angular horizontal edge.

1. Eyes hidden.

22 or 24 scales round middle of body, the diameter of which

is 40 to 50 times in total length ............-.0.ccececeeeee eee eee T. obtusus.

26 scales round middle of body, the diameter of which is

25 times in total Jength.. . TL. tornieri.

34 scales round middle ‘of body, ‘the diameter of which is

23 times in total length.. slg uaiclacleve @oiee nective cesesoeecall MEL ERTILOILLETUSUS

2. Hyes distinct.

a. A small shield separating the preocular from the

upper labials; 28 scales round middle of body,

the diameter of which is 50 times in total

Venoth: cites th soocannadcce — LAL, (OGTR

b. Preocular in samiadt oath a upper Slabiale

Nasal completely divided: 24 scales round middle of body, the

diameter of which is 30 times in total length ............... T. mossambicus.

Nasal incompletely divided, the cleft not quite reaching the

rostral; 24 scales round middle of body, the diameter of

which is 37 or 38 times in total length . . T. tettensis.

Nasal incompletely divided, the cleft not reaching ‘the rostr al;

26 to 30 scales round "middle of body, the diameter of

which is 24 to 30 times in total length . eee DIL IECELLUSS

C. Nostrils inferior; snout with sharply heals horizontal edge; eyes distinct.

Lower surface of rostral broader than long; 38 to 44 scales

round middle of body, the diameter of which is 25 to 32

times in total length ...... . TT. schlegelii.

Lower surface of rostral broader than long ; “34 to 40 scales

round middle of body, the diameter of which is 42 to 46

times in total length . . T.dinga.

Lower surface of rostral as long a as br oad; 30 to 38 scales round

middle of body, the diameter of which is 25 to 37 times in

total length ......... Recess eens sonjecdane cones, C45 TKO PUIO

III. A preocular ; Aes Sa eed ‘pon ithe upper labials by a larger

subocular ; diameter of body 50 to 63 times in total length.

Snout rounded ; eyes hidden ; 22 scales round middle of body. 1. pallidus.

Snout with sharp horizontal edge; eyes hidden; 18 scales

round middle of body... sesseeeeeee LD. lumbriciformis.

Snout with sharp horizontal edges eyes “distinct ; “24. scales

round middle of body ... Baisakivehcoelcslisssisetyarieeteetecer nee) Me amUMeECe ILE ESS

1. Typuiops Gracitis Sternf. Mitth. Zool. Mus. Berl. v. 1$10,

p. 70.

Urungu, German East Africa.

2. TYPHLOPS BRAMINUS Daud.

Bouleng. Cat. Sn. i. p. 16.

Southern Asia ; islands of the Indian Ocean (including Mozam-

bique island); South Africa; Lagos ; Mexico.—Distribution

probably ascribable to transport by human agency.

SNAKES OF EAST AFRICA. 615

3. TYPHLOPS PLATYRHYNCHUS Sternf. Mitth. Zool. Mus. Berl. v.

1910, p. 69. |

Tanga, German East Africa.

4, Typuuors oprusus Peters, Mon. Berl. Ac. 1865, p. 260, pl. —,

fig. 2; Bouleng. t. c. p. 38.

Nyassaland.

5. TYPHLOPS TORNIERI Sternf, Mitth. Zool. Mus. Berl. v. 1910,

p. 69.

Kilimanjaro.

6. TYPHLOPS MANDENSIS Stejneg. Proc. U.S. Nat. Mus. xvi.

1894, p. 725; Bouleng. Cat. Sn. in. p. 587.

Island of Manda, north of Lamu.

7. TYPHLoPS GIERR& Mocquard, Bull. Mus. Paris, 1897,

Peela2e

Tanga, German Hast Africa.

8. T'YPHLOPS MOSSAMBICUS.

Onychocephalus mossambicus Peters, Mon. Berl. Ac. 1854,

p. 621.

Typhlops mossambicus Peters, Reise Mossamb. 111. p. 93, pl. xv.

fig. 2; Bouleng. Cat. Sn. 1. p. 41.

Portuguese Hast Africa and Zululand.

9. 'TYPHLOPS TETTENSIS.

Onychocephalus tettensis Peters, Mon. Berl. Ac. 1860, p. 80.

Typhlops tettensis Peters, Reise Mossamb. ii. p. 92, pl. xv.

fig. 1; Bouleng. Cat. Sn. i. p. 41.

Portuguese Hast Africa.

10. TypHiors puncratus Leach.

Bouleng. Cat. Sn. i. p. 42.

Typhlops bocagii Bethencourt Ferreira, Jorn. Sc. Lisb. (2) vi.

1904, p. 114.

Typhlops adolphi Sternf. Mitth. Zool. Mus. Berl. v. 1910, p. 70.

Tropical Africa.

11. TyPHLoPs SCHLEGELIZ Bianconi, Spec. Zool. Mossamb. p. 13,

pl. ii. fig. 2; Bouleng. Cat. Sn. i. p. 44.

East and Central Africa to Portuguese East Africa and

Southern Rhodesia.

12. TYPHLOPS DINGA.

Onychocephalus dinga Peters, Mon. Berl. Ac. 1854, p. 620.

Typhlops dinga Peters, Reise Mossamb. ii. p. 98, pl. xiv. fig. 1

& pl. xiv. A, fig. 3; Bouleng. Cat. Sn. i. p. 45.

Portuguese Hast Africa.

616 MR. G. A. BOULENGER ON THE

13. TYPHLOPS MUCRUSO.

Onychocephalus mucruso Peters, Mon. Berl. Ac. 1854, p. 621.

Typhlops mucruso Peters, Reise Mossamb. iii. p. 95, pl. xiii.

fig. 3; Bouleng. Cat. Sn.i. p. 46.

Typhlops humbo Bocage, Jorn. Se. Lisb. xi. 1886, p. 171;

Bouleng. 1. c.

Typhlops hottentotus Bocage, op. cit. (2) iii. 1898, p. 117.

Typhlops latirostris Sternf. Mitth. Zool. Mus. ‘Berl. Ve. SIUC.

D200.

Tropical Africa south of the Equator.

14, TYPHLOPS PALLIDUS.

Letheobia pallida Cope, Proc. Ac. Philad. 1868, p. 322.

Typhlops pallidus Bouleng. Cat. Sn. i. p. 54.

Zanzibar and Pemba Id.

15. TYPHLOPS LUMBRICIFORMIS.

Onychocephalus (Letheobia) lumbriciformis Peters, Mon. Berl.

Ac. 1874, p. 377.

Typhlops lumbriciformis Bouleng. Cat. Sn. i. p. 54.

Vyphlops kleebergi Werner, Zool. Anz. xxvii. 1904, p. 664.

East and Central Africa,

16. TyPpHLops unirznratus Peters, Mon. Berl. Ac. 1878,

p. 205, pl. ii. fig. 5; Bouleng. Cat. Sn.i. p. 55, and Ann. Mus.

Genova (3) v. 1892, p. 331 (var. ateniatus).

Somaliland and British East Atvica.

Family GLAUCONIIDS.

A single genus.

1. GLAUCONTA.

Gray, Cat. Liz. p. 139; Bouleng. Cat. Sn. i. p. 59.

Synopsis of the Species.

I. Ocular bordering the mouth ; tail at least three times as long as broad.

A. Rostral more than half width of head, extending backwards beyond level

of eyes.

Diameter of body 50 to 65 times in total length........................ G. distanti.

Diameter of body 80 to 87 times in total length............000.cc G. merkeri.

B. Rostral one third to one half width of head, not extending backwards

beyond level of eyes.

1. Rostral not in contact with supraocular.

Diameter of body 50 to 55 times im total length........................ G. emini.

Diameter of body 30 times in total length ...................000... G. boulengeri.

pune of body 70 times in total length SM eet cea Gare: . G. longicauda.

. Rostral in contact with supraocular ; sPannetiot oF Taaaky

Az to 6O/times im total length. oi c.e. eee e se cee eee cee G. conjuncta.

II. Ocular not bordering the mouth; tail hardly twice as long

as broad ; diameter of body 40 times in total length ......... G. braueri.

SNAKES OF EAST AFRICA. 617

1. GLAUCONIA DiIsTANTI Bouleng. in Distant, Natur. Transvaal,

p. 175, fig., and Cat. Sn. 1. p. 62.

Nyassaland, S. Rhodesia, Transvaal.

2. GLAUCONIA MERKERI Werner, Jahresh. Ver. Nat. Wirttemb.

Iba ISOS), fe 1a

Moschi, German E. Africa.

3. GLAUCONIA EMINI Bouleng, Ann. & Mag. N. H. (6) vi. 1890,

p- 91, and Cat. Sn. i. p. 64, pl. 111. fig. 8.

Kast and Central Africa.

4. GLAUCONIA BOULENGERI Boettg. in Voeltzkoff, Reise Ostafr.

li. p. 354.

Witu, German E. Africa.

5. GLAUCONIA LONGICAUDA.

Stenostoma longicauda Peters, Mon. Berl. Ac. 1854, p. 621, and

Reise Mossamb. iii. p. 102, pl. xv. fig. 5.

Glaucoma longicauda Bouleng. Cat. Sn. i. p. 66.

Portuguese H. Africa and N. Rhodesia,

6. GLAUCONIA CONJUNCTA.

Stenostoma conjunctum Jan, Arch. Zool. Anat. Phys. 1. 1861,

p. 189, and Icon. Gen. |. 2, pls. v. & vi. fig. 9.

Glauconia conjuncta Bouleng. t. c. p. 67. —

South and East Africa.

7. GLAUCONIA BRAUERI Sternf. Mitth. Zool. Mus. Berl. v. 1910,

p. 70.

Bagamoyo, German E. Africa,

Family Borp &,

Two genera :—

Upper surface of head with shields; rostral and anterior upper labials

deeply pitted ; subcaudals in two VOWS .............c.ceceeeeeeeeeeeesereeeee. Python.

Upper surface of head with small scales; tail very short, subcaudal

SIVYEE (Sa) dempeanegacedas DA SeRE ROR BRRCAO SSE AEAABR eer IE ad CHUCCNEREM RO Gos CHa Gon IMM OF gen

1. PyrHon.

Daud. Hist. Rept. v. p. 266; Bouleng. Cat. Sn. i. p. 85.

|. PyvHON SEB.

Coluber sebe Gmel. 8. N.1. p. 1118.

Python sebe Bouleng. t. c. p. 86.

Tropical and South Africa,

618 MR. G. A. BOULENGER ON THE

2. Hryx.

Daud. Hist. Rept. vil. p. 251; Bouleng Cat. Sn. i. p. 122.

1. Eryx rHEBAIcUS Reuss, Mus. Senckenb. i. 1834, p. 134;

Bouleng. t. c. p. 125,

Upper Egypt to German East Africa.

Family COLUBRID &.

Three parallel series :—

No poison-fangs ; all the teeth solid........................... A. Aglypha.

Poison-fangs behind B. Opisthoglypha.

Poison-fangs in front ..... C. Proteroglypha.

A. Aglypha.

I. Loreal present (occasionally absent in Homalosoma, recognisable by the com-

bination of a single nasal shield, paired subcaudals, and a low number of

ventral shields, 1138-144).

A. Snout without angular horizontal edge; internasal and prefrontal paired.

1. Scales in 19 rows or more, not oblique; body not remarkably slender.

a. Pupil round.

a. A single anterior ey

Scales in 19 rows; anal divided ........... ... Tropidonotus.

Scales in 19-25 rows; anal entire; a a deep groove d on side of head,

above upper Te isis ca Rea emai eA a RE Ay Glypholycus.

(3. Two or three superposed anterior temporals.

Scales keeled, in 23 rows; a deep a between nasal and

preocular as seus Bothrophthalmus.

Scales smooth or very -obtusely keeled, in 25 to 31 rows ......... Pseudaspis: zi

Scales smooth, in 21 rows. ........ sotddéoceoss (Clomraqeiiict:

6. Pupil eae alipties Tadales cinogehe in 23 to

33 YOWS ....... anpadong cesses JOOOTKGD.

2. Scales in 138 to 17 rows, or, if i in io, autremely narrow and oblique.

a. Pupil vertically elliptic; snout much flattened ; scales in 15 or 17 rows.

Scallestsmrootihe avescmaccnstinssuisaeiito- <honseeeae ioe nabs Sagwedsecnaece anne Lycophidium.

Scales keeled, vertebrals bicarimate ............ Tessas oe. Nemocephalus.

6. Pupil round; body usually very debe (Ce snakes).

a. Scales in 138 or 15 rows, smooth.

Subcaudals not keeled . Chlorophis.

Subcaudals keeled and with a notch corresponding: to the. keel,

same as on the ventrals 2.0.00... ....c2s1e vec seccteecececteceseastee Philothamnus.

3. Scales in 15 rows, ieselad

Subcaudals keeled and notched ......... .. Gastropyxis.

Subcaudals not keeled ; two super ‘posed. anterior temporals .. Hapsidophrys.

y. Scales in 17 or 19 rows, very narrow, keeled; eye very large.

Juateral scales shorter tham dorsals) <0... 22. 5.0.ccccecce sce cen eeeeen es Thrasops.

Lateral scales as long as dorsals............... Soest sees sen | Leanniophas:

c. Pupil round; nostril in a Aaneles or eenmuided nasal; scales not oblique,

short and smooth, in 15 or 7 rows.

Nostril directed upwards, nasal divided or semidivided; two

superposed anterior temporals ; ventvals 145 or more ...... Grayia.

Nostril lateral, nasal entire ; a single anterior temporal ; ventrals

USS] skein Eas aca dase cocanbedecpacdadesacashebosonidekeioad. Jahounalaxonnay

SNAKES OF EAST AFRICA. 619

-B. Snout with angular horizontal edge, or internasal and prefrontal single.

Eye in contact with labials; internasal and prefrontal single ;

scales in 15 or 17 rows; ventrals less than 170 ............... Prosymna.

Suboculars separate the eye from the labials ; scales in 19 to 25

ROWSE WOMANS TG TREN I70).ccn5cnq6s6oa8 Son pou soo onde OO Scaphiophis.

II. No loreal, nasal in contact with preocular ; pupil vertically

elliptic; scales strongly keeled, some of the laterals very

@OUETEE | sos bbeondoeccon oxo ond dod eoanAd Radmsd ang eos Reb AEDeMATnARCeoosbeBine HOURLY ALUAa.

1. TRoPIDONOTUS.

Kuhl, Bull. Sc. Nat. i. 1824, p. 81; Bouleng. Cat. Sn. i.

pe Loe

1. TROPIDONOTUS OLIVACEUS.

Coronella olivacea Peters, Mon. Berl. Ac. 1854, p. 622.

Tropidonotus olivaceus Bouleng. t. ¢. p. 227.

Grayia giardi Dollo, Bull. Mus. Belg. iv. 1886, p. 158, fig. ;

Bouleng. op. cit. 11. p. 288.

Tropical Africa, from the Soudan to Angola, and Southern

Rhodesia.

2. GLYPHOLYCUS.

Giinth. P. Z.S. 1893, p. 629; Bouleng, Cat. Sn. iii, p. 615.

Two species :—

Scales in 23 or 25 rows; nasal semidivided ............:.:.....00000s00:000. 2 = G bicolor

Sentloss rn ID srogyig mmseall GUIMCE CL adosas dds odo cnnaiy seo beasencudecndeb se cscncann | (Ere @HDORDOs

1. GiypHoLycus BicoLor Giinth. |. ¢., fig.; Bouleng. 1. ¢.

Lake Tanganyika.

2. GuypHoLtycus wuHytTm Bouleng. P. Z. 8. 1897, p. 802,

pl. xlvinifig. 2. .

Nyassaland.

3. BoTHROPHTHALMUS.

Peters, Mon. Berl. Ac. 1863, p. 287; Bouleng. Cat. Sn. 1.

p. 324,

1. BoTtTHROPHTHALMUS LINEATUS.

Elaphis (Bothrophthalmus) lineatus Peters, 1. e.

Bothrophthalmus lineatus Bouleng. 1. c.

From the Gold Coast and Uganda to the Congo.

4. Boopon.

Dum. & Bibr. Mém. Ac. Se. xxiii. 1853, p. 460; Bouleng. Cat.

Sn. i. p. 327.

Synopsis of the Species.

T. Subcaudals in two rows.

Preocular uot extending to upper surface of head; scales in 23 or

25 rows ; ventrals 175-195; no light lines on side of head ...... B, infernalis.

620 MR. G. A. BOULENGER ON THE

Preocular extending to upper surface of head; scales in 23 (rarely

25) rows ; ventrals 186— 220; belly blackish brown with the

middle ne SOULOG DISA. Woasopa.sac oaocoaeen sue eeanentenorp ann denone seoanne tes B. virgatus.

Preocular extending to upper surface of head; scales in 25 to 33

rows ; ventrals BUSS DD AO) ee a ce econ enh ae ea Ge en eS aL CAPE CUC ILS

II. Subcaudals single ; scales in 25 to 29 rows: oe 183-221... B. olivaceus.

1. Boopon INFERNALIS Giinth. Cat. Col. Sn. p. 199; Bouleng.

OMI, S05 TG jos Bai), oll, kexts whe ae

German Kast Africa and South Africa.

2. BooDON VIRGATUS.

Ocelopeltis virgata Hallow. Proc. Ac. Phiiad. 1854, p. 98.

Boodon virgatus Bouleng. t. c. p. 331.

German East Africa (fide Sternfeld); West Africa, from the

Gold Coast to Calabar.

3. Boopon tinEatus Dum. & Bibr. Erp. Gén. vii. p. 363;

Bouleng. t. ¢. p. 332.

Tropical and South Africa and South Arabia.

4, BooDON OLIVACEUS.

Holuropholis olivaceus A. Dum. Rev. et Mag. Zool. 1856,

p- 466.

Boodon olivaceus Bouleng. t. ¢, p. 335.

West and Central Africa, from Nigeria and Uganda to the

Congo.

5. LycoPpHIDIUM.

Dum. & Bibr. Mém. Ac. Se. xxiii. 1853, p. 462; Bouleng. Cat.

Snes paad0.

Synopsis of the Species.

I. Scales in 15 rows ; ventrals 153-165; subcaudals 23-32 ... LZ. meleagris.

Il. Scales in 17 rows.

A. Parietal shields longer than the distance between frontal and end of snout.

Diameter of eye hardly equal to its distance from mouth ; ventrals

140-150 ; subcaudals 18-28 . . LL acutirostre.

Diameter of eye considerably oveater “than its distance from

mouth ; ventrals 146-156 ; subcaudals 20-30 ......... . LL semianniulis.

Diameter of eye considerably greater than its distance from

mouth; ventrals 163-208 ; Subeaudals 21°47 ene LL. capense.

Diameter of eye not greater than its distance from mouth ;

ventrals 164-189 ; Weubcnudales2o233. .<... wie ee LL jacksonii.

B. Parietal shields not longer than distance between frontal

and end of snout; ventrals 188-219; subcaudals

ISIE Sea ee gh eee ern Ren. Shae RRL cee RS CLEC LILCLUUIIEG

1. LycopHrpium MELEAGRIS Bouleng. Cat. Sn. i. p. 337, pl. xxi.

fig. 2.

German East Africa (fide Sternfeld); Angola.

2. LycopHipiuM AcuUTIROSTRE Giinth. Ann. & Mag. N. H. (4) 1.

1868, p. 427, pl. xix. fig. D; Bouleng. t. c. p. 338,

Zanzibar.

SNAKES OF EAST AFRICA. 621

3. LYCOPHIDIUM SEMIANNULIS ae Mon. Berl. Ac. 1854,

p. 622, and Reise Mossamb. ii +e 135, pl. xvi. fig. 2; Bouleng.

PANES [On Sais)

Portuguese Hast Africa.

4, LyYCOPHIDIUM CAPENSE.

Lycodon capensis A. Smith, 8. Afr. Quart. Journ. (1) no. 5,

UerIl5 joe sk :

Lycophidium capense Bouleng. t. c. p. 339.

Tropical and South Africa.

5. LycopHrpium sAcKsoNntI Bouleng. Cat. Sn. i. p. 340, pl. xxi.

fig. 3.

Kast Africa (Kilimanjaro, Lamu).

6. LycopHipDIuM sEmiIcINcTuM Dum. & Bibr. Erp. Gén. vii.

p. 414; Bouleng. t.c. p. 341.

German East Africa (fide Sternfeld); Senegambia, French

Guinea, Northern Nigeria.

6. SIMOCEPHALUS.

Ginth. Cat. Col. Sn. p. 194; Bouleng. Cat. Sn. i. p. 344.

Synopsis of the Species.

I. Ventrals 203-255 ; subcaudals 45-70.

Two) postoculars; a yellow vertebral line’ .............ccues-nsese--s-ss----s. 9S» COPENStS.

Three postoculars ; pi ete) ONHEA | Gasancooeon nos ooussnandtnoscncceudassdcesdos - Ise Qoanoleines

Three postoculars ; two superposed Torealet en ans en Secunia lars

Il: Ventrals 289-262; subcaudals 75-124 2.0... ......c.s.cecensens--s--s-. Se POENSIS.

Ill. Ventrals 171-178; -subcaudals 62-63 .................00.ccce eevee ces Se NYASSE.

_], SEMOCEPHALUS CAPENSIS.

Heterolepis capensis A. Smith, Ill. Zool. 8. Afr., Rept. pl. lv.

Simocephalus capensis Bouleng. t. c. p. 345.

German and Portuguese East Africa, Nyassaland, Gaboon,

Natal.

2. SIMOCEPHALUS CHANLERI Stejneg. Proc. U.S. Nat. Mus. xvi.

1894, p. 726; Bouleng. op. cit. iii. p. 617.

Isiand of Manda, N. of Lamu.

3. SIMOCEPHALUS UNICOLOR Bouleng. Ann. & Mag. N. H. (8) v.

1910, p. 512.

Kenya District, Brit. E. Africa, .

4, STMOCEPHALUS POENSIS.

Heterolepis poensis A. Smith, Il. Zool. 8. Afr., Rept.

Simocephalus poensis Bouleng. op. cit. i. p. 346.

Uganda; Sierra Leone to Congo... . ecg Eat

Proc. Zoou, Soe.—1915, No. XLITI. 43

MR. G. A. BOULENGER ON THE

5, SIMOCEPHALUS Nyass# Giinth. Ann. & Mag. N. H. (6) i.

1888, p. 328; Bouleng. t. c. p. 347, pl. xxii. fig. 2.

Gonionotophis degrijsit Werner, Zool. Anz. xxx. 1906, p. 53.

Kast Africa, Nyassaland, Natal.

7. PsEUDASPIS:

Cope, Proc. Ae. Philad. 1864, p. 168; Bouleng. Cat. Sn.

p. 373.

1. BsEuDASPIS CANA.

Coluber canus Linn. Mus. Ad. Frid. i. p. 31, pl. x1. fig. 1.

Pseudaspis cana Bouleng. 1. c.

East and South Africa, Nyassaland, Angola.

8. CHLOROPHIS.

Hallow. Proe. Ac. Philad. 1857, p. 52; Bouleng. Cat. Sn.

iil Powe

Synopsis of the Species.

I. No trace of ventral keels ; ventrals 147-190.

A. Scales in 15 rows.

9 upper labials, 4th, 5th, and 6th entering the eye ; subcaudals

103-123..

7 upper labials, 3rd, 4th, and 5th entering the eye 5 5 subcaudals

114.

8 upper Inbials, ‘Ath and 5th entering ‘the eye; ; subcaudals 82—

105..

183. chiens in 13st YOws ; io upper labile 5th ‘aud 6th onitie

the eye; subcaudals 75

II. Ventrals with a more or less distinct lateral keel.

7 or 8 upper labials, 4th and 5th (rarely 3rd and 4th) entering

the eye; ventrals 148-169; subcaudals 71-114...............

8 or 9 upper labials, 4th, 5th, and 6th (rarely 3rd, 4th, and 5th)

entering the eye; ventrals 175-190; subcaudals 115-135 ;

body very slender anteriorly ..

9 upper labials, 4th, 5th, and 6th enteri ing > the eye; preeocular

in contaet with or narrowly separated from the frontal ;

ventrals 150-182; subcaudals 90-138 ...............0...00. 22

1. CHLOROPHIS EMINI.

C. emini.

C. schubotzi.

C. hoplogaster.

C. macrops.

C. neglectus.

C. heterolepidotus.

C. irregularis.

Ahetulla emini Giinth. Ann. & Mag. N. H. (6) i. 1888, p. 325.

Chlorophis emini Bouleng. t. c. p. 92, pl. v. fig. 1.

Kgyptian Soudan to Uganda and Ruwenzori.

2. CHLOROPHIS scHUBOTZI Sternf. in Schubotz, Wiss. Ergebn.

Deutsch. Z.-Afr. Exped. iv. Zool. 1. p. 269, fig. (1912).

Near Bukoba, German E. Africa.

3. CHLOROPHIS HOPLOGASTER.

Ahetulla hoplogaster Giinth. Ann. & Mag. N. H. (3) xi. 1863,

p. 284.

Chlorophis hoplogaster Bouleng. t. c. p. 93.

Central, East, and South Africa.

SNAKES OF EAST AFRICA, 623

4, CHLOROPHIS MACROPS.

Oligolepis macrops Bouleng. op. cit. ili. p. 644.

Chlorophis macrops Sternf. Sitzb. Ges. Nat. Fr. Berl. 1908,

joe De

German East Africa.

5. CHLOROPHIS NEGLECTUS.

Philothamnus neglectus Peters, Mon. Berl. Ac. 1866, p. 890,

and Reise Mossamb, iii. p. 130, pl. xix. a, fig. 2.

Chlorophis neglectus Bouleng. op. cit. ii. p. 94.

Kast and Central Africa, S. Rhodesia.

6. CHLOROPHIS HETEROLEPIDOTUS.

Ahetulla heterolepidota Giinth. Ann. & Mag. N. H. (3) xi. 1863,

p. 286.

Chlorophis heterolepidotus Bouleng. t. ¢, p. 95.

Chlorophis gracilis Sternf. Mitth. Zool. Mus. Berl. v. 1910,

p. 64.

Tropical Africa.

7. CHLOROPHIS IRREGULARIS.

Coluber wiregularis Leach, in Bowdich, Miss. Ashantee,

p. 494.

Chlorophis irregularis Bouleng. t. ¢. p. 96.

Senegambia and Uganda to Damaraland and $8. Rhodesia.

9. PHILOTHAMNUS.

A. Smith, Ill. Zool. 8S. Afr., Rept.; Bouleng. Cat. Sn. ii. p. 98.

1, PHILOTHAMNUS SEMIVARIEGATUS A. Smith, t. c. pls. lix., Ix.,

& Ixiv. fig. 1; Bouleng. t. c! p. 99.

Tropical and South Africa.

10. GASTROPYXIS.

Cope, Proc. Ac. Philad. 1860, p. 556; Bouleng. Cat. Sn. ii.

pe LO:

Two species :—

Temporals 1+2; scales strongly keeled 00.0.0... eee 6G smaragdina.

Temporals 2+2; scales feebly keeled......... 00.00... cc. see seeeecee eee eee G. orientalis.

1. GASTROPYXIS SMARAGDINA.

Dendrophis smaragdina Schleg. Phys. Serp. ii. p. 237.

Gastropyxis smaragdina Bouleng. t. c. p. 103.

Tropical Africa, from Sierra Leone and Uganda to the Congo

and Northern Angola.

A3*

G24 MR, G. A. BOULENGER ON THE

2. GASTROPYXIS ORIENTALIS Werner, Jahresh. Ver. Nat. Wiirt-

temb. Ixv. 1909, p. 55.

terman East Africa.

11. Hapsipopurys.

Fischer, Abh. Nat. Ver. Hamb. iii. 1856, p. 110; Bouleng.

Cat. Sn. ii. p. 108.

1. HaprstpopHrys LINEATA Fischer, t.c. p. 111, pl. ii. oe Ie

Bouleng. t. ¢. p. 104.

From the Gold Coast to the Congo, eastwards to Uganda.

12. 'UHRASOPS.

Hallow. Proc. Ac. Philad. 1857, p. 67; Bouleng. Cat. Sn. 11.

p. 104.

}. Turasops roruscuitpit Moequard, Bull. Mus. Paris, 1905,

p. 286.

British East Africa.

13. RHAMNOPHIS.

Giinth. Ann, & Mag. N. H. (3) ix. 1862, p. 129; Bouleng. Cat.

Sn. ui. p. 632.

1. RHAMNOPHIS JACKSONII.

Thrasops jacksonit Giinth. Ann. & Mag. ING Tels (()) sae WW ey

p- 528; Bouleng. 1. e.

’ Uganda, French Guinea, Gold Coast, Kasar.

14. CoRoNELLA.

Laur. Syn. Rept. p. 84; Bouleng. Cat. Sr. i. p. 188.

Two species :—

Rostral much broader than deep .............:.0ecceeceeseseeeereeeeteseueees OL semiornata.

Rostral scarcely broader than deep ....-........0..600eesseeteeeeereeees CL scheffleri.

1. CoRONELLA SEMIORNATA Peters, Mon. Berl. Ac. 1854, p. 622,

and Reise Mossamb. iii. p. 116, pl. xvn. fig. 2; Bouleng. t. c.

p- 195.

East Africa, N. Rhodesia.

2, CORONELLA SCHEFFLERT Sternf Sitzb. Ges. Nat. Fr. Berl.

1908, p. 93. ‘

British East Africa.

SNAKES OF EAST AFRICA. 625

| 15. Grayta.

Gunth, Cat. Col. Sn. p. 50; Bouleng. Cat. Sn. ii. p. 286.

Two species :—

Scales in 17 rows; ventrals 145-168; subcaudals 89-102.................. G. smythii.

Scales in 15 rows; ventrals 130-148; subcaudals 100-128 .............. G. tholloni.

1. GRAYIA SMYTHII. .

Coluber smythii Leach, in Tuckey’s Explor. R. Zaire, App.

jos BOO),

’ Grayia smythii Bouleng. t. c. (part.), and P. Z. 8. +899;

p- 948, figs.

West and Central Africa and Uganda.

2. GRAYIA THOLLONI Mocquard, Bull. Soc. Philom. (8) ix. 1897,

pL Bouleng., P. ZS. soo) p. 951, fig.

orpeiin Soudan, Uganda, Ep ench Congo, Katanga.

16, HoMALOSOMA.

Wagl. Syst. Amph. p. 190; Bouleng. Cat. Sn. ii. p. 273.

1. HoMALosoma LUTRIX.

Coluber lutrix Linn. 8, N. i. p. 375.

Homalosoma lutria Bouleng. t. c. p. 274.

Homalosoma shiranum Bouleng. t. ¢. p. 276.

Homalosoma abyssinicum Bouleng. 1. c.

Hast Africa, Nyassaland, South Africa.

17. PRosyMNA.

Gray, Cat. Sn. p. 80; Bouleng. Cat. Sn. ii. p. 246.

Synopsis of the Species.

Snout with angular horizontal edge; two postoculars; ventrals 131-

IS). aS Hacer and adancia gona Ee dolor nes Moece Me RELA GE LeaE Ura EMME haat men Ae P. ambigua.

Snout with angular horizontal edge; a single noeecas 3 Been

entering the eye; ventrals Tete 167 ito apooanasaacosen darn WOXCRG Id

Snout rounded; two postoculars; ventrals 140-143 . neey aosconws den Gen POGIOCU OG,

1. PRosyMNA AMBIGUA Bocage, Jorn. Se. Lisb. iv. 1873, p. 218;

Bouleng. t. c. p. 248.

Zanzibar Coast to Zululand, N. Rhodesia, Angola.

Prosymna Bocacit Bouleng. Ann. & Mag. N. H. (6) xix.

50. p. 278, fig., and Ann. Mus. Congo, Zool. ii. 1901, p. 9,

pl. iu. fig. 4.

Prosyina vasset Mocquard, Bull. Mus. Paris, 1906, p. 250.

Ubanghi and Mozambique.

626 MR. G. A. BOULENGER ON THE

3. PrRosyMNA VARIABILIS Werner, Jahresh. Nat. Ver. Wiirt-

temb. Ixv. 1909, p. 57,

Moschi, German E. Africa.

18. SCAPHIOPHIS.

Peters, Mon. Berl. Ac. 1870, p. 644; Bouleng. Cat. Sn. ii.

p. 254.

1. SCAPHIOPHIS ALBOPUNCTATUS Peters, t. c. p. 645, pl. i. fig. 4;

Bouleng. 1. c.

Tropical Africa, from the Soudan to the Congo.

19. DAsyPELris.

Wagl. Syst. Amph. p. 178; Bouleng. Cat. Sn. 11. p. 353.

1. DASYPELTIS SCABRA.

Coluber scaber Linn, Mus. Ad. Frid. p. 36, pl. x. fig. 1.

Dasypeltis scabra Bouleng. t. ¢. pe 254.

Tropical and South Africa, nan 8. Arabia.

B. Opisthoglypha.

I. Eye moderate or large, with vertically elliptic pupil; head distinct from neck ;

nostril between two shields ; loreal present (sometimes entering the eye).

A. Nostril directed upwards, between the nasal and the

internasal; parietals broken up into small shields;

scales smooth or obtusely keeled, in 21 rows ............ Pythonodipsas.

B. Nostril between two nasals; scales smooth or faintly keeled.

1. Two superposed anterior temporals; scales oblique.

Scales in 19 or 21 rows, vertebrals not enlarged ..................... Tarbophis.

Scales in 21 to 25 rows, vertebrals enlarged ........................... -Dipsadomorphus.

2. A single anterior temporal.

Scales in 19 rows (rarely 17) ; loreal not entering eye ............ Leptodira.

Scales in 17 rows; loreal entering eye ............ Weer | Chamcetonvuse

II. Eye small, with vertically cine one “heed distinct

from neck ; nasal semidivided, with horizontal cleft ;

scales in 17 or 19 rows. corr an Gree Sane EERE aan eoabe . Hemirhagerrhis.

III. Eye moderate or large, with cae or Siecnenel rane head distinct from

neck; loreal present.

A. Pupil round.

1. Loreal not more than once and a half as long as deep; scales in 17 or 19

rows, not oblique.

Nostril between two nasals; ventrals 128-139; subcaudals

34-44. Pebisnsioiiudees «connie eden See eReee teseanesr wen CRCOCCDSAS.

Nostril in a semidivided ‘nasal ; “internasals shorter than pre-

frontals; ventrals 147-187 ; subcaudals 59-98 ................ Amplorhinus.

Nostril between two nasals and the internasal ; subcaudals less

Gyan W7 On cc escheat ap deny caansteeeone enero scatelscc eee CORRE eee eee Trimerorhinus. .

Nostril between two pee: rostral ah large ; subcaudals 90

or more ......... leesssseeeeee Rhamphiophis.

2. Loreal at tease, once sce a nae as loner as deep. ; scales more or less oblique.

Scales in 17 rows; a single anterior temporal........ .... Dromophis.

Scales in 11 to 17 rows; one or two middle ‘maxillary ‘teeth

much enlarged, fang-like... sensi cigs eas oeeteetete tatoe Ste acters LP RERSCUTNIZOD UGS.

SNAKES OF EAST AFRICA. 627

3. Loreal not more than once and a half as long as deep ;

nostril in an undivided nasal; scales very narrow,

oblique, keeled, in 19 or 21 rows.. eee eee Despholiduss

B. Pupil horizontal; nostril in an eecedel bietle scales

narrow, oblique, feebly keeled, in 19 rows.. tisleteee Lhelotornis:

IV. Eye small or very small ; head not at all aeteee es om ree no loreal.

A. Subcaudals in two rows. ‘

1. One or two upper labials in contact with the parietal ; nasal in contact

with the rostral.

Scales in 19 or 21 rows; no preocular .. : ...... Calamelaps.

Scales in 15 or 17 rows; no preocular ; rostral very ‘large, con-

cave below; a pair of internasals anda pair of pretrontals. Rhinocalamus.

Scales in 17 rows; a large, elongate preocular; rostral very

large, flat below; prafrontals absent..................sseeseeen ee Xenocalamus.

2. Temporals separate the upper labials from the parietals; scales in 15 rows.

Nasal in contact with the rostral; no prwocular..................... Micrelaps.

First upper labial in contact with the internasal; a preocular... Miodon.

B. Subcaudals single; a preocular; scales in 15 rows ....... Aparallactus.

1. GEODIPSAS.

Bouleng. Cat. Sn. 11. p. 32.

1. GEopIpsAs vAUEROCEG# Tornier, Zool. Anz. xxv. 1902,

p. 703.

Usambara, German E. Africa.

2. PyTHONODIPSAS.

Ginth. Ann. & Mag. N. H. (4) i. 1868, p. 425; Bouleng. Cat.

Sn. 111. p. 45.

1. PyrHonopipsas CARINATA Giinth. t.c. p. 426, pl. xix. fig. K ;

Bouleng. 1. ¢.

Zambesi, Damaraland.

3. ‘TARBOPHIS.

Fleischm. Dalm. nov. Serp. Gen. p. 17; Bouleng. Cat. Sn. i.

p- 47.

Two species :—

Scaleshmnell Firowsrmanalecivid edie ener eneneens seeder eee sei dscertsrerae T. semiannulatus.

Scales in 21 rows (rarely 19); anal entire...................0:00:0. TZ. guentheri.

1. TARBOPHIS SEMIANNULATUS.

Telescopus semiannulatus A. Smith, Ll. Zool. 8. Afr., Rept.

pl. Ixxi.

Tarbophis semiannulatus Bouleng. t. c. p. 51.

Central and East Africa, Angola, 8S. Africa.

2. TARBOPHIS GUENTHERI Anders. Proc. Zool. Soc. 1895,

pl. xxxvi. fig. 3; Bouleng. t. c. p. 52

East Africa, Arabia, Syria.

628 MR. G. A. BOULENGER ON THE

4, DrIpsADOMORPHUS.

Fitzing. in Tschudi, Faun. Per., Herp. p.55; Bouleng. Cat.

Sn. ill. p. 99.

1. DrpsADOMORPHUS BLANDINGII.

Dipsas blandingii Hallow. Proc. Ac.’ Philad. 1844, p. 170.

Dipsadomorphus blandingti Bouleng. t. ¢. p. 77.

British East Africa; West Africa, from Senegambia to Congo.

5, LmpropirRA. .

Giinth. Cat. Col. Sn. p. 165; Bouleng. Cat. Sn. ii. p. 88.

Synopsis of the Species.

J. Ventrals less than 200; subcaudals less than 60; 3 to 5 pairs of chin-shields.

One prxocular (rarely divided), separated from the frontal ; frental rd

considerably broader than deep; loreal not or but little longer

HIE) GL} ee epee meee dconconsoadtocnecunetes Scdoestdnccomsijooduaecoatke, Jon LOAM,

One preocular, separated from the frontal; rostral little broader

than deep ; loreal longer than deep ..............0.ceceec eee LD. degeni.

Two preoculars, upper in contact with the frontal ..................... LL. tornieri.

II. Ventrals more than 200; subcaudals 100 or more; two pairs

TINT TIERS OTE) (hoe ees te merlies Roy EUS WE On. eeeneeda neoangsnye 2 Oty ORE OCI- Us

1. LeproprrRA HOTAMBGIA.

Coronella hotambeia Laur. Syn. Rept. p. 85.

Leptodira hotambeia Bouleng. t. c. p. 89.

Tropical and South Africa.

2. LepropIRA DEGENI Bouleng. Proc. Zool. Soc. 1906, 11. p.572, fig.

Leptodira attarensis Werner, Sitzb. Ak. Wien, exvi. 1907,

p. 1875.

Uganda.

3. LEProDIRA TORNIERI Werner, t.c. p. 1876.

Usambara, German Hast Africa.

4. LEPTODIRA WERNERI Bouleng. Ann. & Mag. N. H. (6) xix.

1897, p. 281.

Dipsadomorphus reticulatus Werner, Jahresh. Nat. Ver, Wirt-

temb. Ixv. 1909, p. 55.

Usambara, German East Africa.

6. CHAMATORTUS.

Giinth. Proc. Zool. Soc. 1864, p. 310; Bouleng. Cat. Sn.iu. p. 98.

1. CHAMaAToRTUS AvULICUS . Gunith, 1) e., pl. saat hes 2

Bouleng. I. ec.

German and Portuguese Hast Africa, Transvaal.

baer!

SNAKES OF EAST AFRICA. §29

7. HeMrRHAGERRHIS.

Boettg. Zool. Anz. 1893, p. 119; Bouleng. Cat. Sn. i. p. 119.

1. HEMIRHAGERRHIS KELLERI Boettg. |. c.; Bouleng. 1. ¢.

Somaliland and British East Africa.

8. AMPLORHINUS.

A. Smith, Ill. Zool. 8. Afr., Rept.; Bouleng. Cat. Sn. ii. p. 124.

Two species :—

Yemporals 1+2 or 2+2: ventrals 154-187 ..............ccceeeeee ee. A. nototenia.

Temporals 8+4; ventrals 147 0.0.0.0... cece eect eee terre eeeeeee A, CEMatUS.

1. AMPLORHINUS NOTOLENIA.

Coronella nototenia Giinth. P. Z.5. 1864, p. 309, pl. xxvi. fig. 1.

Amplorhinus nototenia Bouleng. t. c. p. 125.

Amplorhinus guntheri Mocquard. Bull. Mus. Paris, 1896,

p. 251.

Egyptian Soudan to Nyassaland ; Angola.

2. AMPLORHINUS THNIATUS Sternf. Mitth. Zool. Mus. Berl. iv.

* 1908, p. 241. ;

Lamu Id., British East Africa.

9. ‘TRIMERORHINUS.

A. Smith, Dll. Zool. 8. Afr., Rept.; Bouleng. Cat. Sn. ii.

p. 138.

Two species :—

Rostral as deep as broad, its upper portion at least half as long as

THES) CUISINES InN VaR TOV) THROM | AOA Aas An aedecdiedlsnconosoamennssasance Lb MUACeROenADIN.

Rostral slightly broader than deep, its upper portion about one 3

thirdiitsidistance trom the Lronbalsekec.scdhsscrsscessesseesese Os waneabuliss

1. TRIMERORHINUS TRITENIATUS.

Rhagerrhis triteniata Giinth. Ann. & Mag. N. H. (4) 1. 1868,

p. 423, pl. xix. fig. H.

Trimerorhinus triteniatus Bouleng. t. c. p. 139.

Kast, Central, and South Africa.

Probably only a variety of the 8. African 7’. rhombeatus.

2. 'TRIMERORHINUS VARIABILIS.

Psammophylax variabilis Giinth. P.Z.8. 1892, p. 557, pl. xxxy.

Trimerorhinus variabilis Bouleng. t. c. p. 140.

Nyassaland.

630 MR. G. A. BOULENGER ON THE

10. RHAMPHIOPHIS.

Peters, Mon. Berl. Ac. 1854, p. 624; Bouleng. Cat. Sn. iii.

p. 144.

Two species :—

Snout rounded; scales in 19 rows; ventrals 230-241; sub-

é CERI EWES) LIEYTSING0 y tatss cd netoce aor eRe eee seeiceinbed cone se R. rubropunctatus.

Snout somewhat hooked in profile; scales in 17 rows; ventrals

148-192; subcaudals 90-110 ..............ccccceseeeeeeeeeeeee. R. oxyrhynchus.

1. RHAMPHIOPHIS RUBROPUNCTATUS.

Dipsina rubropunctata Fischer, Jahrb. Hamb. Wiss. Anst. i.

ISkSi4b, JO Wo Tolls Me sakes By,

Rhamphiophis rubropunctatus Bouleng. t.c. p. 146.

Egyptian Soudan and East Africa.

2. RHAMPHIOPHIS OXYRHYNCHUS.

Psammophis oxyrhynchus Reinh. Vid. Selsk. Skrift. x. 1843,

p. 244.

Rhamphiophis oxyrhynchus Bouleng. }. c.

Tropical Africa.

11. Dromopais. .

Peters, Mon. Berl. Ac. 1869, p. 447; Bouleng. Cat. Sn. 111

p. 149.

1. DROoMOPHIS LINEATUS.

Dryophylax lineatus Dum. & Bibr. Erp. Gén. vii. p. 1124.

Dromophis lineatus Bouleng. 1. c.

Coast of Guinea to Egyptian Soudan and Zanzibar Coast ;

Nyassaland.

12. PsAMMOPHIS.

Boie, Isis, 1827, p. 521; Bouleng. Cat. Sn. i. p. 152.

Synopsis of the Species.

I. Scales in 17 rows; 2 or 3 superposed anterior temporals.

Rostral broader than deep; ventrals 177-192; subcaudals 130-

178 P. punctulatus.

Rota broader. than deep 5 ventrals ii 1805 ‘subcaudals 100-

19 ee : ; ‘ . P. subteniatus.

Rostral as s deep a as broad . ovis feces neeeennee novonesonapovneen, dey GAODIAIS:

II, Scales in 15 rows; 2 ronan anterior ea saWageec see P. biseriatus. ,

III. Scales in 11 rows; a single anterior temporal..................... P. angolensis.

1. PsAMMOPHIS PUNCTULATUS Dum. & Bibr. Erp. Gén. vii.

p- 897; Bouleng. t. c. p. 159.

East Africa, from the Blue Nile to Mozambique ; Arabia.

SNAKES OF EAST AFRICA. 631

2. PSAMMOPHIS SUBTENIATUS.

Psammophis sibilans, var. subteniata Peters, Reise Mossamb.

WHE; Oy LUZl.

Psammophis subteniatus Bouleng. t.c. p. 160.

Kast Africa, from Uganda to Mozambique; Nyassaland.

3. PSAMMOPHIS SIBILANS.

Coluber sibilans Linn. 8. N. 1. p. 383.

Psammophis sibilans Bouleng. t. c. p. 161.

Psammophis trinasalis Werner, Abh. Bayer. Ak. xxii. 1903,

p. 381.

Tropical and South Africa; Egypt.

4. PSAMMOPHIS BISERIATUS Peters, Sitzb. Ges. Nat. Fr. Berl.

1881, p. 88; Bouleng. t. c. p. 168.

Somaliland and British East Africa.

5. PSAMMOPHIS ANGOLENSIS.

Amphiophis angolensis Bocage, Jorn. Sc. Lisb. iv. 1872, p. 82.

Psammophis angolensis Bouleng. t. c. p. 170.

East and Central Africa, Angola, Orange River Colony.

13. 'THELOTORNIS.

A. Smith, Ill. Zool. 8. Afr., Rept.; Bouleng. Cat. Sn. in.

p. 184.

1, THELOTORNIS KIRTLANDII.

Leptophis kirtlandit Hallow. Proc. Ac. Philad. 1844, p. 62.

Thelotornis kirtlandii Bouleng. t. c. p. 185.

Tropical and South Africa.

14. DispHOLIDUs.

Duvernoy, Ann. Sc. Nat. xxvi. 1832, p. 150; Bouleng. Cat.

Sn. 1. p. 186.

1, DIsPHOLIDUS TYPUS.

Bucephalus typus A. Smith, Zool. Journ. iv. 1829, p. 441.

Dispholidus typus Bouleng. t. ¢. p. 187.

Tropical and South Africa.

15. CALAMELAPS.

Ginth. Ann. & Mag. N. H. (3) xviii. 1866, p. 26; Bouleng.

Cat. Sn. ili. p. 245.

Two species :—

Seals ity WY TROWS ona covagp ano angonn opens baa cn nbogsonsa bo caacobaaposoucoces ansans | (Oo Cmdeollor.

Secular tha IS) Ore PAL SREWYS jsn000 400 oosond bon dou 08000 Sound 00d ace quo sqoUoNaseocE coBade. LOD ARANIQCTaIS.

632 MR. G. A. BOULENGER ON THE

1. CALAMELAPS UNICOLOR.

Calamaria wnicolor Reinh. Vid. Selsk. Skrift. x. 1843, p. 226,

pl. 1. figs. 1-3.

Calamelaps untcolor Bouleng. |. e¢.

Atractaspis hildebrandti Peters, Mon. Berl. Ac. 1877, p. 616;

Bouleng. t. c. p. 512.

British East Africa; West Africa.

2. CALAMELAPS POLYLEPIS Bocage, Jorn. Sc. Lisb. iy. 1873,

p- 216; Bouleng. t. c. p. 246.

Nyassaland, Angola.

16. RHINOCALAMUS.

Gunth. Ann. & Mag. N. H. (6) 1. 1888, p. 322; Bouleng. Cat.

Sn. ili, p. 247.

Two species :—

Scalespinl liarows\-| GMupperlalbials eens ereeneeenee eee eeneer Emme LUAICLnzecacleuss

Scalesiimne5)rows: 7 upper labials) eee ye aeons eee eee emEUSieleamase

1, RHINOCALAMUS DimipiATuS Giinth. ]. c pl. xix. fig. C;

Bouleng. 1. ¢.

Kast Africa.

2. RHINOCALAMUS MELEAGRIS Sternf. Mitth. Zool. Mus. Berl. iv.

1908, p. 244.

Lamu Id., British East Africa.

17. XENOCALAMUS.

Giinth. Ann. & Mag. N. H. (4) i. 1868, p. 414; Bouleng. Cat.

Sn. lil. p. 247.

1. XENOCALAMUS BICOLOR Giinth. t.c. p. 415, pl. xix. fig. A;

Bouleng. t. c. p. 248.

Zambesi.

18. MicrEaps.

Boettg. Ber. Senck. Ges. [eg 80, p. 136 ; nieces Cat. Sn.

ill. p. 948.

1. MicrRELAPS BICOLORATUS Sternf. Sitzb. Ges. Nat. Fr. Berl.

IOS pg ae

British East Africa.

19. Mropon.

A. Dum. Arch. Mus. .x.1859, p. 206 ; ae Cat. Sn. 1

p. 249. fae

Synopsis of the Species.

Frontal slightiy broader than supraocular, 14 to 14 as long as

broad.. M. gabonensis.

Frontal much broader than the supraccnlan a little longer ‘than :

broad; nasal entire ... seoprin | LM OTA IS UDe

Frontal much broader than the supraocular, as broad as slong: wr on) IGT vente

SNAKES OF EAST AFRICA. 633

1. MiropcNn GABONENSIS.

Elapomorphus gabonensis A. Dum. Rev. et Mag. Zool. (2) vii.

1856, p. 468.

Miodon gabonensis Bouleng. t. c. p. 2

Old Calabar to Congo ; German East Africa,

Mropon curisryt Bouleng. Ann. & Mag. N. H. (7) xii. 1903,

p. 354.

Uganda.

3. Mropon GRAvERI Sternf. Sitzb. Ges. Nat. Fr. Beri. 1908,

p. 94, and Faun. Deutsch. Kolon. iii. 2, p. 34, fig.

* Uganda.

Perhaps not specifically distinct from the preceding.

20, APARALLACTUS

A. smith, Ill. Zool. S. Afr., Rept. App. p. 15; Bouleng. Cat.

Sn. ui. p. 205.

Synopsis of the Species.

I. Symphysial not in contact with the chin-shields.

A. Two postoculars, in contact with a temporal; nasal entire, in contact with

) ) >

the preocular.

Third and fourth upper labials entering the eye.,................. A. jacksonit.

Second and third upper labials entering the eye.................. A. werneri.

B. A single postocular ; one upper labial in contact with the parietal.

Nasal entire, not in contact with preocular..... .... A. concolor.

Nasal divided, in contact with preocular; “upper. “part “of

rostral about one third its distance from frontal ......... A, lunulatus.

Nasal divided, in contact with preocular; upper part of

rostral two thirds its distance from frontal .................. A, christyi.

II. Symphysial in contact with the chin-shields.

A. Third and fourth upper labials entering the eye.

Nasal divided; swbcaudals 49-59 ..................c1.s022-0teees- Ae guentheri.

Nasal entire; subcaudals 87—58).............ceceeceecceeesceeeee sees. An CApensis.

B. Second and third upper labials entering the eye.

Wemninrallss TOSS) sn s86 ob ono oon sos act suse angdbbe sadeceacdlasnestacaKes, 2k OCKPIAZIS>

Wem WS7ANG® soc cocgas seo quo ane aonsSobosess! obasacadacepnancensccaqcs 24, /OCMIGRUONIMARUUS

1, APARALLACTUS JACKSONIL. ; im

Uriechis jacksonti Gitinth. Ann. & Mag. N. H. (6) i. 1888,

p. 325, pl. xix. fig. E.

Aparallactus jacksonit Bouleng. t. ¢. p. 256.

East Africa.

2. APARALLACTUS WERNERI Bouleng. Ann. & Mag. N. H. (6) xvi.

1895, p. 172, and Cat. Sn. it. p..257, pl.-xt. fig. ie Pi: ee

U Snir German Bast Africa.

634 MR. G. A. BOULENGER ON THE

3. APARALLACTUS CONCOLOR.

Uriechis concolor Fischer, Jahrb. Hamb. Wiss. Anst. 1. 1884,

05 4h Jolls we

Aparallactus concolor Bouleng. Cat. Sn. i. p. 257.

Uganda and Somaliland.

4, APARALLACTUS LUNULATUS.

Uriechis lunulatus Peters, Mon. Berl. Ac. 1854, p. 623, and

Reise Mossamb. 11. p. 113, pl. Xviil. fig. 2.

Aparallactus lunulatus Bouleng. 1s Gs [On PADIS

Lake Tanganyika, Nyassaland, Mozambique.

5. APARALLACTUS CHRISTYI Bouleng. Ann. & Mag. N. H. (8) v.

1910, p. 512.

Uganda.

6. APARALLACTUS GUENTHERI Bouleng. Ann. & Mag. N. H. (6)

xvi. 1895, p. 172, and t. c. p. 259.

Kast and Central Africa, S. Rhodesia, Angola.

7. APARALLACTUS CAPENSIS A. Smith, Ill. Zool. 8. Afr., Rept.

App. p. 16; Bouleng. Cat. Sn. ii. p. 259.

East and South Africa, Katanga.

8. APARALLACTUS NIGRICEPS.

Uriechis nigriceps Peters, Mon. Berl]. Ac. 1854, p. 623, and

Reise Mossamb. 111. p. 111, pl. xviii. fig. 1.

Aparallactus nigriceps Bouleng. t. c. p. 260.

Mozambique and Nyassaland.

9. APARALLACTUS PUNCTATOLINEATUS Bouleng. Ann. & Mag.

N. H. (6) xvi. 1895, p. 173, and Cat. Sn. iii. p. 261.

Nyassaland, Angola.

C. Proteroglypha.

(Loreal absent in all the genera.)

J. Tail compressed ; no distinct ventrals (Marine) .................. HAydrus.

II. Tail not compressed ; ventrals large. 3

A. Head short; snout broader than long; body cylindrical; subcaudals less

than 80.

Scales not at all oblique; ventrals 192-193; subcaudals 67-78 ...... Boulengerina,

Scales more or less oblique, sometimes very slightly ; ventrals 14.1

172; subcaudals 13-25 ...... nooosuccAecnceosnn LONI,

Scales oblique ; ventrals 180-228 ; ; “ subcaudals 50-72 CRC Naia.

B. Head long, narrow; snout not broader than long; body

slightly compressed ; scales very jobliaues ventrals 202-

270; subcaudals 97-121.. Sitecck seeincnnoannooeabucces, » LOXHEKRUS RE

- SNAKES OF EAST AFRICA. 635

1. Hyprus.

Schneid. Hist. Amph. 1. p. 233; Bouleng. Cat. Sn. ii. p. 266.

1. Hyprvus PLATURUS.

Anguis platura Linn. 8. N. 1. p. 391.

Hydrus platurus Bouleng. t. c. p. 267.

Indian and Pacific Oceans.

2. BouLENGERINA.

Dollo, Bul. Mus. Belg. iv. 1886, p. 159; Bouleng. Cat. Sn. iii.

p- 357.

1. BouLENGERINA sTorMsI Dollo, 1. c.; Bouleng. 1. ¢.

Lake Tanganyika.

3. HLAPECHIS.

Bouleng. Cat. Sn. ii. p. 358.

Synopsis of the Species.

Scales in 13 rows ; internasals much shorter than the prefrontals.. EH. guentheri.

Scales in 13 rows; internasals three fourths the length of the

Puce kro Nall set ee ceneeree eee nessa acme ee ate nee oee teeta snek care EL. niger.

Scales in 15 rows; internasals hardly half as long as the pre-

fOTNGALS Heres eeteorea cee sercer seen sess teaa seaatceeaeeanan scene Le Doulenge ri

1. ELAPECHIS GUENTHERT.

Hlapsoidea guenthert Bocage, Jorn. Sc. Lisb. i. 1866, p. 70,

pl. i. fig. 3.

Elapechis guentheri Bouleng. t. ec. p. 359,

Tropical Africa, from the Gaboon and Uganda to Angola and

Nyassaland.

. ELAPECHIS NIGER.

leet nigra Giinth. Ann. & Mag. N. H. (6) i. 1888, p. 332.

Elapechis niger Bouleng. |. c. pl. xx. » fig. i

East Africa, Congo, N. Rhodesia.

3. HELAPECHIS BOULENGERI.

Hlapsoidea boulengeri Boettg. Zool. Anz. 1895, p. 62.

Elapechis boulengert Bouleng. ter ps ole

Zambesi.

4, Nata.

Laur. Syn. Rept. p. 90; Bouleng. Cat. Sn. iii. p- 372.

Synopsis of the Species.

Sixth (rarely seventh) upper labial largest and deepest; eye

separated from the upper labials by suboculars; a single

aniberionybemp orale iacuceuen ce ened cacsuatan seni hal. cea See aa coke ae N. hate.

Sixth upper labial largest and deepest, in contact with lower

postocular, third and fourth Sees the eye; a single

anterior temporal ....... . WN. melanoleuca.

Third upper labial deepest, sixth not in contact with postocular,

third or third and fourth entering the ae 2 or 3 super shee

anterior temporals ........ .. DW. nigricollis.

636 MR. G. A. BOULENGER ON THE

1. NAIA HAIE.

Coluber haie Linn. in Hasselq. Reise Palest. p. 366.

Naia haie Bouleng. t. c. p. 374.

Borders of the Sahara, East Africa southwards to the Trans-

vaal and Zululand; Palestine, Arabia.

2. NAIA MELANOLEUCA. :

Naia haie, var. melanoleuca Hallow. Proc. Ac. Philad. 1857,

pp- 61 & 72. ’

Naia melanoleuca Bouleng. t. c. p. 376.

Tropical Africa, from Uganda and the Gold Coast to Nvassaland

and Angola.

3. Nara nigricoLiis Reinh. Vid. Selsk. Skrift. x. 1843, p. 269,

pl. iil. figs. 5-7; Bouleng: t. c. p. BOF. 37%

Senegambia and Upper Egypt to Bechuanaland and Natal.

5. DENDRASPIS.

Schleg. Vers]. Zool. Gen. Amsterd. 1848 ; Bouleng. Cat. Sn. 111.

p. 434.

Synopsis of the Species.

A large upper temporal, in contact with the whole outer border of

the parietal; scales im 15 to 19 rowWS......... 0.0.0... see seer

Two upper temporals, in contact with the outer border of the

parietal, the second separated from its fellow on the other

side by 3 or more scales or shields; scales in 19 to 23 rows... D. angusticeps.

Two upper temporals, in contact with the outer border of the

parietal, the second separated from its fellow on the other side

by a single shield ; scales in 19 rowS........0... 0002.0 cseeeeeeeee es D. sjoestedti.

D, jamesonii.

1. DENDRASPIS JAMESONIIT.

Elaps jamesonit Traill, in Schleg. Phys. Serp., Engl. Transl.

p- 179, pl. i. figs. 19 & 20.

Dendraspis jamesonii Bouleng. t. c. p. 436.

Tropical Africa, from Uganda and Nigeria to the Congo and

Angola.

2. DENDRASPIS ANGUSTICEPS.

Naia angusticeps A. Smith, Il. Zool. S. Afr., Rept. pl. Ixx.

Dendraspis angusticeps Bouleng. t. ¢. p. 437.

East and Central Africa, Angola, South Africa.

3. DENDRASPIS sJoESTEDTI Lonnb. Wiss. Ergebn. Exped.

Kilimandj.-Meru, Rept. p. 17.

British East Africa.

Perhaps. based on an aberrant specimen of the preceding

species.

Genaien Pear mone

SNAKES OF BAST AFRICA. 637

Family VIPERID &.

Synopsis of the Genera.

1. Eye moderate or large, separated from the upper labials by suboculars; ventrals

less than 180.

Upper surface of head covered with large symmetrical shields;

pupil round mertetcarielostate syacrsteni ate sia

Upper surface of head covered with ‘scales ; ‘nasal in contact with

‘yostral or separated by a single (naso-rostral) shield; pupil

WGI 8 FRIAR NEG ENIS WV LANKO) TRONS -50 conece ana asdosn op bon ene hoanoR boDOHe Vipera.

Upper surface of head covered with scales; nasal separated from the

rostral by small scales ; pupil vertical; subcaudals in two rows. Bitis.

Upper surface of head covered with scales ; pupil vertical; sub-

caudals single; tail prehensile .............. Poet szaitaticameenehiiaey a, AGH ents

Il. Eye minute, with round pupil; upper aes of ica covered

with large symmetrical shields ; no loreal ; a sma!l prxocular

usually present; ventrals TERA cee Wt OR a ee Atractaspis.

Causus.

1. Causus.

Wagl. Syst. Amph. p. 172; Bouleng. Cat. Sn. i, p. 465.

Synopsis of the Species.

T. Scales in 17 rows or more; subcaudals all or greater part in two rows.

Snout obtuse, moderately prominent; ventrals 120-155 ............ C. rhombeatus.

Snout prominent, often more or less distinctly turned up at the

end; scales in 19 to 22 rows; ventrals 134-152 ............... C. resimus.

Snout prominent, more or less turned up at the end; scales in

TPP ROMS 3 WETMTTENS TOONS... por cacconosdascoss.er cop atvarasennccsences — On hahaa aids

IJ. Scales in 15 rows; subcaudals single .........................---.. GG. lichtensteinii.

CAUSUS RHOMBEATUS.

Sepedon rhombeatus Licht. Verz. Doubl. Mus. Berl. p. 106.

Causus rhombeatus Bouleng. t. c. p. 467.

Tropical and South Africa.

2. CAUSUS RESIMUS.

Heterophis resimus Peters, Mon. Berl. Ac. 1862, p. 277, pl. —

fig. 4,

Causus resimus Bouleng. t. c. p. 468.

Kast and Central Africa, Angola.

3. CAUSUS DEFILIPPII.

Heterodon defilippii Jan, Arch. Zool. Anat. Phys. 1. 1862,

p- 225.

Causus defilippti Bouleng. t. c. p. 469.

East and Central Africa, 8. Rhodesia, Transvaal.

4, CAUSUS LICHTENSTEINII.

Aspidelaps lichtensteinii Jan, Rev. et Mag. Zool. 1859, p. 511.

Causus lichtensteinit Bouleng. t. ¢. p. 470.

Tropical Africa, from the Gold Coast and Uganda to the Congo.

Proc. Zoon. Soc.—1915, No. XLIV. A4

638 MR. G. A. BOULENGER ON THE

2. VIPERA.

Laur. Syn. Rept. p. 99; Bouleng. Cat. Sn. ii. p. 471.

Two species :—

No supraocular; anterior nasal separated from the rostral by a

MASO-VOSLIA een cees ee aatem nia oe eae teen cee SER CenU Meme mebreneimen | MUGSRIDEIUCI Ds

A large supraocular; anterior nasal in contact with the rostral... WV. superciliaris.

1. VIPERA HINDII Bouleng. Ann. & Mag. N. H. (8) v. 1910,

Po SIS.

Kenya district, East Africa.

2. VIPERA SUPERCILIARIS Peters, Mon. Berl. Ac. 1854, p. 625,

and Reise Mossamb. tii. p. 144, pl. xxi.; Bouleng. Cat. Sn. i.

p. 491.

German East Africa at Cape Delgado.

3. BItis.

Gray, Zool. Miscell. p..69 ; Bouleng. Cat. Sn. iil. p. 492.

Synopsis of the Species.

One or two series of scales between the nasal and the rostral;

MOSS GhirseECl WHONTETRES oc. c06 cosone aap con con cor nee one coo ponoHAaAASe Coe B. arietans-.

4 or 5 series of scales between the nasal and the rostral; a single

enlarged, sometimes horn-like scale above the internasal, in

COMGDGH CHUAN MES TAMOKT ccoosacoscossopned cnn ons oegabesneaas ood suaoDsoD9099 B. yabonica.

4 or 5 series of scales between the nasal and the rostral; 2 or 3

enlarged, horn-like scales above the internasal, usually with

small scales between them and their fellows ........................ B. nasiceriis.

1. Brris ARIETANS.

Vipera arietans Merr. Tent. p. 152.

Bitis arvetans Bouleng. t.c. p. 493.

Tropical and South Africa; Southern Arabia.

2. BITIS GABONICGA.

Echidna gabonice Dum. & Bibr. Erp. Gén. vii. p. 1428,

dla keen OS

Bitis gabonica Bouleng. t.c. p. 499.

Tropical Afriea.

3. BITIS NASICORNIS.

Coluber nasicornis Shaw, Nat. Miscell. 111. pl. xeiv.

Bitis nasicornis Bouleng. t.c. p. 500.

Tropieal Africa.

SNAKES OF BAST AFRICA. 639

4, ATHERIS.

Cope, Proc. Ac. Philad. 1862, p. 337; Bouleng. Cat. Sn. iii.

p- 508.

Synopsis of the Species.

No horn-like superciliary scales; scales in 15 to 25 rows; gular

Scalesistronclynkeeled ee teers oaecerce ase sects sae seas seule sles

No horn-like superciliary scales; scales in 25 to 32 rows ; gular

scales smooth or very feebly keeled ...................000000:00000.. A. nitschei.

Several erect, horn-like superciliary scales; scalesin 25 rows ... A. ceratophorus.

A. syuamiger.

1, ATHERIS SQUAMIGER.

Echis squamigera Hallow. Proc. Ac. Philad. 1854, p. 193.

Atheris squamiger Bouleng. t.c. p. 509.

Uganda, Central Africa, West Africa from Calabar to Angola.

2, ATHERIS NITSCHEI Tornier, Zool. Jahrb., Syst. xv. 1902,

p. 589, fig.

Atheris woosnami Bouleng. Ann. & Mag. N. H. (7) xviii. 1906,

p. 37, and Tr. Zool. Soc. xix. 1909, p. 246, pl. ix.

N.W. and E. of Lake Tanganyika, and Mt. Ruwenzori.

3. ATHERIS CERATOPHORUS Werner, Verh. Zool.-bot. Ges.

Wien, xlv. 1895, p. 194, pl. v. fig. 1; Bouleng. Cat. Sn. i.

p- 510.

German East Africa and Togoland.

5, ATRACTASPIS.

A. Smith, Ill. Zool. 8. Afr., Rept.; Bouleng. Cat. Sn. iii.

p. 510.

Synopsis of the Species.

I. Anal divided ; all or most of the subcaudals paired ; scales in 23 to 27 rows.

Snout rounded ; one postocular; third lower labial largest ...... A, irregularis.

Snout pointed; one postocular; third lower labial largest ...... A. conradsi.

Snout rounded; two postoculars ; fourth or fifth lower labial

I EAWe22S Fee ae cemmmRneeans coe melee A. bipostocularis.

II. Anal entire ; all or most of the subcaudals single.

A. Postocular in contact with a large temporal.

Snout cuneiform ; first lower labial in contact with its fellow

behind the symphysial; scales in 23 rows ; ventrals 227—

DAS Mire sececO ees te mat esc creer Hanae oimeie mesonsts od wee auabeltea = Ales MOSENUEGS

Snout rounded; first lower labial in contact with its fellow

behind the symphysial; scales in 19 or 21 rows; ventrals

DAS IIEE OO) Bacmne toniecoe oda evio Seon donoton AopdoneeenOmrACco SORT EGC HERO ECR cone CAAT a UNTG Ts

Snout rounded; symphysial in contact with the chin-shields;

scales in 25 rows; ventrals 242 0.000. oer. A. Katange.

B. Temporals small, 2 or 3 superposed in front; snout sub-

cuneiform ; scales in 29 to 37 rows; ventrals 212-245. A. microlepidota.

44%

640 ON THE SNAKES OF EAST AFRICA,

1. ATRACTASPIS IRREGULARIS.

Elaps irregularis Reinh. Vid. Selsk. Skrift. x. 1843, p..264,

pl. 11. figs. 1-3.

Atractaspis irregularis Bouleng. t.c. p. 513.

From the Gold Coast and Uganda to the Congo.

9, ATRACTASPIS CONRADSI Sternf. Sitzb. Ges. Nat. Fr. Berl.

1908, p. 94.

Uganda and German Hast Africa.

3. ATRACTASPIS BIPosTocuLARIS Bouleng. Ann. & Mag. N. H.

(0) 2a USO) yon LO

Mount Kenya.

4. ArRAcTAsPIs RosTRATA Ginth. Ann. & Mag. N. H. (4) 1.

1868, p. 429, pl. xix. fig. 1.; Bouleng. Cat. Sn, 11. p. 514.

East Africa and Nyassaland.

5, ATRACTASPIS ATERRIMA Giinth. Ann. & Mag. N. H. (3) xii.

1863, p. 363; Bouleng. t.c. p. 515.

Uganda and West Africa, from the Gold Coast to the Niger.

6. ATRACTASPIS KATANG® Bouleng. Ann. Mus. Congo, Zool. ii.

1901, p. 13, pl. v. fig. 2; Sternf. Mitth. Zool. Mus. Berl. v. 1911,

p. 385.

Katanga and German East Africa.

7. ATRACYASPIS MICROLEPIDOrA Giinth. Ann. & Mag. N. H.

(3) xviii. 1866, p. 29, pl. vil.; Bouleng. Cat. Sn. 1. p. 517.

Kast and Central Africa.

ON THE SNAKES OF NORTH-EAST AFRICA. 641

46. A List of the Snakes of North-Hast Africa, from the

Tropic to the Soudan and Somaliland, including

Socotra. By G. A. BouLenenr, F.R.S., F.Z.8.*

[Received October 4, 1915 ; Read November 23, 1915. |

INDEX.

(FEOGRAPHICAL : Page

INL, Aige@a, Wists ot Simvalkes cosscoenoscoubansocgdaceueosbngaagadaréoonn asbil

SYSTEMATIC : 4 ;

Keys to the identification of the Snakes of N.H. Africa ......... 641

With this list the Ophidian fauna of Hast and Central Africa

is completed, only two divisions remaining to be dealt with,

viz., West Africa north of the Congo, and North Africa.

Synopsis of the Families.

I, Worm-like, with small inferior mouth ; eyes hidden or visible under the head-

shields, and body covered with uniform imbricate scales above and beneath.

22 or more scales round middle of body ; ocular not bordering the

mouth; tail not or but little longer than broad .................. TYPHLOPID®.

Te emlas, mowwanel wavKWEllke Ove LOCK oo ececeacoocrocnacuscoceccocacsaoaessancooa, (Can CONGUUDZE,

II. Mouth large, eyes exposed; body with enlarged shields beneath (except in

the marine genus Hydrus).

Ventral shields much narrower than the body; supraocular, if

distinct, broken up into two or more sbields: scales on

anterior part of body smooth ...... . Borpm.

Ventral shields at least nearly as broad as the pody ; ; “supraocular

single; poison-fangs, if present, not ma very large sheath .... CotuBRIp®.

Ventral shields at least nearly as broad as the body, or upper sur-

face of head vovered with small scales; poison-fangs in a very

kaa) SETI TSMOWS THO GO nooscosno si cccecqoess edocaensenoaeeasasacacco — \/ NDITURILEEL

Family TyPHLOPID &.

A single genus.

1. TypHuops.

Schneid. Hist. Amph. i. p. 339; Bouleng. Cat. Sn. 1. p. 7.

Synopsis of the Species.

I. Snout rounded or with obtusely angular horizontal edge.

A. Rostral not more than one-third width of head; eyes distinct.

Snout rounded ; 24 scales round middle of body, the diameter of

which is 37 to 50 times in total length.. ee T. socotranus.

Snout much depressed; 22 seales round middle of body,” ‘the

diameter of which is 20 to 30 times in total length ............ LZ. euneirostris.

B. Rostral very broad.

Rostral half width of head; 30 scales round middle of body, the

diameter of: which is 30 to 40 times in total length ......... T. blanfordii.

Rostral more than half width of head; 26 to 30. scales round

middle of body, the diameter of which is 24 to 30 times in

{nO nee) LICH KH Ie coapuanes cantonsrhonalcdede eubonaccandtnceeudtuonpcee oamucrson:, | lla OOM Le RID

* Published by permission of the Trustees of the British Museum.

642 MR. G. A. BOULENGER ON THE

II. Snout with sharp horizontal edge.

No subocular; 28 scales round middle of body, the diameter of

VuVlOWCG] OL HIS) fs) wees) rool woyil ESOS Cos goa qenonboce boscoa eo S00 soo cooENs T. acutirostris

No subocular; 24 scales round middle of body, the diameter of

which is 90 times in total length ....... . T. somalicus

A large subocular, separating the small ocular from the ‘upper

labials ; 24 scales round middle of body, the diameter of

which is 50 to 68 times in total length ............. wae. LD. unitent

1. TypHiops socorranus Bouleng. Ann. & Mag. N. H. (6) iv.

1889, p. 362, and Cat. Sn. i. p. 21, pl. il. fig. 2.

Socotra.

2. TypHiops cunnIRostRis Peters, Mon. Berl. Ac. 1879, p. 775,

pl. —, fig. 4; Bouleng. Cat. Sn. i. p. 32, and Ann. Mus. Genova

(2) xvii. 1896, p. 10.

Somaliland.

3, TypHLops BLANFORDIT Bouleng. Ann. & Mag. N. H. (6) iv.

1889, p. 363, and Cat, Sn. 1. p. 39; Peracca, Boll. Mus. Torin. xii.

no. 273, p- i

Abyssinia, EKrythrea.

4, TypHiors punctatus Leach.

Bouleng. Cat. Sn. 1. p. 42.

1 yphbops adolphi Sternf. Mitth. Zool. Mus. Berl. v. 1910, p. 70.

Tropical Africa, as far north as the Gambia and the [Gabe el

Gebel; not recorded from Somaliland.

5. Typunops acurirosrris Mocquard, Bull. Mus. Paris, 1905,

Ts C0

Shoa.

6. TypHtops somaticus Bouleng. Proc. Zool. Soc. 1895, p. 536,

pl. xxx. fig. 1, and Cat. Sn. 11. p. 589.

Somaliland.

7. TYPHLoPs UNIT#NIATUS Peters, Mon. Berl. Ac. 1878, p. 205,

pl. ii. fig. 5; Bouleng. Cat. Sn. i. p. 55, and Ann. Mus. Genova

(3) v. 1912, p. 331 (var. ateniatus).

Somaliland and British Hast Africa.

Family GLAUCONIIDA.

A single genus.

1. GLAUCONIA.

Gray, Cat. Liz. p. 139; Bouleng. Cat. Sn. i. p. 59.

Synopsis of the Species.

I. Ocular bordering the mouth.

A. Snout more or less strongly hooked in profile.

Rostral extending posteriorly to the level of the eyes; diameter

of body 70 to 113 times in total length ..............:000e = G. macrorhynchus.

SNAKES OF NORTH-EAST AFRICA. 643

Rostral not extending to the level of the eyes; diameter of

body 100 to 140 times in total length ................. ce G. filiformis.

Rostral not extending to the level of the eyes; diameter of

body 40 to 48 times in total length ...........0.....ccc eee =GL macrura.

B. Snout not hooked.

Supraocular nearly twice as broad as long; diameter of body

50 to 55 times in total length . G. emini.

Supraocular small, longer than broad ; “diameter of ‘body 38

times in total length FRR ee ect ee ee a LE aa high iereimeticies G. reticulata.

Supraocular small, not or but little broader than long; diameter

of body 65 to 90 times in total length ....................8.. G. cairi.

II. Ocular not bordering the mouth; diameter of body 42

HUINOS Tn HOA EMEA scoco0 cou vooasd sav osvAnoANeoageen nse snestoconons |» Gro GuSSoMDAlds:

1. GLAUCONIA MACRORHYNCHUS.

Stenostoma macrorhynchum Jan, Arch. Zool. Anat. Phys. 1.

1862, p. 190.

Glauconia macrorhynchus Bouleng. Cat. Sn. 1. p. 61.

Glauconia algeriensis Jacquet, Bibl. Anat. iv. 1896, p. 79, figs.

ao Algeria, Mesopotamia (?), Persia.

. GLAUCONIA FILIFORMIS Bouleng. Bull. Liverp. Mus. i. 1899,

De 7, and Rep. Sokotra Exped. p. 88, JON, tly Hees He

Socotra.

3. GLAUCONIA MACRURA Bouleng. Rep. Sokotra Exped. p. 89,

pl. xi. fig. 3.

Socotra.

4. GLAUCONIA EMINI Bouleng. Ann. & Mag. N. H. (6) vi. 1890,

p. 91, and Cat. Sn. i. p. 64, pl. i. fig. 8.

Conte and Hast Africa, 2 as far north as Somaliland and

Abyssinia.

5. GLAUCONIA RETICULATA Bouleng. Ann. & Mag. N. H. (7) xviii.

1906, p. 441.

Goolis Mts., Somaliland.

6. GLAUCONIA CAIRI.

Stenostoma cari Dum. & Bibr. Erp. Gen. vi. p. 323.

Glauconia cairt Bouleng. Cat. Sn. i. p. 66.

Egypt, Nubia, Abyssinia, Somaliland, Mauritania.

7. GLAUCONIA DISSIMILIS.

Stenostoma dissimile Bocage, Jorn. Sc. Lisb. xi. 1886, p. 174.

Glaucenia dissimilis Bouleng. t.c. p. 70.

White Nile.

Family Borps.

Two genera : —

Upper surface of head with shields; rostral and anterior upper labials

deeply pitted; subcandals im two rOWS .......00....ceecceeeeeeeeesseesseeseee Python.

Upper surface of head with small scales; tail very short, subcaudals

SIONS (ey, aeree ieee ice neacircis NEC MBER eu om Gms GCE a seer etiotsen Spat TOR ORCC ERC Ge ee He

644 MR. G. A. BOULENGER ON THE

l. PyrHon.

Daud. Hist. Rept. v. p. 266; Bouleng. Cat. Sn. i. p. 85.

‘Two species :—

Two upper labials pitted ; 79-95 scales across thickest part of body;

ventrals 260-286; subcaudals 58-82 ... . P.sebe.

Four upper labials pitted ; 53-63 scales across thickest t part o of f body’

ventrals 196-207 ; subcaudals BOLSIT Serecce oe eens iactine EPXreguus:

PYTHON SEB.

Coluber sebe Gmel. 8. N.i. p. 1118.

Python sebe Bouleng. t.c. p. 86.

Tropical and South Africa, as far north as Senegambia and

the White Nile.

. Pyrox REGIUS.

oe regia Shaw, Zool. ui. p. 347, pl. xevi.

Python regius Bouleng. t.c. p. 88.

Senegainbia to Niger, eastwards to the Hgyptian Soudan,

2. Kryx.

»)

Daud. Hist. Rept. vii. p. 251; Bouleng. Cat. Sn. i. p. 122.

Two species :—

Tail ending in a conical scute; scales in 47 to 53 rows, keeled on

posterior TENA OF LOCK, coe cco once ..... DH. thebaicus.

Tail ending in a curved, claw-like scute ; ‘scales in 41 to ‘45 row vs, all

smooth.. SE CHSEE L5G ds BaRuOR GanlcnM Mansa ccySacadda 1a Maced eomO Ua ABObASSEe nc ial COA Os

|. Kryx rHEparcus Reuss, Mus. Senckenb. i. 1834, p. 134;

Bouleng. t.c. p. 125

Upper Egypt to German East Africa.

HRYX MUELLERI.

Gongylophis muelleri Bouleng. Ann, & Mag. N. H. (6) ix. 1892,

p- 74.

Hryx muellert Bouleng. Cat. Sn. i. p. 128, pl. v. fig. 2.

Egyptian Soudan, Mauritania, N. Nigeria, Togoland.

Family CoutuBRID g.

Three parallel series :—

No poison-fangs; all the teeth solid .......0.........0c.... A. Aglypha.

JANOS [SING 5. aosoncacaoavdcaoasaenonsencoopcceneaccosns Joh Opisthoglypha.

JERDISOMHAYSS TN IRON G so sacndo ccarooanaccnennesecosomevooscnsacoan Ch Proteroglypha.

A. Aglypha.

J. Nostril between two nasals, or between two nasals and the internasal; loreal

present.

A. At least one upper labial bordering the eye.

1. Pupil round; scales in 19 or 21 rows; no subocular; a single anterior

temporal.

Seales in 19 rows; ventrals 131-150; anal divided ; subcaudals

DHCD eaasttna Senades Sake CERRO Ute ene PR Ml UC a an Tropidonotus.

SNAKES OF NORTH-EAST AFRICA,

Scales in 21 rows; ventrals 174-198 ; anal entire; subcaudals 36-

41; a deep groove above 5th and 6th upper labials

2. Pupil vertically elliptic.

a. Anal entire; a single anterior temporal.

SGHIES SUN OUI. in PUSSIES} IROWIS 2. coves > socecdescnadsos S60 ons a5vosb ean DUNSoDNES

Scales smooth, in 17 rows SEG HneC OR aOno ARR soneed eS

Scales strongly keeled, vertebrals bicarinate

b. Anal divided.

Scales in 21 rows; ventrals 216 ; subcaudals 67-70 ..................

Scales in 19 rows; ventrals 160-188; subcaudals 35-46; rostral

angularly bent, detached at the sides

3. Pupil round ;

scales in 15-23 rows;

640

Pseudoboodon.

Boodon.

Lycophidium.

Simocephalus.

Aeluroglena.

Lytorhynchus.

ventral 150 « or more, or, if less,

subcaudals 100 or more; loreal not more than twice as long as deep.

a. One or two suboculars below the ee ae ; frontal

not twice as long as broad

6. No subocular ; frontal not twice as long as ‘oad

Scales in 15 rows; Fonte 150-193 ; subcaudals 77-123, without

Eby) g=12) linea caesar Mum Toh

Scales in 15 rows; ventrals 166-207 ;

strong keel .

Scales in 21 rows; -ventrals 176-

“subcaudals 112— -157, with a

224; subcaudals 63-88 ............

ce. No subocular; frontal at least twice as long as broad ;

scales in 15 rows; ventrals nies subcaudals

100-128 . sees Sastre

3. Eye separated Ay the upper _jeiviels es series of sub-

oculars ; snout with angular horizontal edges; scales in

19-25 rows

JI. Nostril in a single or sonatill Ped a

B. Scales smooth, in 15 rows.

1. Nasal semidivided, with a longitudinal cleft behind the nostril ;

and prefrontal single.

Rostral with angular horizontal edge

Rostral with obtuse horizontal edge

2. Nasal entire; internasal el -peaitroathal Pore

WVentirals 162) anal divided’: subcandals)/69s.s-0- se eesees ese snee oe

Ventrals 113-144; anal entire; subcaudals 19-46 ...........0.000..0..

B. Scales strongly keeled, in 23-27 rows ; no loreal ............

1. TROPIDONOTUS.

Kuhl, Bull. Sc. Nat. ii, 1824, p. 81;

p. 192.

1. TROPIDONOTUS OLIVACEUS.

Bouleng.

Zamenis.

Chlorophis.

Philothamnus.

Coronella.

Grayia.

Scaphiophis.

internasal

Prosymna.

Asthenophis.

Contia.

Homalosoma.

Dasypeltis.

Oris, SiGs ay

Coronella olivacea Peters, Mon. Berl. Ac. 1854, p. 622.

Tropidonotus olivaceus Bouleng. t. ¢. p. 227.

Tropical Africa, from the Soudan to Angola, Damaraland, and

Southern Rhodesia.

2, PsEUDOBOODON.

Boll. Mus.

Peracca, Torin.

PSEUDOBOODON GASCA.

Peracea, t.c. p. 2, fig.

Lamprophis abyssinicus Mocquard,

p. 249.

Erythrea,

Bull. Mus.

Abyssinia.

siti, SIS) f5 TOE, AM oy Ala

Paris,

1906,

646 MR. G. A. BOULENGER ON THE

3. Boopon.

Dum. & Bibr. Mem. Ac. Sc. xxii. 1853, p. 460; Bouleng.

Cat. Sn.1. p. 327.

Synopsis of the Species.

I. Scales in 21 or 23 rows; preocular not extending to upper surface of head.

Three upper labials entering the eye; a single anterior temporal . B. lemniscatus.

Two upper labials entering the eye; 2 or 3 superposed anterior

Cemap onallsmaeeenenamasatensat: socines «rere seen adcleaciuea teins B. erlangeri.

II. Seales in 25 to 33 rows ; preocular extending to upper surface of head.

Parietals longer than the distance between the frontal and the end

Orsi MeNSTV OW beMmmep er recor a ticc ene ene ce erate epee a B. lineatus.

Parietals as long as the distance between the frontal and the end

OfgbeFemOUt pH eta ccse secece od sues ise nolo ceca Mee eee SEER eres ED a ULEGUILOSUS =

1. Boopon Lemniscatus Dum. & Bibr. Erp. Gén. vii. p. 365 ;

Bouleng. t.c. p. 329.

Lamprophis rogert Mocquard, Bull. Mus. Paris, 1904, p. 307,

and 1905, p. 289.

Abyssinia.

2. BooDON ERLANGERI Sternf. Sitzb. Ges. Nat. Fr. Berl. 1908,

p. 92.

Somaliland.

3. Boopon LinEATUS Dum. & Bibr. Erp. Gén. vi. p. 363;

Bouleng. t. ¢. p. 332.

Tropical and South Africa; 8. Arabia.

4, BooDOoN FULIGINOSUS.

Lycodon fuliginosus Boie, Isis, 1827, p. 551.

Boodon fuliginosus Bouleng. t.c. p. 334.

Mauritania to N. Nigeria, eastwards to the Egyptian Soudan.

4, JLyYcoPHIDIUM.

Dum. & Bibr. Mém. Ac. Sc. xxiii. 1853, p. 462; Bouleng.

Cat. Sn. i. p. 336.

Two species :—

Rostral nearly twice as broad as deep...............s0ecceeeeeeeeeee De. capense.

Rostral but little broader than deep ..............:0:ccceeeeeeee DL. abyssinicum.

1. LycoPHIDIUM CAPENSE.

Lycodon capensis A. Smith, S. Afr. Quart. Journ. i. no. 5,

Soh, Toy Ws.

Lycophidium capense Bouleng. t.c. p. 339.

Tropical and South Africa.

2. LiycoPHIDIUM AByssINIcUM Bouleng. t.c. p. 342, pl. xxii.

mee, Me

Abyssinia.

SNAKES OF NORTH-EAST AFRICA. 647

5, SIMOCEPHALUS.

Giinth. Cat. Col. Sn. p. 194; Bouleng. Cat. Sn. in. p. 344,

1. SmvocePHALus BUTLERI Bouleng. Ann. & Mag. N. H. (7) xx.

1907, p. 489, and Ann. Mus. Genova (3) v. 1911, p. 164.

Mehelya somaliensis Linnb. & Anderss. Ark. f. Zool. viii. 1913,

no. 2, p. 2.

Egyptian Soudan, Southern Ethiopia, Somaliland.

6. ZAMENIS. ;

Wagl. Syst. Amph. p. 188; Bouleng. Cat, Sn. i. p. 379.

Synopsis of the Species.

io SGMES 1 1H TOWIES WEMURAIS T7/S sco nceceg coaconoso concenead ene aoNecc Z. somalicus.

IL. Scales mn 19 rows.

Ventrals 213-262; subcaudals 113-154 .............2..cceeeeeeeee. Z. rhodorhachis.

Womtinalle 19s BmIt eR) [email protected] Co UREOIS:

ILI. Scales in 21 (rarely 23) rows; two labials entering the eye.

9 or 10 upper labials; ventrals 171-187 ...... «02.0... 2.2.00 ens eee Z. smithii.

9 upper labials; ventrals 200-228 ..........:0ceceeeeeeeee eee GZ, florulentus.

SB} mypjeaie MNS S wetaeills WES) so sconce adaccaconpaneee ap toncuoepnosccoaen 4a CUUGIAOd

IV. Scales in 23 rows; a single labial entering the eye;

WO MTTIIS DNO-OS) 0 docinsactsous onersaconosebodcodeasanbenuegeedesqoo An SOAOURE

1. ZAMENIS soMALICUS Bouleng. Ann. Mus. Genova (2) xvii.

T8965 p. LL

Somaliland.

2. ZAMENIS RHODORHACHIS Jan, in De Filippi, Viagg. Pers.

p- 356; Bouleng. Cat. Sn. 1. p. 398.

Egypt to Somaliland ; 8.W. Asia.

3. ZAMENIS BREVIS Bouleng. Ann. Mus. Genova (2) xv. 1895,

p- 13, pl. iii. fig. 3, and Cat. Sn. i. p. 623.

Somaliland.

4. ZAMENIS sMITHII Bouleng. P. Z.8. 1895, p. 536, pl. xxx.

fig. 2, and Cat. Sn. iii. p. 624.

Somaliland.

5. ZAMENIS FLORULENTUS.

Coluber florulentus Geoftr. Descr. Egypte, Rept. p. 146,

pl. vii. fig. 2.

Zamenis florulentus Bouleng. Cat. Sn, 1. p. 402.

Egypt to Somaliland.

6. ZAMENIS CITERNIT Bouleng. Ann. Mus. Genova (3) v. 1912,

p- 331.

Somaliland.

648 MR. G. A. BOULENGER ON THE

7. ZAMENIS socorR# Giinth. P. Z. 8. 1881, p. 463, pl. xli.;

Bouleng. Cat. Sn. i. p. 408.

Socotra.

7. AELUROGLENA.

Bouleng. Ann. & Mag. N. H. (7) ii. 1898, p. 132.

1. AELUROGLENA cucuLLATA Bouleng. 1. ec.

Somaliland.

8. LyroRHYNCHUS.

Peters; Mon. Berl. Ac. 1862, p. 272; Bouleng. Cat. Sn. i.

p- 414.

1. LyvoRHYNCHUS DIADEMA.

Heterodon diadema Dum. & Bibr. Erp. Gén. vii. p. 779.

Lytorhynchus diadema Bouleng. t. c. p. 415.

Algerian and Tunisian Sahara, Tripoli, Egypt, Nubia; Arabia,

Syria.

9. CHLOROPHIS.

Hallow. Proc. Ac. Philad. 1857, p. 52; Bouleng. Cat. Sn. il.

pa Je

Two species :—

3 labials entering the eye; ventrals 151-193, without lateral keels ... C. emini.

2 labials entering the eye; ventrals 148-169, with a slight lateral

J RIS Le aero nt cae tari orieasaeaaencices eo sanee ace bo suan ode cbneudtnsuaeebutoMareomadsoon, | Cn ROAMUACEOIN

1. CHLOROPHIS EMINI.

Ahetulla emint Giinth. Aun. & Mag. N. H. (3) xi. 1863,

p. 289.

Chlorophis emini Bouleng. t. c. p. 92, pl. v. fig. 1.

Egyptian Soudan to Uganda and Ruwenzori.

2. CHLOROPHIS NEGLECTUS.

Philothamnus neglectus Peters, Mon. Berl. Ac. 1866, p. 890.

Chlorophis neglectus Bouleng. t. c. p. 94.

Central and Kast Africa, as far north as Somaliland.

10, PHILOTHAMNUS.

A. Smith, Ill. Zool. S. Afr., Rept.; Bouleng. Cat. Sn. 11. p. 98.

1. PHILOTHAMNUS SEMIVARIEGATUS A. Smith, op. cit. pls. lix.,

lacs, Leia gions Boulemesmtecs (ook

Tropical and South Africa.

SNAKES OF NORTH-EAST AFRICA. 649

11. Coronrua.

Laur. Syn. Rept. p. 84; Bouleng. Cat. Sn. ii. p. 188.

1. CoRONELLA SEMIORNATA Peters, Mon. Berl. Ac. 1856, p. 622 ;

Bouleng. t. c. p. 195.

N. Rhodesia and EK. Africa, as far north as the Blue Nile.

12. Grayia.

Giinth. Cat. Col. Sn. p. 50; Bouleng. Cat, Sn. it: p. 286.

1, GrayiA THOLLONT Mocquard, Bull. Soc. Philom. (8) ix. 1897,

jp LI Ieomllenies IP. 41S, O08), jo, SDI, tee

Egyptian Soudan, Uganda, Katanga, French Congo,

13, SCAPHIOPHIS,

Peters, Mon. Berl. Ac. 1870, p. 644; Bouleng. Cat. Sn. ii.

p. 254.

1. SCAPHIOPHIS ALBOPUNCTATUS Peters, t. c. p. 645, pl. i. fig. 4;

Bouleng. 1. ¢.

Tropical Africa, from the Soudan to the Congo.

14. ProsymMna.

Gray, Cat. Sn. p. 80; Bouleng. Cat. Sn. ii. p. 246.

1. PrRosyMNA MELEAGRIS.

Calamaria meleagris Reinh. Vid. Selsk. Afh. x, 1843, p. 238

pl. i. figs. 4-6.

Prosymna meleagris Bouleng. t. ¢. p. 249,

Egyptian Soudan and Sierra Leone to the Congo.

15, Astamnopnts.

Bouleng. Ann. Mus. Genova (2) xvii. 1896, p. 12.

1. ASTHENOPHIS RUSPOLIT Bouleng. ]. ¢.

Somaliland,

16. Contra.

Baird & Gir. Cat. N. Am. Rept. p. 110; Bouleng. Cat. Sn. ii.

p. 255.

1, Conria arricana Bouleng. Ann. & Mag. N. H. (8) xiv. 1914,

p. 485.

red Sea Province of the Soudan.

650 MR. G. A. BOULENGER ON THE

17. Homanosoma,

Wagl. Syst. Amph. p. 190; Bouleng. Cat. Sn. i. p. 273.

1. HoMALOSOMA LUTRIX.

Coluber lutria Linn. 8. N. i. p. 375.

Fomalosoma lutrix Bouleng. ‘ ce. p. 274.

Homalosoma shiranum et abyssinicum Bouleng. t. c p. 276,

pl. xi. figs. 1 & 2

Kast, Central, and South Africa, as far north as Abyssinia.

18. DAsyPELTIs.

Wagl. Syst. Amph. p. 178; Bouleng. Cat. Sn. i. p. 353.

1. DASYPELTIS SCABRA.

Coluber scaber Linn. Mus. Ad. Frid. p. 36, pl. x. fig. 1.

Dasypeltis scabra Bouleng. t. c. p. 354.

Tropical and South Africa, Egypt; South Arabia.

B. Opisthoglypha.

I. Eye rather small, moderate, or larze, with vertically elliptic or subelliptic pupil;

loreal present; nasal, if not completely divided into two, with horizontal cleft.

A. Nostril between two nasals.

Scales in 21 or 23 rows; ventrals rae subcaudals single,

35-44 0... peudadtven aagrmeadsepnennuonsse, 2 LUA DOODES:

Scales in 19 to 23 LOWS; ; ventrals 2 20 5-974; subcaudals 61-86

pairs. sae dates aoe er eme eS lsc ence diets eee LU DODILESS

Scales in 19 TOWS; . ventrals 144-180; subcaudals 32-54 pairs... Leptodira.

B. Nasal semidivided ; scales in 17 (rarely 19) rows; ventrals

148-173; 187: subcaudals 61-78 ........................... Hemirhagerrhis.

II. Eye moderate or large, with round or horizontal pupil; loreal present.

A. Pupil round.

1. Loreal not more than once and a half as long as deep; scales not oblique.

a. Nostril round ; scales in 17 rows.

Nostril in a semidivided nasal; rostral broader than ee just

visible from above... .. Amplorhinus.

Nostril hetween two nasals ‘and. the. ipter aacall: ; ‘rostral’ as : deep

as broad, its upper potion at least half as long as its

distance from the frontal.. Beles a: pspavcasnooeoesce LU PRPOROPIDOISS

b. Nostrila GremeaHte. Sits in a aie idea or Siconiidinidad nasal; scales in 17

or 19 rows.

SiuleamGlalls Ness nara 0) ccoccaanccce sauonsenaeanancednepaboancesnnccosze0s — ClaAlOV RANTS.

aes S0lorsmorepeeeeeee veseeeseeee Rhamphiophis.

. Loreal at lense « once ane a ial: as eilenen as sdéeer scales more or less oblique;

subcaudals 78 or more.

Scales in 17 rows; a single anterior temporal ......... Dromophis.

Scales in 18 to 19 rows ; usually two super posed ‘anterior ‘tem-

porals; one or two anterior ees teeth much enlarged,

fang-like . pasaglsseets ..... Psammophis.

3. Tigveal not more ein ae once andl a half a as Metiea as eases

nostril in an undivided nasal; scales very narrow,

very oblique, keeled, in 19 or 21 rows ......... . Dispholidus.

B. Pupil horizontal ; nostril in an undivided nasal ; asin

narrow, oblique, feebly keeled, in 19 rows ............... Thelotornis.

SNAKES OF NORTH-EAST AFRICA. 651

TII. Eye small or very small; no loreal; nasal, if present, single; scales in

y y ? ) p )

15 rows.

A. Subcaudals paired ; ventrals 171-256

Nasal and internasals present... ere Unenelaniss

No nasal or internasals ; nostril in the first upper Tabialo. ee Chilorhinophis.

B. Subcaudals single.

Internasal in contact with first upper labial ; ventrals 104-115 ;

Subcaudal sSNA a wemene eee ce eencec te asics g.scar dele cclaisiae sulsiot aller Brachyophis.

Nasal in contact with rostral; ventrals 145-165; subcaudals

AORTA Pea. aos CAR BREE Re Lae A pan allaclUs,

1. Dirypopuis.

Ginth. P. Z.S. 1881, p. 462; Bouleng. Cat. Sn. i. p. 46.

1. Drryroruis vivax Giinth. 1. c. pl. xl.; Bouleng. |. e.

Socotra.

2. 'TARBOPHIS.

Fleischm. Dalm. nov. Serp. Gen. p. 17; Bouleng. Cat. Sn. iii.

p- 47.

Two species :—

Scaleshimbli9iors2 lerows))anallentinesnn.eeeseeensseeeees secre sete ese eee NOLEN LE 120.

Scalesnml2von 23mowsis) anal ainvidedmese use neeeesceseeteeee ee eceeeeeehee nn eELEOULUSIHS.

1. TARBOPHIS GUENTHERI Anders. P. Z. 8. 1895, p. 656,

pl. xxxvi. fig. 3; Bouleng. t. c. p. 52.

Hast Africa, as far north as Somaliland; Arabia, Syria.

2. TARBOPHIS OBTUSUS.

Coluber obtusus Reuss, Mus. Senckenb. i. 1834, p. 137.

Tarbophis obtusus Bouleng. |. ¢.

Egypt to Somaliland, Mauritania and N. Nigeria.

3. LEPTODIRA.

Giinth. Cat. Col. Sn. p. 165; Bouleng. Cat. Sn. in. p. 88.

1. LEPTroDIRA HOTAMBGIA.

Coronella hotambeia Laur. Syn. Rept. p. 85.

Leptodira hotambeia Bouleng. t. c. p. 89.

Tropical and South Africa.

4. HEMIRHAGERRHIS.

Boettg. Zool. Anz. 1893, p. 119; Bouleng. Cat. Sn. iii. p. 119.

1. HEMIRHAGERRHIS KELLERI Boettg. t.c. p. 129; Bouleng. l.c.

Somaliland and British Hast Africa.

652 MR. G. A. BOULENGER ON THE

5, AMPLORHINUS.

A. Smith, Ill. Zool, 8. Afr., Rept.; Bouleng. Cat. Sn. i.

"p. 124,

1. AMPLORHINUS NOTOTENIA.

Coronella nototenia Ginth. P. Z. 8. 1864, p. 309, pl. xxvi.

fie. 1.

Amplorhinus nototenia Bouleng. t.c. p. 125.

Amplorhinus giintheri Mocquard, Bull. Mus. Paris, 1906,

[Os 2Ole

Kgyptian Soudan to Nyassaland; Angola.

6. TRIMERORHINUS.

A. Smith, Ill. Zool. S. Afr., Rept.; Bouleng. Cat. Sn. 11.

p. 138.

1, TRIMERORHINUS TRITENIATUS.

Rhagerrhis tritenata Giinth. Ann, & Mag. N. H. (4) 1. 1868,

p. 423, pl. xix. fig. H.

Trimerorhinus triteniatus Bouleng. t.¢. p. 139.

Africa south of the Equator; Hast Africa as far north as

Abyssinia.

Perhaps not specifically separable from 7’. rhombeatus L.

Ca:LOPELTIS.

Wagl. Syst. Amph. p. 189; Bouleng. Cat. Sn. ii. p. 141.

1. C@LOPELTIS MOILENSIS.

ee moilensis Reuss, Mus. Senckenb. 1. p. 142, pl. vii. fig. 1,

Ceelopeltis moilensis Bouleng. t.c¢. p. 1438.

Nocihersn Sahara, from Algeria to Egypt and Nubia ; Apap

western Persia.

RHAMPHIOPHIS.

Peters, Mon. Berl. Ac. 1854, p. 624; Bouleng. Cat. Sn. iii

p. 144.

Two species :—

Seales in 19 rows on the body ; ventrals 230-241; subcaudals

154-160... : sopepoesnescens JAte TRUDI O/OUNOCUTIS.

Scales in 17 rows on n the body ; ventrals. 148-192, ; subcaudals

90-110 . pg acd ace aoplagd ca von odoaaouds oka ragunmmntnnruncacasasace oe fi CROP MNOTAOdISS

]. RHAMPHIOPHIS RUBROPUNCTATUS.

Dipsina rubropunctata Fisch. Jahrb. Hamb. Wiss. Anst. 1.

1884, p. 7, pl. i. fig. 3.

Rhamphiophis rabr opunctatus Bouleng. t. c. p. 146.

Egyptian Soudan to Kilimanjaro,

SNAKES OF NORTH-EAST AFRICA, ‘ 653

2. RHAMPHIOPHIS OXYRHYNCHUS.

Psammophis oxyrhynchus Reinh. Vid. Selsk. Skrift. x. 1843,

p- 244, pl. i. figs. 10-12.

Phamphiophis oxyrhynchus Bouleng. |. c.

Tropical Africa.

% 9. DRoMOPHIS.

Peters, Mon. Berl. Ac. 1869, p. 447; Bouleng. Cat. Sn. u1.

p. 149.

1. DROMOPHIS LINEATUS.

Pryophylax lineatus Dum. & Bibr. Erp. Gén. vii. p. 1124.

Dromophis lineatus Bouleng. |. ¢.

Coast of Guinea to Eastern Soudan, Central Africa.

10. PSAMMOPHIS.

Boie, Isis, 1827, p. 521; Bouleng. Cat. Sn. ii. p. 152.

Synopsis of the Species.

I. Scales in 17 rows, rarely 19.

Rostral a little broader than deep; 9 (rarely 8 or 10) upper labials,

sixth shorter than eye; subcaudals 93-154. . P. schokari.

Rostral a little broader than deep; 9 (rarely 8) upper Jabials, ‘sixth

(or fifth) as long as eye; subcaudals 130-178 .. P. punctulatus.

Rostral as deep as broad ; 8 ee 9) upper labials ; ‘subcaudals

85-116 .. Leper tok Maire MEN ae ETE P. sibilans.

II. Scales in 15 rows P. biseriatus.

IIT. Scales in 13 rows P. pulcher.

1. PSAMMOPHIS SCHOKARI.

Coluber schokari Forsk. Descr. Anim. p. 14.

Psammophis schokari Bouleng. t. c. p. 157.

North Africa to Mauritania and Somaliland; Arabia and Syria

to Afghanistan and Sind.

PsaAMMOPHIS PUNCTULATUS Dum. & Bibr. Erp. Gén. vii.

p- 887; Bouleng. t. ¢. p. 159.

East Africa, from the Blue Nile to Mozambique ; Arabia.

3. PSAMMOPHIS SIBILANS.

Coluber sibilans Linn. S. N. 1. p. 383.

Psammophis sibilans Bouleng. t. ec. p. 161.

Tropical and South Africa, Egypt.

_ A, PSAMMOPHIS BISERIATUS Peters, Sitzb. Ges. Nat. Fr. Berl.

1881, p. 88; Bouleng. t. c. p. 168.

Somaliland and British East Africa.

Proc. Zoot. Soc.—1915, No. XLV. 45

654 MR, G. A. BOULENGER ON THE

5, PsAMMOPHIS PULCHER Bouleng. P. Z.S. 1895, p. 537, pl. xxx.

fig. 3, and t. ec. p. 169.

Somaliland.

11. DispHoLIDus.

Duvernoy, Ann. Se. Nat. xxvi. 1832, p. 150; Bouleng. Cat.

pSbaly WL, 7D, ltsko),

1. DisPHoLIDUS TYPUS.

Bucephalus typus A. Smith, Zool. Journ. iv. 1829, p. 441.

Dispholidus typus Bouleng. t. ec. p. 187.

Tropical and South Africa.

12. THELOTORNIS.

A. Smith, Ill. Zool. S. Afr., Rept.; Bouleng. Cat, Sn. iii.

p. 184.

1. THELOTORNIS KIRTLANDII.

Leptophis kirtlandii Hallow. Proc. Ac. Philad. 1844, p. 62.

Thelotornis kirtlandii Bouleng. t. ec. p. 185.

Tropical and South Africa.

13. Micrenars.

Boettg. Ber. Senckenb. Ges. 1879-80, p. 136; Bouleng. Cat.

Sn. ii. p. 248.

Two species :—

3 upper labials entering the eye; ventrals 171........................... MM. vaillanti.

2 upper labials entering the eye; ventrals 203-232........0.0............ MM. boettgeri.

1. MiIcRELAPS VAILLANTI.

Elaposchema vaillanti Mocquard, Mém. Cent. Soe. Philom.

1888, p. 123, pl. xu. fig. 1.

Micrelaps vaillanti Bouleng. t. c. p. 249.

Somaliland,

2. MrcreLars BoErTGERI Bouleng. Ann. Mus. Genova (2) xvii.

1896, p. 13.

Micrelaps nigriceps Sternf. Mitth. Zool. Mus. Berl. v. 1910,

p- 69.

Somaliland, Abyssinia.

14. CHILORHINOPHIS.

Werner, Sitzb. Ak. Wien, cxvi. 1. 1908, p. 1881.

1. CHILORHINOPHIS BUTLERIT Werner, |. c. pl. iii. fig. 8.

Egyptian Soudan.

‘hi

SNAKES OF NORTH-EAST AFRICA. 654

15. BRACHYOPHIS.

Mocquard, Mém. Cent. Soc. Philom. 1888, p. 125; Bouleng.

Cat. Sn. i. p. 254.

1. Bracuyopruis REvoILI Mocquard, |. c. pl. xi. fig. 3; Bouleng.

Somaliland.

16. APARALLACTUS.

A. Smith, Ill. Zooi. S. Afr., Rept., App. p. 15; Bouleng. Cat.

Sn. il. p. 255.

1, APARALLACTUS CONCOLOR.

Uriechis concolor Fisch, Jahrb. Hamb. Wiss. Anst. i. 1884, p. 4,

jOlle ale, aes Al

Aparallactus concolor Bouleng. t. ¢. p. 257.

Lado, Uganda, Somaliland.

C. Proteroglypha.

(Loreal absent in all the genera.)

J. Tail ae oar-shaped ; scales juxtaposed ; no ventrals

(Sea-snake) .. SRC haa be sooopuccseosaca « LELOE PUIG

I Tail ey. diese eeaales mibtcaiee ; ee Taree!

Head short, snout broader than long; scales oblique, 17—25 across

middle of body, 21-29 across neck; anal entire ....... . Naia.

Head short, snout broader than long; scales not oblique, 23 a across

middle ot body, 27 across neck; anal divided .................-... Walterinnesia.

Head long and narrow, snout not broader than long; scales very

ClO G MS, thd NO HO WB WOW coo sooceanoosvcosssnaeosoadcootcoceanogavescoose DR ORRESDIS

: 1. Hyprus.

Schneid. Hist. Amph. 1. p. 233; Bouleng. Cat. Sn. 11. p. 266.

1. HyDRUS PLATURUS. °

Anguis platura Linn. 8. N.1. p. 391.

Hydrus platurus Bouleng. t. c. p. 267.

Indian and Pacific Oceans; E. Africa as far north as the

coast of Obok.

2, Nata.

Laur. Syn. Rept. p. 90; Bouleng. Cat. Sn. iii. p. 372.

Two species :—

Eye separated from the upper labials by suboculars; 6th or 7th

upper labials largest and deepest ........ sacoo ly LAGI

Third or third and fourth upper labials deepest and entering the. eye. NV. nigricollis.

‘1. NAIA HAIE.

Coluber haie Linn. 8. N. 1. p. 387.

Naia haie Bouleng. t. c. p. 374.

Borders of the Sahara, Hast Africa southwards to the Transvaal

and Zululand; Arabia, Palestine.

AD*

656 MR. G. A. BOULENGER ON THE

2. Nara niericoniis Reinh. Vid. Selsk. Skrift. x. 1843, p. 269,

pl. iii. figs. 5-7; Bouleng. t. c. p. 378.

Senegambia and Upper Egypt to Bechuanaland and Natal.

3. WALTERINNESIA.

Lataste, Le Natur. 1887, p. 411; Bouleng. Cat. SMe Wl. Pyaar.

1. WALTERINNESIA 2GyPTIA Lataste, 1. c.; Bouleng. 1. c.

Nubia? Egypt ?

4, DENDRASPIS.

Schleg. Versl. Zool. Gen. Amsterd. 1848; Bouleng. Cat. Sn. ui.

p. 434.

Two species :—

Second upper labial much deeper than first ............... 000 scene eee D. angusticeps.

Second upper labial not deeper than first ...........0.::0eceeee eee D. antinori.

1. DENDRASPIS ANGUSTICEPS.

Naia angusticeps A. Smith, Ill. Zool. S. Afr., Rept. pl. Ixx.

Dendraspis angusticeps Bouleng. t. ¢. p. 437.

East and Central Africa, Angola, South Africa.

. DENDRASPIS ANTINORIT Peters, Mon. Berl. Ac. 1873, p. 411,

pl. 1. fig. 2; Bouleng. 1. c.

Abyssinia.

Family VIPERID.

Synopsis of the Genera.

I. Eve moderate or large; separated from the upper labials by suboculars.

A. Upper surface of head covered with large symmetrical

shields ; loreal present ; pupil round; ventrals 120-155 . Causus.

B. Upper surface of head covered with scales; pupil vertical; scales strongly

keeled ; ventrals 130-205.

Lateral scales not smaller than dorsals, without serrated keels ; sub-

caudals in two rows .... seeenuce JS0GIS.

Lateral scales smaller than “dorsals, ‘oblique, “with serrated. ‘keels ;

subcaudals in two rows ...... .... Cerastes.

Lateral scales smaller than dorsals, ‘oblique, “with serrated ‘keels ;

SIDCROLDIIS SIVA, Coocde osasodcocoenag soocenesacogaad noses jessesccosnasascce | JaADAS

II. Eye minute, with round pupil; upper surface of head covered

with large symmetrical shields ; no loreal; ventrals 212-245. <Atractaspis.

1. Causus, ;

Wagl. Syst. Amph. p. 172; Bouleng. Cat. Sn. iii. p. 465.

Two species :—

Snout obtuse, moderately Yee nents dorsal scales more or less ee

distinctly Keeled .. 43 cu. OL rhombeatus.

Snout prominent, more or » less distinctly. tmmed up at the end; i

scales smooth or faintly keeled 0. ci Gevestsss ssnesuter ess .s. Ce PESiINMSy.

SNAKES OF NORTH-EAST AFRICA. 657

1. CausuUS RHOMBEATUS.

Sepedon rhombeatus Licht. Verz. Doubl. Mus. Berl. p. 106.

Causus rhombeatus Bouleng. t. c. p. 467.

Tropical and South Africa.

CAUSUS RESIMUS.

Heterophis resimus Peters, Mon. Berl. Ac. 1862, p. 277, pl. —,

fie. 4.

Causus reszmus Bouleng. t.¢. p. 468.

Eastern Soudan, East and Central Africa, Angola.

2. Brivis.

Gray, Zool. Miscell. p. 69; Bouleng. Cat. Sn. ii. p. 492.

1. Brivis ARIETANS

Vipera arietans Merv. Tent. p. 152.

Bitis arietans Bouleng. t. ce. p. 493.

Tropical and South Africa; Southern Arabia.

3. CBRASTES.

Wagl. Syst. Amph. p. 178; Bouleng. Cat. Sn. ni. p. 501.

1. CERASTES CORNUTUS.

Coluber cornutus Linn. 11 Hasselq. Reise Paleest. p. 315.

Cerastes cornutus Bouleng. t. ¢. p. 902.

Soudan and borders of the Sahara; Arabia and Palestine.

4. Kouts.

Merr. Tent. p. 149; Bouleng. Cat. Sn. i. p. 504.

Two species :—

Scales on snout and vertex more or less strongly keeled; 2 (rarely 1

or 3) series of scales between eye and upper yes. ventrals

NePSIB PA abanoce soespese Jeb AH RUR MIS

Scales on snout and. vertex. smooth or 1 obtusely ‘keeled ; 3 or 4 series

of scales between eye and upper labials ; ventrals 174-205 eee EL. coloratus,

1. EcuIs CARINATUS.

Pseudoboa carinata Schneid. Hist. Amph. 11. p. 285.

Iichis carinatus Bouleng. t. c, p. 505.

Desert and sandy districts of Africa north of the Equator ;

Southern Asia, from Transcaspia and Arabia to India.

2. Ecuis cotoratus Gunth. P. Z.8. 1878, p.978; Bouleng. t.c.

p- 507, pl. xxv. fig. 1.

Eeypt, Socotra, Arabia, Palestine.

658 ON THE SNAKES OF NORTH-EAST AFRICA.

5. ATRACTASPIS.

A. Smith, Ill. Zool. 8. Afr., Rept.; Bouleng. Cat. Sn. iil.

p- 510.

Synopsis of the Species.

I. Anal divided ; subcaudals paired ; 4th upper labial entering

the eye; scales in 19 rows ........ coe A. engdahlii.

IT. Anal entire; subcaudals all or satan fens

4th upper labial entering the eye; a single large anterior

temporal; scales in 31 rows ee cssbseeeeteeensne UAauphoitipse:

3rd and 4th upper labials. entering the eye ; ; temporals small ;

scales in 23 rows ...... ... <A, lewcomelas.

8rd and 4th upper labials entering the eye; temporal small ;

SEMIS Uy AY tO BP TOWS coocoocovesncnoscoasecooene ado coocv vanoonceoonn Ale PHACROUGOUCaLiE,

1. ATRACTASPIS ENGDAHLIIT Loénnb. & Anderss. Ark. f. Zool. viii.

1913, no. 20, p. 5.

Somaliland.

2. ATRACTASPIS PHILLIPSI Barbour, Proc. Biol. Soc. Washingt.

1913, p. 148.

Sennar.

3. ATRACTASPIS LEUCOMELAS Bouleng. Ann. Mus. Genova (2) xv.

1895, p. 16, pl. iv. fig. 2, and Cat. Sn. iii. p. 517.

Somaliland.

4, ATRACTASPIS MICROLEPIDOTA Giinth. Ann. & Mag. N. H. (3)

xvill. 1866, p. 29, pl. vil. fig. 3; Bouleng. t.c. p. 517.

Egyptian Soudan, Hast and Central Africa,

ON TWO NEW COLOMBIAN REPTILES. 659

47. Descriptions of a new Amphisbena and a new Snake

discovered by Dr. H. G. F. Spurrell in Southern

Colombia. By G. A. BouLEencsr, F.R.S., F.Z.8.*

{ Received October 29, 1915: Read November 23, 1915. |

(Text-figures 1 & 2.)

AMPHISB£NA SPURRELLI, sp. n. (Text-fig. 1.)

Snout rounded, prominent. Rostral rather small, triangular,

just visible from above; nasals forming a suture; a pair of

Text-figure 1.

' Amphisbena spwrrelli. Upper, lower, and side views of head, and lower view

of posterior part of body and tail.

660 MR, G. A. BOULENGER ON

preefrontals, the largest head-shields, the suture between them

once and a half to once and two-thirds as long as that between

the nasals and slightly longer than that between the frontals,

which are much smaller and a little longer than broad * ;

a pair of small occipitals; eye distinguishable under the ocular,

which is situated between the prefrontal and the second and

third upper labials and followed by two superposed postoculars ;

four upper labials, fourth small, second very large; symphysial

trapezoid, the sides slightly concave, followed by a longer chin-

shield ; four lower labials, second largest. 213 or 214 annuli on

the body, 21 to 23 on the tail; an annulus in the middle of the

body contains 34 or 36 segments, 16 or 18 dorsal and 18 ventral,

the dorsals at least twice as long as broad, the median ventrals

once anda half to twice as broad as long ; the dorsal segments on

the distal half of the tail tubercular or subconical. Lateral Jine

very distinct. 6 anal segments. 4 preanal pores. Whitish,

each dorsal segment with an oval or squarish purplish-brown

spot.

Length to vent 320 mm. ; tail 28; diameter of body 10.

Two specimens from Anda Goya, at the junction of the R.

Condoto and San Juan.

Distinguished from A. darwint D. & B. by the broader ven-

tral segments and the tubercular segments on the tail.

HERPETODRYAS VICINUS, sp. n. (Text-fig. 2.)

Eye large, as long as the snout. Rostral once and one-third

as broad as deep, just visible from above ; internasals a little

broader than long, a little shorter than the prifrontals ; frontal

Text-figure 2.

Herpetodryas vicinus. Side view of head.

once and a half as long as broad, longer than its distance from

the end of the snout, slightly shorter than the parietals; loreal

as long as deep; one pra- and two postoculars; temporals 2+ 2 ;

nine upper labials, fourth, fifth, and sixth entering the eye;

five lower labials in contact with the anterior chin-shields, which

are shorter than the posterior. Scales in 10 rows, all smooth or

the vertebral pair very faintly keeled; scales gradually decreas-

ing in size towards the ventrals, the largest not half the size of

the parietal shields. Ventrals not angulate laterally, 163; anal

* In the specimen figured the left frontal has tused with the left occipital.

TWO NEW COLOMBIAN REPTILES. 661

divided ; subcaudals 149. Reddish brown above, anterior two-

thirds of body with darker lines on the borders of the scales,

forming faint cross-bands terminating in a spot on the outer

series of scales; a series of irregular yellow spots or streaks along

the vertebral line; posterior part of body and tail uniform brown

with darker edges to the scales; a round blackish spot on each

parietal shield ; a short black streak behind the eye above the

lip which is white, some of the shields finely black-edged ; belly

white in front, brownish behind; a dark streak along the tail at

the junction of the subcaudal shields.

Total length 490 mm.; tail 190.

A single young specimen, also from Anda Goya.

Closely allied to H. fuscus L. Distinguished by the number of

temporals (2+2 instead of 141), the greater extent to which

the fourth upper labial borders the eye, the divided anal, and

the more numerous subcaudals.

haa eve av

he June 6 (i ny

hie nage alanis,

i eas, davies” 1 Mes

Bry ai peat lrbias) pre

heap

ON SECONDARY SEXUAL CHARACTERS IN BIRDS. 663

48. The Distribution of Secondary Sexual Characters

amongst Birds, with relation to their Liability to

the Attack of Enemies. By J. C. Morrram,

M.B. (Lond.)*.

[Received June 25, 1915: Read November 9, 1915.]

INDEX.

Morpnoboey : Page

Secondary sexual characters. Distribution among birds ... 663

ETHOLoeGY :

Secondary sexual characters. Relation to enemies............ 663

GEOGRAPHICAL:

Secondary sexual characters. Insular and continental ...... 671

This paper endeavours to show that there is a correlation

between extra-sexual dimorphism among birds and their vulner-

ability to enemies. No attempt is made to account for this

correlation. However, in the light of this correlation, the more

important theories which have been put forward to account for

the presence of secondary sexual characters are briefly con-

sidered.

Collection of Material.

In collecting material for classification, works were selected in

which many facts were collected on broad lines and with no

particular object in view. References are given for all the

material used. The work from which by far the most material

was taken is ‘The World’s Birds, by F. Finn. This is a small

volume, in which the families are briefly described in a syste-

matic manner under the headings—diagnosis, size, form, plumage,

young, nest, eggs, incubation, courtship, food, gait, flight, notes,

disposition and habits, economic qualities, captivity and dis-

tribution. A wealth of reliable information is here laid down

in a condensed manner, and is, therefore, at once available for

classification.

Apart from this work, descriptive monographs have been

chiefly consulted. Only families containing more than ten

species are considered, as it was thought that a lesser number

could not give a reliable picture of the characters of a family.

In order to escape any misunderstanding, the factors to be

correlated must first be clearly defined. A species is said to

show extra-sexual dimorphism when there isa difference between

the somatic characters of male and female. Attention is usually

directed to differences in form and colour, but it must be remem-

bered that almost any character may show a difference. It is by

no means uncommon to find birds in which the form and colour-

ing of the sexes are identical, but it is quite rare to observe birds

* Communicated by the SECRETARY.

664 DR. J. C. MOTTRAM ON SECONDARY

the males and females of which are entirely alike in every respect

apart from their sexual and reproductive organs. Many, in

describing the Song-Thrush, remark “sexes alike,” forgetting

the male’s song; nevertheless, it is necessary in a consideration

of this kind to accept the classifications of others.

Vulnerability to enemies is essentially a relative term. Un-

fortunately a classification of birds according to their vulnerability

to enemies has not been found. It follows that this subject must

be considered in detail.

Birds are subject to attack from the air by hawks, whilst

feeding in- trees from carnivorous mammals and reptiles, and

when on the ground from all three. It follows that birds

entirely aerial are more free from enemies than are arboreal or

terrestrial ones, and birds entirely arboreal more free than

terrestrial. All birds, with very few exceptions, are to some

extent aerial. Sea-birds must be almost as free from enemies as *

aerial birds, as there is little evidence that they are preyed upon

by any aquatic animals; and of sea-birds, the oceanic ones must

be especially free from enemies, as hawks are not found far from

land. The following table, compiled from Finn’s ‘ World’s Birds’

(see Table I.), sets out the families from this point of view, and

shows that birds relatively free from enemies present little

sexual difference, whereas those presenting great sexual differ-

ences are only to be found amongst terrestrial and arboreal

birds.

TABLE I.

Terrestrial. Arboreal. Aerial. Aquatic. Maritime.

1. No sex wh On 2 ae es] Ba j ! / /

| difference ... 38% 8 [35%] | 25% 4 [17%] | 50% 4 [17%] | 100% 2 [9%] | 83% 5 [22%]

12. Usually no 24.0), 5 [33%

sex difference

. Sometimes

sex difference

Oceanic and

31% 5 [33% |

50 %/p 4 [27%]

16% 1 [7 %pj

9% 2 [40%]

19%, 3 [60%]

. Usually sex

difference

| 29% 6 [60%] |

25 0% 4, [40 0 0]

Norrs.—Compiled from F. Finn’s ‘ The World’s Birds.’

2= Usually no sex difference, and includes the following descriptions :—“ Hardly

ever a sex difference,

difference,

DIS

22 6

alike,” “ usually small s.d.”

4=Usually sex difference, and includes the following descriptions :—“ Sexes

different,” “ often sex difference.”

The above method of classifying secondary sex characters is used throughout the

paper. ; :

seldom a sex difference,

small sex difference,’ “little s.d.,” “rare s.d.,” “ oftenest

22 66

(Continued at foot of next page.)

very slight or no sex

SEXUAL CHARACTERS IN BIRDS. 665

The subject may be approached from other points of view.

Many birds are more or less free from enemies on account

of possessing powerful weapons of defence — either beaks,

talons, wings (e. g., swans), or legs (é. g., cassowaries). Sexual

dimorphism should be less commonly found among these birds

than among birds in general; and such is the case, as shown

in Table II., only hornbills and toucans being to some extent

Tase IT.

Po Neaeerave | With defensive

| weapons. weapons.

——— | —

1. No sex difference............ a 38%) 11 | 520, 11

2. Usually no sex difference | 24% 7 88% 8

A i | > § Toucans &

- : 0/ | 0 5

3. Sometimes sex difference “| 10% 3 10% 2 { TELNnDNIDe

4. Usually sex difference ...... | 28% 8 — (0)

Norrs.—Compiled from F. Finn’s ‘The World’s Birds.’

Secondary sex characters classified as before.

Reference Nos. :—1 no def. 4, 8, 22, 25, 28, 29, 30, 32, 40, 45, 46: 2 no def. 10, 26,

31, 37, 39, 42, 44: 3 no def. 11, 21, 38: 4 no def. 5, 14, 16, 19, 24, 34,

41,49: 1def. 1, 2, 6. 9, 13, 17, 18, 35, 36, 43, 48: 2 def. 3, 7, 12, 15, 20,

27, 38, 50: 3 def. 23, 47.

exceptional. On referring to Table III., it can be seen that

these two families have been correctly described as exhibiting

“sometimes sex difference,” especially in the case of hornhills.

As regards toucans, it can be seen that it is chiefly among the

smaller and therefore less powerful species that sex dimorphism

is found. This, however, is not the case in the hornbills.

Although their bills are apparently very powerful, it is doubt-

ful whether they are so in practice; keepers are of opinion that

Notes to Table J., continued.

In a few cases birds have been placed in two columns, e. g. Pigeons in Tervestrial

and Arboreal, Passerine Birds ditto, Divers in Aquatic and Maritime.

Ducks have been placed under Terrestrial, as they are especially lable to the

attacks of carnivorous mammals whilst feeding on dry land or in shallow

water; they are considered in detail elsewhere.

The bracketed percentages refer to the horizontal lines, the unenclosed percentages

to the vertical columns.

Reference numbers for Table I. :—

1 Terr. 6, 8, 9, 25, 29, 48, 45, 46: 2.Terr. 20, 37, 39, 42, 33: 3 Terr. 11,

23: 4 Terr. 34, 6, 14, 16, 19, 41: 1 Arb. 4, 28, 30, 48: 2 Arb. 3, 26, 33, 37,

50: 3 Arb. 23, 38, 47: 4 Arb. 24,49, 34,16: 1 Aer. 1, 4, 12, 82: 2 Aer. 15,

31, 40, 44: 1 Aq. 18, 17: 1 Ocea. 2, 18, 18, 35, 36: 2 Ocea. 7: Nos. 10, 21,

22, 27 omitted.

These numbers refer to the list of birds at the end of the paper, and show how

they are distributed in the table; they enable others to criticise the distri-

bution.

666 DR. J. C. MOTTRAM ON SECONDARY

a severe bite can only be given when the finger is seized far back

in the bill: at the tip the leverage against the jaw-muscles must

be very great, and, unlike most long bills, they are unsuitable for

stabbing.

Tasxe IIT.

| as | ssl | sl.d. | so.d. d |

| | | |

; | ae el

No. of species ......... eels Wf | Bl 3 11 ”) |

| | Hornbills. |

| Average wing-length. 12°5 154 108 | Ils 151 i) |

| |

} | |

| No. of species ......... | PE Hh 2) 1 6 ? |

| | ! Toucans. |

_ Average total length. 18°3 | Wer Teles 13°5 13 , |

Norszs.—Compiled from Monographs on the Bucerotide by D. G. Elliot and the

Ramphastide by J. Gould.

a.=sexes alike; v.sl.d.=sexes very slightly different; sl.d.=slightly different ;

so.d.=some difference; d.=different; v.d. (used in other tables) =very

different.

If, instead of considering birds from the point of view of

defence we classify them according to their offensive powers,

the same conclusion results. Offensive powers are divided

according as to whether the birds’ diet consists of vertebrates,

small vertebrates, invertebrates, or vegetable matter, as set out

in Table ITI. a.

Tasie III. a. (Food of Birds.)

| | Food consisting of :—

| | Tne Small Vertebrates Small Vertebrates,

| | Vertebrates. | and Invertebrates. | me veeene

| 1 8 7 | 8

DQ. 4 L 3 | 7

3 , " | 2 3

A. “| = — 8

Norrs.—Compiled from F. Finn’s ‘The World’s Birds.’

Numbers at column headings refer to secondary sex characters as before.

Reference Nos. :—1 vert. 1, 2, 13, 18, 32, 35, 36, 43: 2 vert. 7, 12, 20, 27: 1 vert.

& invert. 4, 8, 22, 25, 40, 44, 45: 2 vert. & invert. 15, 26, 31: 3 vert. &

invert. 21, 28: 1 vert. & veg. 6, 9, 17, 28, 29, 30, 46, 48: 2 vert. & veg. 3,

10, 33, 37, 39, 42, 50: 3 vert. & veg. 11, 23, 47: 4 vert. & veg. 5, 14, 16, 19,

9A, 34, 41, 49.

Further, certain birds for other reasons are especially free

from enemies. Diving-birds have always a safe refuge at hand,

SEXUAL CHARACTERS IN BIRDS. 667

and should, therefore, present little secondary sexual differ-

entiation. This is shown to be the case in Table IV. In this

Tasie IV.

| : |

Diving Birds. | Non-diving Birds. |

lenlit ecrcc asc taca eee eel 4 66% | 18 42%

2s Aen Rr itil fots:| UO) a EPs

| 3 o = | 5 12%

eh ekae eonibeneeeeprncs one i (Of, 7 VA,

| (Ducks)

Nores.—Compiled from F. Finn’s ‘ The Worlds Birds.’

Sexual classification as before.

Reference Nos. :—1 Div. 3, 13, 17, 36: 2 Div.7: 4 Div. 14: 1 Non-div. 1, 4, 6, 8,

9, 18, 22, 25, 28, 29, 30, 32, 36, 40, 43, 45, 46, 48: 2 Non-div. 3, 10, 12, 15,

20, 26, 27, 31, 33, 37, 39, 42, 44, 50: 3 Non-div. 11, 21, 23, 38, 47: 4 Non-

div. 5, 16, 19, 24, 34, 41, 49.

table ducks form an exception—for, though capable of diving,

they often present sex dimorphism. Ina previous table they

were placed with difficulty, so that it is necessary to study them

in more detail (see Table V..). This table shows that the diving-

TABLE V.

| Diving Ducks. | Non-diving Ducks.

| |

| ae, Meld: slide de vd: ||

}

— 2 3 3 a |

P wellol Sl cb v.d.

— — — — 10

Norrs.—Compiled from J. Gould’s ‘ Birds of Europe,’ vol. v.

Swans, Geese, and Sheldrakes. presenting little or no sex difference, were omitted.

The following list shows the placing of the ducks in the above table :—

Div. v.sld. Fuligula leucophthalmos, Undina leueocephala: Div. sl.d. F. cris-

tata, Oidemia fusca, O. nigra: Div. d. F. ferina, F. rufina, Mergus

albellus: Diy. v.d. F. marila, Clangula vulgaris, C.histrionica, Harelda

glacialis, M. merganser, M. serrator, M. cucullatus: Non-div. v.d.

Mareca peneiope, Spatula clypeata, Anas boschas, Querquedula crecca,

Q. glocitans, Q. circia, Dafila caudacuta, Chauliodes strepera, Somateria

mollissima, S. spectabilis.

habit is associated with a reduction in the amount of secondary

sexual difference. No doubt, had a larger amount of material

been examined, exceptions would have been encountered ; for

instance, in the Spot-bill (Anas pecilorhyncha) the sexes are

668 DR. J. C. MOTTRAM ON SECONDARY

alike, and perhaps the duck might be considered to he a non-

diver, but KE. C. Stuart Baker in ‘Indian Ducks’ says “it isa

most expert diver.”

The Marbled Duck (Marmaronetia angustirosiris) forms a like

exception, but has unusual habits, which to a large extent

protect it from enemies. It never goes ashore; it remains

nearly always in rushes. The Andaman Teal (WVettion albigulare)

is another, but this bird is an insular species and is said to be

entirely nocturnal.

However, examples more difficult to account for are to be

found ; for instance, the Whistling Ducks (Dendrocygna). These

birds have arboreal habits, and at a sign of danger take to the

trees. On the whole, therefore, although at first sight appa-

rently serious exceptions, on more detailed examination ducks

are found to fall into line with the rule that diving-birds exhibit

less secondary sexual differences than non-divers.

Very few, if any, birds can entirely escape from hawks by

means of rapid flight (arboreal birds are to some extent free

from their attacks). Two of the favourite quarries of the

HKastern hawkers are teal and sand-grouse, among the most rapid

fliers. There are, however, a few birds which are able by .

means of skilful dodging to evade the attacks of hawks—for

instance, the hoopoe, of which Finn says “it is easily able to

escape from hawks.”

TABLE VI.

Birds mentioned as | ot Bact a

| forming food for man. | No note re edibility. |

eat eae cn 44 0/, 12 | 430), 10

OF hae eae 220), 6 39% 9

Honey-Guides).

9% 2 (Trogons and

Autre hal Seba ae Name 229%, 6

Hummine-birds).

Sos Renae 11%, 3 | 9%) 2 (Puff-birds and

Norrs.—Compiled from F. Finn’s ‘ The World’s Birds.’

Honey-Guides are not eaten by natives because they are useful in guiding them

to honey ; Humming-birds possibly because of their small size. Puff-birds

are doubtfully described as presenting “sometimes sexual differences,’ as

shown in the following table compiled from P. Li. Sclater’s ‘ Bucconida’ :—

P a. sl.d.

PuttebmdSieee es aoe 8 g

2—no note made as to whether sexes are different or not.

As regards Trogons, other works have been consulted but with no result.

Reference Nos. :—1 edib. 2, 9, 22, 25, 29, 30, 32, 35, 36, 45, 45, 46: 2 edib. 7, 20,

33, 37, 39, 42: 3 edib. 11, 23, 47: 4 edib. 5, 14, 16, 19, 34, 41: 1 non-edib.

1, 4, 6. 8, 13, 17, 18, 28, 40, 48; 2 non-edib. 3, 10, 12, 15, 26, 27, 31, 44, 50:

3 non-edib, 21, 28: 4 non-edib. 24, 49.

SEXUAL CHARACTERS IN BIRDS. 669

Birds may avoid enemies in other ways; there is little doubt

but that ill-flavour playsa part. Under the heading “ Economic

Qualities,” Finn, in the ‘ World’s Birds,’ mentions those birds

which form food for man, and this to some extent will separate

' the well-flavoured from the ill-flavoured. This material is set

out in Table VI., which shows that secondary sexual dimorphism

is especially to be found among table-birds.

Birds may also be said to avoid enemies by protective color-

ation. However, it is not possible to examine this aspect statis-

tically, because protective coloration often forms a part of sex

dimorphism, the female presenting this coloration and the male

not doing so. ‘This is in itself, however, evidence that sex

dimorphism is in some way related to enemies.

Size.—Upon size must depend to some extent aspecies’ vulner-

ability to enemies. Large birds are only open to the attack of

large carnivorous animals, whereas small birds are preyed upon

by both small and large. Large birds should, therefore, show

less secondary sexual dimorphism than medium-sized birds, and

these less than birds of small size. Finn describes the size of

birds by comparing them with well-known English ones—for

instance, of parrots he says “less than tit to that of an ordinary

fowl,’ of hornbills “from that of a hen turkey to that of a

pigeon.” Birds thus used in description may be classified

into large, medium, and small, and, with this as a measure,

Table VII. has been compiled. Parrots would fall, therefore,

into all three columns, and hornbiils into medium and large.

Tasue VIL.

| ut Large. | Medium-sized. | Small. |

ee eg BOO oil vim (480 18 Buy |

| |

OR ics ate Neh AY, 2 | BO TS 49/0), 4 |

5 le ieee 10%, 2 10% 4 Ome

Ala vconnseee NY es NO RIE OFAN. WO, i Nl OR |

| |

Norts.—Compiled from F. Finn’s ‘The World’s Birds.’

Table of measure for classification :—Large=swan, goose, turkey, “upwards,”

heron, 3 ft. and above in length: Medium=jay, dove, raven, rook,

mallard, partridge, crow, fowl, pheasant, “ medium size,” pigeon, jackdaw,

hen, duck: Small=lark, “very small,” thrush, sparrow, blackbird, tit

swallow, quail, starling, linnet, “3 inches.”

Reference Nos.:—1 L. 1, 2, 6, 9, 18, 18, 25, 32, 35, 36,43: 2 L. 7, 12, 20, 37:

3 L. 11, 23: 4 L. 5, 14, 16: 1 M. 1, 2, 4, 8, 13, 17, 18, 22, 25, 28, 29, 32, 35,

36, 40, 45, 46, 48: 2 M. 3, 7, 10, 12, 15, 20, 27, 31, 33, 37, 39, 42, 50:

3 M. 11, 23, 38, 47: 4M. 5, 14, 16, 19, 34, 41, 49: 1S. 2, 4, 8, 17, 18, 22,

28, 30, 32, 36, 46: 2S. 3, 10, 12, 15, 20, 26, 27, 31, 33, 37, 39, 42, 44, 50:

3S. 21, 38, 47: 4.8. 16, 19, 24, 34, 49.

Secondary sex characters classified as previously.

Proc. Zoon. Soc.—1915, No. XLVI. AG

670 DR. J.-C. MOTTRAM ON SECONDARY

The table shows that 15 per cent. of large birds, 17 per cent.

of medium, and 15 per cent. of small often present sex differences.

In this respect there is little difference, but of large birds

55 per cent., of medium-sized 43 per cent., of small 33 per cent. —

present no sex difference. Examination horizontally also shows

that large birds less often present secondary sexual dimorphism

than the medium-sized or small birds.

Instead of studying birds as a whole, certain families, chosen

at random, have been examined by themselves with respect to

the question of size. In barbets, jacamars, puff-birds, wood-

peckers, and toucans the Tables III. and VIII. show that the

smaller species present the most secondary sexual dimorphism.

In hornbills and kingfishers, however, this association is not

found, though the reverse does not hold.

Other families show the effect of size without any detailed

analysis ; for instance, in the Anatide swans and geese present

little or no sex difference, and in the Falconide it is only the

smaller species which show any sex difference.

On the whole, therefore, the consideration of size appears to

support the thesis.

Vase VIil.

ae vesl.d. sl-di’ sozd! d. v-d..

(No. of species ...... 29 — 1 — il 3

| | Average total length 8°0 — 7-4, —- 6:0 58

Barbets ...... ———— : : : SCE GS a ee

| No. of species ...... 43 1 1 1 3 4,

LAverage wing-length 344 ol oA a3 3:23 2°69

No. of species ...... 1 8 5 1 — —

Jacamars ... f I

@ Averaze total length 11 8-2 ae 8 — —

Pareniede ; No. of species ...... 8 — 2 — — ==

| CAverage totalleneth 7°5 = 70 = = --

Raneheier § No. of species ...... 13 2 a ill 1 1

a. CU Average length ...... 95 10:2 120 100 85 9-0

7. § No. of sjecies --.... 1 103 100 if 3 ==

eee arate ayant ae 24 | ea

Norrs.—Compiled from Monographs on the Capitonidze by C. T. & G. lL. Marshall ;

Galbulide aud Bueconide by FP. L. Sclater; Alcedinidie by R. B. Sharpe;

Picidee by Alf. Malherbe.

For column headings, see Table III.

Sociability.—This is another factor which should protect birds

from enemies to some extent. Numbers may be able to resist

an enemy to which one would fall a victim. Sociable birds

should, therefore, show less secondary sexual dimorphism—as 1s

the ease, see Table LX.

SEXUAL CHARACTERS IN BIRDS. 671

TABLE LX.

Sociable Birds. | Not Sociable,

en —a Eee ee

Wyre Ds seas cals Leer 12 | 4

Sh eM Ries 1 | 1

Nores.—Compiled from F. Finn’s ‘The World’s Birds.’

Of the remaining birds no note is made except Passerine birds, which are described

as sometimes sociable.

Some birds form coveys or family associations out of the breeding-season—geme-

birds, for instance, and other birds form flocks, e. g. ducks.

Reference Nos. :—I Soc. 1, 2, 4, 9, 18, 22, 25, 30, 35, 36, 43, 48: 2 Soc. 7, 33, 42,

44; 3 Soc. 47: 1 Non-soc. 8, 28, 32,45: 2 Non-soc. 3, 10, 12, 20, 26, 27,

31, 39,50: 3 Non-soc. 21: 4 Non-soc. 5, 16, 19, 24, 49.

Secondary sex characters classified as before.

Birds may also avoid enemies by having a habitat where they

are free from enemies—for instance, mountain-tops, islands,

arctic regions, oceans. Oceanic species are included in Table J.

The only other material with which it has been found possible to

examine this question is Seebohm’s monograph of the Turdide ;

for in this family there is a sufficient number of insular and

continental forms for comparative purposes. Table X. sets oat

the material, and it can be seen that secondary sexual dimorphism

is much less common in insular than in continental species. Of

the four continental species of the genus Merula which present

little secondary sexual dimorphism, it is noteworthy that there

are three living high up on volcanic mountains.

TABLE X.

on ae = eS See E see oe Sh

| Insular. Continental.

Hee NES a eu |

| | a wesilel, lll sod Gh wel |i eo spell sl.d. so.d. d.

| Merula ........| 6 9 3 Wet OG) 2 2 A Wile aS

| | 2 volcanic 1 volcanic

species. species.

Geocichla ...| 14 0) 2 (0) @ i 18 0) 2 0 it

| LLOUREMIS orpocs sce 8 36 = 6 = 2

| Mimocichla 3 Hee —=

Nort.—Compiled from H. Seebolhm’s ‘ Turdidee.’

Endeavours were made to find other material in order to sub-

stantiate or not this remarkable correlation, but none was found

suitable for classification.

46*

672 DR. J. C. MOTTRAM ON SECONDARY

Song, another secondary sexual character, was also treated in

the same way, and clearly substantiates the former finding, as is

shown in Table XI.

TABLE XI.

Insular. Continental.

| Sing. ? No song. Sing. ? No song.

Mle alee eee se 5 18 3 17 21 1

Gori et 28 15.) ly org, ache ae

TUceestene ah iho Lie i ayqb Ol eek: MSD RSL RAN

Nore—Compiled from H. Seebohm’s ‘ Turdidee.’

2—No uote was made with reference to song. No song includes such descriptions as

“song not known,” “bird is silent,” “never heard.”

Birds may be looked at with advantage from another point of

view. In Table XII. they are classified according to their sexual

dimorphism without reference to any other factor, Examin-

ation of this table (XII.) shows that among birds presenting no

sexual difference (column 1) 10/22nds 45 per cent. are relatively

free from enemies; among birds usually presenting no sex dif-

ference (column 2) 5/15ths 33 per cent.; among birds sometimes

presenting sex difference 2/5ths 40 per cent., and among birds

usually presenting sex difference 0/8ths 0 per cent. are free from

enemies.

Taste XIT.

ay eldemier® 3. 4

Nate UNO ie | INST ol | Wale

WO 12) 5. 10 mare On ec

45 9 33. %/p 40% | 0%

Nores.—Compiled from F. Finn’s ‘The World’s Birds.’

Numbers at column headings refer to secondary sex characters as before. N.L.=not

liable to attack ; L. liable to attack ; °% = percentages of birds not especially

liable to the attack of enemies. ‘This is a synopsis of the previous table.

Reference Nos. :—1 N.L. 1, 2, 6, 18, 17, 18, 32, 35, 36, 43: 1 L. 4, 8, 9, 22, 25, 28,

29, 30, 40, 45, 46, 48: 2 N.L. 7, 12, 20, 33, 44: 21. 8, 10, 15, 26, 27, 31,

37, 39, 42,50; 3.N.L. 23,47: 3 L. 11, 21, 38: 4 L. 5, 14, 16, 19, 24, 34,

41, 49.

Thus it is clear that birds liable to attack more commonly

present secondary sexual differences than birds not so lable to

attack. If, instead of considering birds from a broad standpoint,

a more detailed analysis be made, the same result is arrived at. In

SEXUAL CHARACTERS IN BIRDS. 673

the case of passerine birds, powerful species are only to be found

amongst those showing no sex dimorphism, e. g., crows, drongoes,

shrikes. Swallows, which are aerial birds, present little or no

sex dimorphism. |

The classification of sexual dimorphism chosen for analysis

refers almost entirely to sexual dimorphism of colour; neverthe-

less, it will be advantageous to consider briefly other secondary

sexual differences—song and precopulatory displays.

As regards song, this character is frequently to be found in

birds presenting no sex differences in colour and form, but which,

nevertheless, do not appear to be especially free fromthe attack

of enemies—-for instance, among warblers, American warblers,

wrens, cuckoos, larks, pigeons; whilst among birds not lable to

attack, song-birds are very uncommon, if not entirely absent.

It thus appears that this dimorphic character is correlated to

vulnerability to enemies, and accounts for some of those cases

of birds presenting little or no sexual dimorphism of form or

colour, but which are yet liable to the attack of enemies.

Precopulatory displays, or the displays of courtship, is another

secondary sexual character open to investigation. Here, again,

birds showing little or no display are birds not lable to attack,

whereas birds showing great display are especially lable to attack.

In the ‘ World’s Birds,’ under the heading ‘‘ Courtship,” sexual

displays are briefly described in 25 of the 50 families under con-

sideration. In the case of 19 of the 25 the usual actions are

described : against diurnal birds of prey there is the remark

“never seen the display,” divers “said to be on the wing,”

horn bills “ little known”; whilst against bustards ‘‘very elaborate,”

game-birds “ elaborate,” cranes “ very elaborate.” Cranes, then,

form an exception, for they possess weapons of defence. Never-

theless, certain factors might lead one to suspect that they

are not entirely free from attack; they are terrestrial birds, they

are excellent-eating (their food being mostly vegetables, seeds,

etc.),and in the East they are a favourite quarry of the hawker ;

they are also usually not sociable during the breeding-season,

and, lastly, in most species both sexes take equal part in the

displays, so that in truth they do not present secondary sexual

dimorphism in this respect.

It is very fortunate that cranes needed this detailed considera-

tion at the end of the examination, which has covered a great

deal of material, as it is easy to draw attention to the fact that

this is almost the only oecasion (ducks also) when such treatment

became necessary.

Summing up, it may be said that for certain reasons—namely,

because of aerial, aquatic, and oceanic habits, possession of defen-

sive and offensive weapons, of diving-habits, of ill-flavour to man,

of large size, of insular habitat, of social disposition, of carni-

vorous habits—certain birds must be relatively free from the

attacks of enemies; and it has been shown that these birds

present less secondary sexual dimorphism than birds which ‘for

674 DR. J. GC. MOTTRAM ON SECONDARY

other reasons—namely, possession of terrestrial and arboreal

habits, of no defensive weapons, of non-diving habits, of no ill-

flavour to man, of small size, of continental habitat, of solitary

disposition, and of non-carnivorous habits—have been considered

to be relatively lable to the attack of enemies. These two

factors thus appear to be correlated, but it is obvious that, though

erveat sexual dimorphism is only to be found amongst birds lable

to attack, nevertheless the converse does not hold, for some

birds which do not appear to possess any special characters

which would protect them from attack, none the less fail to

present any secondary sexual dimorphism, e.g. ibises, mound-

birds, mouse-birds, rollers, barbets, and coursers.

Conclusions may be briefly stated :—

(1) Among birds specially liable to the attack of enemies,

secondary sexual dimorphism is chiefly to be observed.

(2) Among birds not specially liable to attack of enemies,

secondary sexual dimorphism is rarely found.

(3) In both cases, birds presenting no sexual differences are

‘represented, but much more commonly in birds more or

less free from attack of enemies.

Having come to these conclusions, it will not be out of place

to examine briefly the more important theories put forward to

account for secondary sexual dimorphism, in order to discover

whether any of them be substantiated or not.

Huuberance of Vital Energy in’ Males.—Wallace, in ‘ Tropical

Nature,’ believed that natural selection could account for second-

ary sexual dimorphism, by supposing that only the strongest and

most virile males gained mates, and that these mates thus came

to possess an excess of vital energy which became manifest in

bright plumes, ete. If this be so, it is difficult to see the reason

why birds vulnerable to enemies should be specially picked out.

One would think that it would be dangerous for these birds to

make themselves thus conspicuous, and would have anticipated

that birds not lable to attack could have done so with more

impunity.

Sexual Selection (Darwin, ‘ Descent of Man’)—The argument

used above apples similarly against Darwin’s theory, which in

no way can account for the correlation under examination. One

would have thought that female selection would have had freer

play among birds more or less free from enemies than among

birds frequently subject to their attack.

Nesting-Habits Theory (Wallace and others).—It has been noted

that birds lable to the attack of enemies, and making open nests,

more frequently present secondary sexual dimorphism than birds

making closed nests. If, however, birds liable to the attack of

enemies more often present secondary sexual dimorphism than

birds in general, then it follows that birds making open nests,

and also lable to the attack of enemies, must do likewise.

It is thus obviously necessary to decide which is the more

SEXUAL CHARACTERS IN BIRDS. 675

important correlating factor—vulnerability to enemies or open-

nesting habits. Table XIII. deals with birds making open nests

and in no way helps to decide the point; coursers, ibises, mouse-

birds, tinamous, touracous, pigeons, rails, shore-birds, according

to the “ nesting-habit hypothesis,” should present sex dimorphism,

but they also fail to fall into line with the other correlation.

Taste XIII. (Open-nesting birds.)

Not especially liable to attack. | Liable to attack.

| jo ES CS eee

American Vultures, Cassowaries . : :

ces : a “4,2 | Coursers, Ibises, Mouse-birds, Tina-

1. Cranes, Divers, Grebes, Gulls. i x

CeO DAneuinie 8 mous, Touracous. 5

s, ouins.

Cormorants, Birds of Pre ners Q .

2. eicrona! TETAS 5) ay Pigeons, Rails, Shore-birds. 3

s ehtjars |

et ee oe a | - iu

Bh - | Curassows. 1

| Bustards, Game-birds, Humming-

A, rere | We J 4 2) =

| birds, Sand-Grouse.

If, however, one turns to closed-nesting birds (Table XIV.),

it is at once obvious that open-nesting habit plays no part,

because several of the groups present considerable secondary

sexual dimorphism, and quite a large number a small amount.

Taste XIV. (Closed-nesting birds.)

1. Bee-Haters, Hoopoes, Rollers, Motmots. 4

2 Barbets, Jacamars, Kingfishers, Parrots,

~ Woodpeckers. 5

3. Hornbills, Toucans. 2

4. Trogons. 1

Nors.—Compiled from F. Finn’s ‘The Worlds Birds.’

Table XV. continues the two previous tables, and also deals

with families of birds making both open and closed nests. It

shows that the relation between open-nesting habits and second-

ary sexual dimorphism is, if not entirely negligible, at any rate

not nearly so close as with vulnerability to enemies. It may be

676 DR. J. C. MOTTRAM ON SECONDARY

mentioned that the Australian warblers (Maluride) present great

xex difference (the males are amongst the most brilliant of birds),

and yet they make closed nests. They are small defenceless

birds, and so they not only form exceptions to the open-nesting

theory, but support the correlation under consideration.

TABLE XV. a cane )

ier |

Opeu nests. Cine aodyclosed Closed nests. |

nests. |

gee eal eels 65 9p 3 WO, | 4 209/, |

520), | 330 | 330 i

ELAR aor (ered orem 2 Fe

| | |

Deanne siies a ATO, | 3 20° | 3) 383% |

28 0 0 33 0; 42,0 0 |

2 er ee te Waar Ade ee S|

pS ic leeae lh Rt 33% abd 2 66%)

4% 14 % |

AN ie heed RO ORRAE fa8 37 3 on

| | 116° 0 | 33 0 0 8 0% }

Norrs.—Compiled from F. Finn’s ‘The World’s Birds.’

Honey-guides, parasitic; the rest, nesting-habits not mentioned.

Reference Nos. to middle column, the others are to be seen in Tables XIII. and

XIV. 1 open and closed, 2, 32, 86: 2 open and closed, 10, 15, 44: 4 open

and closed, 14,19, 34. Percentages to right of figures = horizontal per-

centages, percentages below = vertical percentages.

Further, it is noteworthy that Honey-guides (Indicatoride)

sometimes present sexual differences (Finn), and yet these birds

are parasitic. This exception is important because it shows that

the rearing of young as well as the open-nesting habits cannot

account—in this case, at any rate—for secondary sexual di-

morphism.

The Aphrodisiac Theory.—Those who believe in the necessity

for an aphrodisiac (bright feathers, sexual display, and song)

must assume coyness on the part of the female. Little, if any,

attempt, however, is made to establish this important sexual

difference ale and an impudent male. There does

not appear to be any veason why the females of birds especially

subject to the attack of enemies should be especially coy, or that

among birds not especially subject to attack the females should

not require a powerful stimulant.

The Catabolism Theory (Prof. Geddes and Thomson, ‘ Evolu-

tion of Sex,’ 1889).—The secondary sexual structures found in the

males are considered to be due to a katabolie constitution, and

their absence in the female due to an anabolic tendency.

No attempt is made to explain the distribution of secondary

sexual characters among birds on this theory, or to give a reason

SEXUAL CHARACTERS IN BIRDS. 677

why some birds present great extra-sexual difference and others

little or none. The conclusions arrived at in this paper do not

appear to be explicable on this theory; there seems to be no

reason, for instance, why birds especially subject to the attack of

enemies should be more katabolic than birds not especially so

subject.

The Mendelian Theorrces.—Although the followers of Mendel

have carried out very important investigations with regard to

secondary sexual characters, as far as I am aware, they have

made no attempt to account for the distribution of these charac-

ters in the animal kingdom.

The Hormone Theory (¢ Archiv fiir HEntwicklungsmechanik,’

1908), J. T. Cunningham.—-According to this theory, secondary

sexual characters become established in the following manner :—

Asa result of certain mechanical and functional stimuli, confined,

for instance, to the male, certain organs of the male take on new

characters: these are acquired characters. During their pro-

duction a specific hormone is produced, which stimulates the

corresponding determinant in the gametes and alters them in

such fashion that, during development, they reproduce the

acquired character ; but it also alters them in another direction——

it unplants in them an ability to reproduce the acquired character

only in the presence of the male sexual hormone. In the absence

of the male sexual hormone, the character cannot develop: for

instance, if a stag be castrated the antlers cannot develop on

account of the absence of the male hormone—the character

remains dormant. Obviously the material here dealt with

applies only to the first part of the theory, that dealing with

mechanical and functional stimuli.

In the case of birds the functional stimuli, which eall forth the

bright plumes of the males, are those associated with the display

of these ornaments during courtship, “the tail of the peacock

became brilliant and hypertrophied because it had been con-

stantly erected.” ‘The stimulus calling upon the male to display

seems, therefore, to be the deciding factor. Thus in order to

support this theory, it would be necessary to show that the males

of birds subject to enemies suffer more from such stimuli than

do those birds relatively not liable to be attacked.

Finally, it is necessary to point to the importance of the

correlation under consideration with respect to the possibility of

it being able to account for the presence of secondary sexual

characters.

The activities of animals fall under three headings ;—

(1) Those associated with the procuring of food.

(2) 5 - escape from enemies.

(3) i 95 reproduction.

Tt has usually been thought that the sexes bear the same

relation to the environment as regards the procuring of food and

the escape from enemies ; and therefore to account for secondary

678 ON SECONDARY SEXUAL CHARACTERS IN BIRDS.

sexual characters, search has always been made in the repro-

ductive environment. But the correlation between secondary

sexual dimorphism and vulnerability to enemies indicates that a

solution to the problem may possibly be found to be related to

the activities associated with the escape from enemies, and

especially if it could be shown that the sexes do not bear the

same relation to the environment as regards escape from enemies.

The author has attempted to show elsewhere (‘ Controlled

Natural Selection,’ 1914) the lines along which solution may

possibly lie.

Reference List of Birds.

1. American Vultures (Cathartide). 26. Jacamars (Galbulidz).

2. Auks (Alcide). 27. Kingfishers (Alcedinide),

3. Barbets (Capitonide). 28. Motmots (Momotide).

4. Bee-eaters (Meropide). 29. Mound-birds (Megapodiidze).

5. Bustards (Otidide). 30. Mowse-birds (Coliidee).

6. Cassowaries (Casuariide). 31. Nightjars (Caprimuleide).

7. Cormorants (Phalacrocoracide). 32. Owls (Strigidee).

8. Coursers (Glareolid). 33. Parrots (Psittacide).

9. Cranes (Gruidz). 34. Passerine Birds (Passeride),

_ 10. Cuckoos (Cuculide). 35. Penguins (Spheniscidze).

11. Curassows (Cracidz). 36. Petrels (Procellariide).

12. Birds of Prey (Falconidz). 37. Pigeons (Columbide).

13. Divers (Colymbide). 38. Putf-birds (Bueconide).

14. Ducks (Anatide). 39. Rails (Rallide).

15. Frogmouths (Podargide). 40. Rollers (Coraciidee).

16. Game-birds (Phasianidz). 41. Sand-Grouse (Petroclide).

17. Grebes (Podicipedide). 42. Shore-birds (Charadriide).

i8. Gulls (Laridz). 43. Storks (Ciconiide).

19. Hemipodes (Turnicide). 44. Swifts (Cypselidee).

20. Herons (Ardeide). 45. Thick-knees ((Hdicnemidz).

21. Honey-guides (Indicatorida). 46. Tinamous (Tinamidz).

22. Hoopoes (Upupide). 47. Toucans (Rhamphastide).

23. Hornbills (Bucerotide). 48. Touracous (Musophagide).

24. Humming-birds (Trechilidee). 49. Trogons (Trogonide).

25. Ibises (Ibidide). 50. Woodpeckers (Picide).

ON PATTERN-BLENDING. 679

49. Some Observations on Pattern-Blending with reference

to Obliterative Shading and Concealment of Outline.

By J. C. Morrram, M.B. (Lond.) *.

[Received June 25, 1915: Read November 9, 1915.]

(Text-figures 1-5.)

INDEX.

MorPHOLOGY : Pages

Patterns. Experimental analysis of ...................5. 679, 683, 687

Patterns. Arrangement of in Mammals, Birds, and Insects. 681, 686

ErHoLoey:

enuicrans., — (Cloraeenbbiner JOON? OP 355 sno ga0-sasutqnaas5 290 385 00 NN One 689

This paper endeavours to show that the patterns found on

animals may in many cases be of use in concealment after they

have become blended with distance, and that the patterns of

many animals are not intended to represent pictures of their

backgrounds, but are aids to concealment only after blending.

Distance at which Blending takes Place.

Tf a surface of black and white squares or of black and white

lines, or black and white spotted surfaces be examined from suc-

cessively increasing distances, a point will be reached where the

Text-figure 1.

800

700

es.

M7 117.

600

500

400

300

200

Blending distance

100

Side of square and small side of stripes in millimetres.

Oa hee ee

| 2 3 4 8 16

Diagram showing blending distances. Experimental conditions as in text-fig. 5, 7.

pattern can no longer be seen but is replaced by an even grey tone.

The distance at which this change takes place was found to depend

on the size of the black and white areas. The diagram (text-

fig. 1) indicates the distances at which blending takes place in

* Communicated by the SECRETARY.

680 DR. J. C. MOTTRAM ON

the case of alternate black and white squares and stripes under

the experimental conditions stated.

Further experiments showed that the blending distance of

different shapes depends upon the relative concentration of the

black and white areas, the more concentrated the greater the

blending distance. If, for instance, a checkered surface composed

of squares of 2 square millimetres be compared with a surface

composed of black and white oblongs of 1 multiplied by 4 mm. or

‘*) multiplied by 8 mm. or 025 by 16 mm., as regards the blending

distance, it will be found that the distance is greatest for the

squares and least for the narrowest oblongs. On comparing

squares with other figures in this respect, for instance, with circles

or triangles, it was found that the blending distance was related

to the concentration of the various shapes.

The Surface which kesults after Blending.

If a patterned surface of black and white be gradually

approached from a distance, the surface which at first appeared

of a perfectly even grey tone will, at a certain point, become gran-

ular, and a very short distance in front of this the pattern will

suddenly shine forth clear and defined.

By varying the relative proportion of black to white in the

pattern, all tones of grey can be produced when the pattern is

blended. It is thus possible to match a given grey surface by

means of black and white pattern, and also possible to reproduce

a graded surface of grey: text-fig. 3, 1, shows without any de-

scription how this can be done. If these copies of greys with

patterns be fixed to solid figures such as a cylinder and be then

examined, it will be found that after blending, the copies are

undisturbed. It follows that the obliterative shading so com-

monly seen on animals and consisting of low tones where the

light strikes, high tones in the shadows, and intermediate tones

between the two, could be reproduced by means of pattern.

Search has therefore been made amongst animals to discover

whether this method exists. ‘

Some Hxamples of Animals which appear to show Obliterative

Shading by means of Pattern- Blending.

Grant's Zebia (Hquus burchelli granti).— This animal’s coat is

coloured black and white, black stripes on a white background :

the background, unlike that of the vast majority of animals, pre-

sents no obliterative shading, it has the same tone throughout ;

the stripes are somewhat darker on the back than they are below.

On veferring to text-fig. 2. 3, it can be seen that on the dorsal

surface of the animal and on those parts of the body where the

hight strikes, the stripes are broad, whereas on those parts which

under natural conditions would be in shadow, the stripes are

narrow : further, on those areas which would be more or less

PATTERN-BLENDING. 681

evenly lit, as for instance the vertical surfaces of the legs and

sides of the face, the stripes maintain a more or less constant

width. From what has been observed under experimental

conditions as regards pattern-blending, it is obvious that, if one

cousiders only the trunk of this animal, the varying widths of

the stripes after blending must produce a condition of grey

similar exactly to obliterative shading as seen, for instance, in

the ass.

Text-figure 2.

2 The same view : ae ne sous chief Per under top lighting,

3. Lateral view of same animal.

4. Hind view of base of tail, showing pattern.

}. Wind view of base of tail, pattern omitted, showing shading as in 2.

6. Hind view of Tiger (Helis tigris), showing pattern.

7. Front view of Grant’s Zebra, showing pattern.

Amongst big-game hunters there is a difference of opinion as

to ieee this animal is easily seen or not, under natural con-

ditions. It appears that this difference of opinion is largely due

to the district in which the hunter has observed these animals.

In some districts the zebra is relatively tame and the hunter

can frequently approach to within the blending distance of the

pattern, which has been variously estimated, according to the

lighting : under these conditions the zebra will of course appear

a very conspicuous animal. On the other hand, in those districts

where near approach cannot be gained on account of the open

nature of the country or wildness of the animal, the hunter will

682 DR. J. C. MOLTRAM ON

look upon the zebra as one of the most difficult wild animals to

pick up.

Of the zebras, #. burchelli presents between the stripes con-

siderable obliterative shading, as well as faint stripes of buff.

H. grevyi also has some obliterative shading, H. zebra very little,

and H#. burchelli granti none at all. Complete absence is, however,

not uncommonly seen in all the species.

Thayer, in ‘ Concealing Coloration in the Animal Kingdom,’

looks upon the striped coat of this animal as an obliterative

picture-pattern of rank grasses and tree-stems: it is noteworthy

that he makes no mention of the absence of obliterative shading,

which he elsewhere insists always forms the canvas for picture-

painting. If, however, examination be made of the pattern to

be seen on viewing a zebra from behind (see text-fig. 2, 7.), it is

at once evident that this does not represent a picture-painting of

reeds, herbage or trees. On the other hand, it illustrates in a

remarkable way obliterative shading by means of pattern. Text-

fig. 2, 2, is a modelled drawing of the same view with the stripes

omitted, the lighting being from above, and shows the major

shadows which are cast. On comparing these two figures it can

be seen that in the high lights, the stripes are broad, and that as

one passes into the shadows, they fade away: where the high

hight strikes the hocks the bands are broad, whereas below the

hocks where the leg is in the shadow, the stripes are narrow.

The striping on the tail even falls into line, as shown in text-

YR, 2 AB, Bs

The Guineafowl (Vumida meleagris).—This bird illustrates

obliterative shading by means of white spots on an even blue-grey

background (text-fig. 3, 7, bottom right-hand square). On the

back the white spots are small, but become larger as one

passes to the ventral aspect, where they are four times the

diameter of those on the back. In some species the wings are

covered with a series of short white bars, narrow on the back

and broader below, as the wing lies covering the body during

rest. These markings on the bird blend at a comparatively

short distance; within the blending distance the bird appears

conspicuous, beyond, difficult to see. The bird lives in open

country, and there does not appear to be anything in its surround-

ings which this pattern emulates.

The Cheetah (Cynelurus jubatus).—-The pattern, as shown in

text-fig. 3, 7, bottom left-hand square, consists of black spots on

a pale fawn background: the background presents either no

obliterative shading or very slight; the spots are close together

on the back, gradually becoming more distant towards the ven-

tral surface, which is hidden by a fringe of long hairs projecting

downwards from the flank.

The Serval (Felis serval) presents a similar pattern to the

Cheetah except that in the dorsal region, spots are replaced by

short bars. This kind of pattern is commonly seen in the

Civets.

PATTERN-BLENDING. 683

The Jaguar (felis oncw).—Here the pattern is laid on a back-

ground presenting very little obliterative shading : on the back

are closely placed black spots ; as the ventral surface is approached

the spots are seen to contain a central light brown area which

gradually increases In size, whilst the surrounding dark ring

diminishes and on the ventral surface becomes broken up into a

number of separate spots irregularly arranged around the central

area. This central area is always somewhat darker than the

background and usually contains a small central black spot.

Viewed as a whole this rather complicated pattern (see text-fig.

3, 17) shows a decreasing quantity of black from above, down-

wards, and when blended at distance will produce a graded tone

similar to obliterative shading. A similar pattern is found in the

Ocelot (7. pardalis) and in several varieties of the Leopard, where

it is often associated with an entire absence of obliterative

shading.

These patterns found in the genus Melis have been considered

to be picture-paintings of a checkered background, for instance,

leaf-shadows on bare ground; but the extreme regularity of these

patterns is so unlike the very irregular nature of such backgrounds

and so unlike the patterns which undoubtedly do simulate these

backgrounds, that this deduction does not appear to be well

founded ; especially as these patterns blend at a comparatively

short distance, probably within the charging length of the cat:

and lastly, several of these animals inhabit open country.

However, apart from this consideration, the facts remain that

these patterns will produce after blending obliterative shading,

which is otherwise not presented ; and these patterns must in this

respect be powerful aids in concealment.

Sufficient examples have now been given to illustrate that this

method of concealment is by no means uncommonly utilised.

Black and white has been chiefly dealt with, but any colour can

be produced by the blending of patterns consisting of two or

more colours: examples of this are not difficult to find, for

instance brown is often produced by the blending of buff and

black.

Experiments were carried out to discover whether this method

of coloration had any advantage over an unpatterned one. Ifa

grey disc be examined in front of a series of backgrounds varying

from black to white, a particular background will be found against

which the grey disc is invisible, this being of course that back-

ground which is of an exactly similar tone to the disc: against

the other backgrounds the dise will be visible at varying dis-

tances. If the results so obtained be compared with those obtained

when a disc of black and white squares, which after blending

produce the same tone of grey as the original disc, is substituted,

itis found that the checkered disc possesses no advantage as

regards the distances at which it can be seen, over the plain disc.

There is, however, an exception : against the background on which

the plain dise is invisible, the checkered dise is of course visible

684 DR. J. C. MOTTRAM ON

within blending distance, and against backgrounds closely similar

to this background the checkered disc is at a disadvantage as

regards invisibility. Over a long series of experiments there was

some indication that against other backgrounds the checkered dise

was slightly less visible, the measure being the greatest distance

at which the discs could be seen; but although great care was

taken with the constants of the experiment, the differences were

too small and the experimental error too large, to justify a definite

conclusion, particularly as the appearance of the checkered disc

after blending was remarkably different from that of the plain

disc. When the plain dise was visible, its outline was seen to be

sharp, so that its shape could easily be recognised, but with the

Text-figure 3.

TBE

Examples of obliterative shading by means of pattern-blending.

checkered disc the outline was remarkably indistinct, so that

it was frequently impossible to tell whether a square, circle

or equilateral triangle had been placed against the background.

Experiments were carried out in order to determine the factors

controlling this obscuring of form. Four squares were prepared

as shown in text-fig. 3, 2-5: as can be seen, these squares consist

ot black and white areas of equal proportions. In fig. 3 the areas

are interrupted along the two opposite margins, in fig. 4 along

the two adjacent margins, in fig. 6 along all four edges, and in

fig. 2 there is no interruption at the margins. These squares.

were examined under different conditions of lighting and against

PATTERN-BLENDING. 685

different toned backerounds, and it was found that on all occa-

sions marginal indistinctness after pattern-blending occurred

where the black and white areas were interrupted at the edges ;

for instance, fig. 2 appeared as a grey square whilst fig. 5 showed

an ill-defined grey area fading into the background. It was

noted that when the background was near in tone to the squares

atter blending of the pattern, this obscuring effect was most

marked: when the background was far removed in tone, for

instance was black, or white, very little obscuring was noticeable.

Other experiments showed that the greater the width of the

black and white areas passing out at the margin, the greater the

obscuring effect after pattern-blending.

One is now in a position to conclude that the method of

obliterative shading by pattern-blending, as seen for instance in

the Zebra, if it does not possess an advantage over the usual

method as regards visibility, as measured by the distance at which

the object is visible, nevertheless would seem to have a decided

advantage in that the contour or silhouette of the animal against

backgrounds approaching it in similarity, is made blurred Saidne

than sharp. It is noteworthy that the stripes of the Zebra

everywhere pass out at the margins (see text-fig. 2, 7, 3, and 7).

Referring to the possibility of the Zebra’s stripes being

pictures of reeds in high light and shadow, as has been suggested

by Thayer, it is noteworthy that the black stripes decrease in

breadth from above, down, which is contrary to expectation and

to what actually is found among other animals. Thayer has

pointed out that the markings on the backs of animals are

usually smaller than those on the ventral aspect, because they

represent objects in the landscape more distant: they depict the

foreshortening of the ground. If reeds are painted on the zebra’s

back, one w ould at least have expected the stripes to be of equal

ria.

The Tiger presents an excellent example of reed-painting: on a

coat showing well-marked obliterative shading, dark stripes are

to be seen, they are irregularly distributed, they for the most

part increase in width from above, down, and on the belly

they terminate in a large dark mass which, cumouslyy enough,

often sae a small white centre (compare text- fig. 2, 1 with text-

fig. 2

serie of outline-masking by means of pattern-blending at

the margins are very commonly seen amongst Lepidoptera, but

before considering instances, some experiments must be referred

to. If one takes two flat surfaces, one an even dark grey and

the other an even light grey, and brings them in apposition as

shown in text-fig. 3, 6, the junction will appear sharp, ne matter

how viewed. Jf now, at the junction a narrow band of black

and white squares, which after blending will have the same tone

as the light square, be interposed, see text-fig. 3, 6, it will be

found that the junction no longer remains sharp ; : if the outer

square be replaced by areas of different tones varying from light

Proc. Zoou. Soc.—1915, No. XLVI. 47

686 DR. J. C. MOTTRAM ON

grey to very dark grey, the same result will be seen. It is easy

to see that, supposing the central square was an insect and that

the different tones placed outside were the various backgrounds

against which it would be likely to be seen, then the possession

of a checkered margin would be a great aid towards its con-

cealment.

‘Yo further test this property of an interrupted margin, squares

were made as shown in text-fig. 3, 7-1/0: these squares were

made so that after pattern-blendimg they all appeared of the

same tone. They were examined against a series of backgrounds

and it was found that, whereas 70 remained for the most part

sharp, the others showed blurred outlines; the blurring effect in

the case of square 7 was not appreciably different from that of

Sand 9. It appears therefore that a comparatively narrow

broken margin is effective.

Text-figure 4.

ait

Examples of marginal patterns: F=fringe of scales projecting from wing-margin.

1, Dingy Skipper, Hesperia tayes 2, dorsal surface of anterior wing.

2. Grizzled Skipper, H. malve 2, dorsal surface of anterior wing: pattern con-

fined to fringe.

3, Common Blue, Lycena icarus 3, ventral surface of anterior wing: pattern of

marginal eye-spots.

4. Chalk-hill Blue, L. corydon 9, dorsal surface of anterior wing: pattern chiefly

‘ confined to fringe.

5. Painted Lady, Pyrameis cardui, dorsal surface of posterior wing.

6, Long-tailed Blue, L. betica, ventral surface of posterior wing: pattern at some

distance from margin.

A few examples of this mode of concealment are given in

text-fig. 4: it may be mentioned that outline-masking in insects is

PATTERN-BLENDING. 687

also brought about by scalloping of the margin and by a fringing

of projecting scales ; other methods are utilised, but these two are

specially mentioned as they enter the chosen examples.

Examination of the wings of insects showed that this marginal

pattern is often a short distance from the extreme edge; experi-

ments were therefore carried out to discover at what distance

the marginal pattern would be effective. A grey square (text-

fig. 3,6) was examined against varying backgrounds with the

checkered margins (equal in tone after blending) placed at

varying distances opposite free margins, as shown in the figure.

Jt was found that the marginal blurring, produced by the

checkered patterns, was effective when they were placed a very

short distance from the margin. Up to the width of the squares

used some effect was noticeable, but was much more marked

when the distance was produced to one half or a quarter of this.

The single row of squares appeared to be almost as effective as

the double ; attempts were made to obtain more definite results,

but in the absence of some method for estimating the amount of

blurring, this was found to be impossible. Text-fig. 4, 6, is an

example in which the marginal pattern is separated a short

distance from the free edge.

As eye-spots are frequently to be seen near the margins of the

wings in Lepidoptera, it was thought that they might play some

part in concealment of form by means of outline-blurring. A

few experiments were carried out to discover whether this be

so or not. In the centre of a small grey area, an eye-spot was

placed consisting of a black centre surrounded by a white ring,

the proportion of black to white being so arranged that, after

blending, they were equal in tone to the surrounding grey; a

similar grey area was prepared but with no central eye-spot :

these two squares were then examined against various back-

grounds; at the same time the squares were gradually reduced

in size: it was then found that when the square (text-fig. 3, 77)

was reduced to 12°75 mm.,, the edges began to be blurred as

compared with the edges of text-fig. 3, 72, especially against

backgrounds not widely different in tone. Further reduction of

the square to 10°3 mm. heightened the difference.

In this particular experiment the size of the eye-spot was:

total diameter 6°75 mm., black centre diameter 4 mm. HKye-

spots of different diameters were also used, and it was found that

the larger the spot, the greater the distance at which mar-

ginal blurring was produced. Some experiments were also

made, in which the eye- spots were drawn so that after blending

they were different in tone from the surrounding grey. It was

found that they were not very effective in margin-obscuring

unless the background was closely similar to their tone, after

blending : they appeared as either light or dark areas on the grey

square. An experiment was next devised to discover whether

this outline-blending, by means of an eye-spot, was more effective

than other patterns. As shown in text-fig. 3, 73, a grey square

47*

688 DR. J. C. MOLTRAM ON

was prepared, in one corner of which a square eye-spot was

placed, in another corner a square of the same size and containing

the same proportiom of black and white but distributed on either

side of the diagonal, in a third angle a similar square divided

down the middle, the fourth angle was left unoccupied. This

square was then examined against various backgrounds. Two

facts were recorded : first, the eye-spot blended at a much shorter

distance than the other two black and white squares, and secondly,

the eye-spot square more effectively blurred the angle of the

grey square than did the other two patterns. It was, however,

difficult to decide how much differenee there really was, because

at the distance at which the eye-spot was blended and producing

blending at the corner of the square, the other two patterns had

not blended. In order to show the differences in the blending

distance : under the conditions of the experiment, the blending

distances of these patterns (text-fig. 3, 74-76) were as follows:

eye-spot 184 feet, diagonally divided square 323 feet, medially

divided square 35 feet. Kye-spots distributed over a surface are

often used for producing a pattern; in Lepidoptera they often

form marginal patterns, but more frequently form patterns some

distance from the edge, as is also the case in the Jaguar (see

text-fig. 5, 17).

It appears that a given quantity of black and white, used as

circular eye-spots, forms a pattern which obscures the outline, if

not better, at any rate as well as other patterns. In order to

be effective in this respect the eye-spot, after blending, should

not make a contrast in tone with its ground; this was found to

be the case as regards the eye-spots on the under wings of a

number of butterflies examined: for instance; when the ground-

work was light, the dark m the eye-spot was either small or not

deep in tone.

Other possible uses for eye-spots have been put forward : they

have been thought to be attraction marks which induce the

attacking enemy to strike them rather than vulnerable parts ;

they have been looked upon as “dazzlers” to divert the enemy's

eye from the inseet as a whole. If one examines the larger eye-

spots and those on the dorsal surface of the wings of many

butterflies, one finds that they do not, after blending, harmonise

with the rest of the wing; thus it would appear that they pro-

bably have a different function from those found on the under

wings, which it has been thought are for concealment.

Patterns have been considered as regards their powers of con-

cealing form, against plain backgrounds only. Against mottled

backgrounds, which probably are more common in nature, these

powers of concealment ave as effective, not because they may

copy or be paintings of anything in the background (they may

be), but because after blending they will cause the animal to fade

into its surroundings.

Beyond the blending distance of @ pattern, the question as to

PATTERN-BLEN DING. 689

whether the pattern does or does not imitate some part of the

animal’s surroundings, does not come in: pattern can only be

effective in concealing form by means of mimicry or protective

coloration, within its blending distance; nevertheless, beyond

blending distance, as has been seen, pattern still has powers of

concealing form. It is possible that a given pattern may com-

bine both these uses. Suppose AE (text-fig. 5, 3) be the extreme

Text-figure 5.

ty Ground plan of apparatus for experiments with artificial patterns.

2. The same, modified for measuring the blending distance of the patterns of

insects’ wings.

Notes referring to 1 and 2.

B=backgrounds made of ‘ Velox’ paper exposed for varying lengths of time.

C & C’=standard candles.

S & 8’=screens of “ Bristol” board painted black.

obj.=object to be examined.

G=glass plate on which is glued a small bead of cork to which the insect

is pinned,

obs.=observer along the dotted line.

Using the following distances, an illumination is obtained not unlike that under

a hedge on a dull day :—

C-S and C’=S'= 6 cm.

C-obj. and C—obj.=40 cm.

C-C'=40 cm.

G-B=20 cm.

distance at which animal A is visible, and AF he the distance at.

which the pattern blends, then within the circle FGH the animal

690 DR. J. C. MOTTRAM ON

will be protected from enemies by mimicking its surroundings,

and within the ring defined by FGH and EJ by means of pro-

tective pattern-blending. The relative importance of these two

methods of concealment will be according to the likelihood of

enemies finding their way into the two areas, and the chance

will be as the relative size of the two areas: the central area

will, however, be at a somewhat greater disadvantage, because

the enemy in arriving there, must pass through the outer area,

and in so doing may discover its prey before reaching the centre.

The relative importance will also depend upon the eyesight of the

enemy and the distance at which it commonly looks for its prey ;

those who have watched birds feeding upon insects have often

remarked the long distances at which they can see their quarry.

In order to illustrate the distance at which the patterns of

insects blend, measurements were made in the case of sixteen

insects, as shown in the following table.

It is obvious that the size of an animal must affect the size of

its pattern: for instance, a pattern which blends at a few feet

would be useless to an animal the size of the zebra. It appears,

therefore, that it is necessary, whilst giving the blending distance

of a pattern, to also state the size of the animal, and the relation

of these two in the form of a ratio as is done in the table.

Whether by comparing these pattern-blending ratios or indices,

it is possible to separate animals presenting conspicuous patterns

from those presenting inconspicuous ones, is beyond the scope of

this paper. In the case of the insects examined, in some cases

the dorsal surfaces of the wings have a high index, whereas the

ventral aspects of the lower wings have alll low imifeas, with the

exception of the Queen of Spain Fritillary (4. lathonia). In

this insect the high figure is probably due to the reflection of the

candle-light from the ‘‘ mirror” spots on the wing., It has been

noted that the larger the pattern, the more effective the outline

blending ; on the other hand, the greater the blending distance of

the pattern, probably the more conspicuous the animal: other

things being equal, it follows, therefore, that a pattern may be so

adjusted that the danger of it, Short of blending, is counter-

balanced by its concealing powers after blending. Finally,

attention must be directed to one other aspect of these experi-

ments. Deductions have been drawn from experiments carried

out with the human eye: thus it is entirely a matter of opinion

whether they would apply in the case of the eyes of animals.

It is known that many animals are short-sighted compared with

man, for instance, those of short stature and which for this

reason have a near horizon: it may be that the lion at night

cannot see the zebra’s stripes until within close range. On the

other hand, there is some evidence that in the case of certain

birds, such as falcons, vision is more piercing than in man.

However, these deductions from experiments with the human

eye are illustrated in the markings of animals’ coats.

691

PATTERN-BLENDING.

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Conclusions.

(1) Obliterative shading in animals is sometimes effected by

means of pattern-blending.

(2) The outlines of animals are frequently masked by the

blending of patterns at or near their margins.

(3) Patterns having these effeets are usually unlike the

animal’s surroundings and therefore cannot be of use in conceal-

ment by means of mimicry (using mimicry in the broadest sense

of the word),

ON LAND-PLANARIANS FROM WEST AUSTRALIA. 693

50. On some Land-Planarians collected in Western Australia

and Tasmania by Members of the British Association

for the Advancement of Science. By Arraur DENDY,

D.Se., F.R.S., F.Z.8., Professor of Zoology in the

University of London (King’s College).

[Received November 3, 1915: Read November 23, 1915.]

INDEX.

Page

GEOGRAPHICAL AMEE EN nena) aan oe eee ta te Mei ad Nyse OOS

SYSTEMATIC :

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(Ch IOUS DOCG! B05 THs sodoode: canneoacounnyaacsadcesoosqoscaneseds OLE

GSicomitacisas putts wey Mon oka eine eases ahh vas gateenneen se eb ore MOG

(GEOL IA OUISONS Sealand eae seehae cc eoemhocssuaosesecmen se uninGanereBes, OL

GFA INTES DEMO ec AH dee Se aie aie ree AOL

During the visit of the British Association to Australia in

1914, exceptional facilities were afforded to those members

who visited Western Australia and Tasmania for the collection

of ‘zoological material for future investigation. In Western

Australia various zoological expeditions were most successfully

organized by Professor W. J. Dakin, and in Tasmania similar

excursions were admirably arranged and carried out by Pro-

fessor Flynn. Some of these expeditions were devoted to

the collection of marine animals, while on others the members

who took part in them had ample opportunities for the inves-

tigation of the terrestrial invertebrates.

During my residence in Australia, many years ago, I had

already paid particular attention to a section of the terrestrial

fauna which I [1895 bis] termed ‘“ Cryptozoic,” comprising those

small animals, for the most part invertebrates, which habitually

hide away beneath logs and stones, or under the bark of trees.

This faunistic assemblage includes many lizards, frogs and toads,

centipedes and millipedes, scorpions, spiders, Peripatus, insects

of many kinds—especially cockroaches—many slugs and snails,

a large number of land-planarians, and, more rarely, land-

nemertines.

Of the Land-Planarians, the species of which are more or

less readily distinguished by their beautiful colour-markings,

I have on previous occasions described many different kinds

from Australia, Tasmania and New Zealand, while many

others have been described by Messrs. Fletcher and Hamilton,

Professor Baldwin Spencer and Mr. Thomas Steel. It was

naturally, therefore, a great pleasure to have the opportunity

of returning once more to my old pursuits and collecting again

species that were long ago more or less familiar to me. In

addition to these, however, several hitherto undescribed forms

694 PROF. ARTHUR DENDY ON

were met with, so that altogether we obtained three species in

Western Australia and six in Tasmania.

From Western Australia only four species have hitherto been

described, all by Mr. Steel [1901], and all three obtained by us

appear to be new.

From Tasmania, Steel [1901 dcs] gives a list of twelve species,

six of which were originally described by myself. Of these twelve,

the British Association party obtained three, together with one

previously known only from Australia and two new ones.

By far the commonest species in Tasmania seems to be

Geoplana tasmaniand, originally collected on the historic voyage

of the ‘ Beagle’ and described by Charles Darwin.

Tt is curious that so few Australian zoologists have con-

cerned themselves with the study of the Land-Planarians.

This is the more to be regretted inasmuch as the opportunities

for collecting these animals are rapidly passing away with the

clearing of the bush. Moreover, much remains to be done in

the investigation of these and other Cryptozoic animals. The

Land-Planarians, in particular, still demand thorough compara-

tive anatomical investigation with a view to revising the generic

classification. Thus von Graff [1899] has proposed the genus

Artioposthia, based on material supplied by myself, to include

certain forms with a remarkably complex copulatory apparatus,

but until many more species have been anatomically investigated

it is difficult to estimate the value of this suggestion and impos-

sible to say in many cases to which genus a given species should

be referred. For this reason I retain the generic name Geoplana

in its older and wider sense.

I desive to express my great indebtedness, not only to our

generous Australian and Tasmanian hosts, but also to those

members of the British Association who were good enough to

hand over to me the Land-Planarians which they collected.

A. Species collected in Western Australia.

GEOPLANA DAKINI Sp. 0.

When crawling, very long and narrow, as much as three

or four inches in length; strongly convex dorsally, flattened

ventrally ; without longitudinal ridges. A well-grown specimen

in spirit measures about 74 mm. in length by 3 mm. in width in

the middle, and is approximately oval in transverse section.

The eyes are arranged as usual. The peripharyngeal aperture

is situated about the middle of. the body; the pharynx when

protruded in spirit is subcylindrical. I have been unable to

make out the genital aperture.

In life the dorsal surface is pale yellow, mottled and striped

with olive-brown ; the ventral surface is white, without pattern,

and the anterior tip 1s pink.

The colour-markings on the dorsal surface are typically

LAND-PLANARIANS FROM WEST AUSTRALIA. 695

arranged as follows :—There is a narrow median stripe of brown

and on each side of it a band of clear yellow ground-colour of

about twice the width. This is followed by a band of brown,

less sharply defined than the median stripe and of about twice

the width. We may call this the inner paired stripe; it is

followed by a somewhat wider band of ground-colour mottled

with minute brown specks, and then comes an outer paired stripe

similar to the inner one. In life there is a fairly wide band of

clear ground-colour visible on the dorsal surface outside each

outer paired stripe, but in spirit this tends to be turned in by

contraction, so that the outer paired stripe comes to he at the

margin of the dorsal surface. The degree of distinctness of

the paired stripes varies considerably, and they sometimes tend

to merge into the mottling that lies between them.

There are thus five narrow longitudinal dark stripes on the

dorsal surface, and in this respect the species resembles the

common G. quinquelineata of Kastern Australia, to which it is

probably clesely related. It differs from that species, however,

in that the dark stripes are not placed at equal distances apart,

and in this respect it agrees with G. mediolineata Dendy,

var. simularis Steel [1901], from South and Western Australia.

It differs from both these species, however, in the presence of

the mottling between the inner and outer lateral stripes.

This species was common in the neighbourhoed of the

Mundaring Weir, near Perth, W.A., upwards of a dozen

specimens having been collected by our party. I have much

pleasure in naming it after my friend Professor W. J. Dakin,

D.Sec., who did so much to bring about the striking success of

our zoological expeditions in Western Australia.

GEOPLANA FLAVILINEATA Sp. Nn.

When crawling, the dorsal surface is convex, the ventral flat:

there are no longitudinal ridges. In spirit the ventro-lateral

margins are rather prominent and the ventral surface may be

concave. The larger of the two specimens in spirit measures

40 mm. in length by 3°5 mm. in width in the middle. The eyes

are numerous, but it is difficult to make out their arrangement

in spirit specimens. The peripharyngeal aperture is situated

somewhat in front of the middle of the ventral surface, the

genital a little nearer to the peripharyngeal than to the posterior

extremity of the body.

In life the dorsal surface is dark olive-grey, or purplish, with

five very narrow longitudinal stripes of pale yellow, not at all

conspicuous. The ventral surface is yellowish white, without

markings ; and the horseshoe-shaped anterior tip is pink.

The colour-markings on the dorsal surface are seen, under a

lens, to be arranged as follows in the most strongly marked

specimen :—The very narrow median yellow stripe is edged on

either side by a very narrow dark grey band ; this is followed by

a, very narrow yellow stripe (the inner paired stripe), narrower

696 PROF. ARTHUR DENDY ON

and less distinct than the median stripe. Then comes a much

broader band of finely mottled grey, bounded on the outside by

another very narrow yellow stripe (the outer paired stripe). The

outer paired stripe is much more distinct than the inner one and

is accentuated by the presence of a very narrow, almost black

edging to the broad band of dark grey, finely mottled ground-

colour which lies outside it and extends to the ventro-lateral

margin of the body.

The-characteristic feature of the pattern seems to be the

presence of five very narrow, pale yellow stripes on a dark

background of finely mottled grey, the interval between the

median stripe and the inner paired stripe being considerably

less than that between the two paired stripes. In the specimen

described above the inner paired stripe is almost obsolete, while

the outer one is quite clearly defined; in the other specimen both

paired stripes are equally strongly developed, but neither of them

is nearly so distinct as the median stripe, and the outer one has

only a feebly developed dark edging on the outside.

This species seems to be closely related to Steel’s Geoplana

fusco-dorsalis [1901], also found near Perth.

I have only two specimens, both of which were obtained 1

the neighbourhood of the Mundaring Weir.

GEOPLANA COMITATIS sp. 0.

Body in life comparatively short and broad, Dorsal surface in

life pale dusky yellow, with olive-brown stripe and mottling ;

ventral surface white or grey, without markings; anterior tip

pink.

The arrangement of the pattern on the dorsal surface is as

follows :—There is a narrow median dark stripe. The interval

between this and the margin of the body is divided into three

bands of about equal width, viz., an inner band of clear ground-

colour, an outer band of clear ground-colour, and an ill-defined

intermediate zone in which the ground-colour is thickly sprinkled

with minute dark specks.

The eyes are arranged as usual. The peripharyngeal aperture

is situated about the middle of the body and the genital aperture

somewhat nearer to it than to the posterior end. There is,

however, a good deal of variation in this respect, depending upon

the state of contraction.

The body in spirit is rather short and thick, averaging about

25 mm. in length by 4 mm. in width in the middle; it is approxi-

mately oval in transverse section, with no trace of marginal

ridges.

This species bears a strong resemblance to Geoplana tasmaniana

Darwin, but may be distinguished by the following features.

Although there is one pair of ill-defined, broad dorsal bands of

brown mottling, very much as in G. tasmaniana, there never

appears to be a marginal (or submarginal) mottling or stripe.

The general colour of the dorsal surface in spirit specimens is

LAND-PLANARIANS FROM WEST AUSTRALIA. 697

much yellower than in G. tasmaniana. The body (in spirit)

is thicker, less flattened dorsally, and much more rounded at the

margins. Steel [1901 bis} has described a supposed variety of

Geoplana tasmaniana which he calls flavicincta, which seems to

come a good deal nearer to G. comitatis than the typical form.

It seems to me hardly likely, however, that the West Australian

species is really identical with the Tasmanian “ variety.”

J have eleven specimens of this species, all collected in the

neighbourhood of Mundaring Weir. The specific name comitatis

is given m allusion to the British Association.

B. Species collected in Tasmania.

XHOPLANA TASMANIANA (Darwin) *.

Planaria tasmaniana Darwin [1844].

Geoplana tasmaniana Fletcher & Hamilton [1887 ].

Geoplana tasmaniana Dendy [1893 |.

Geoplana balfourt von Graft [1899 },

Geoplana tasmaniana Steel [1901 bis}.

This certainly seems to be by far the commonest of the

Tasmanian Land-Planarians, and it has been obtained from many

widely separated localities. In September 1914 Dr. Nicholls

collected a number of specimens on Maria Island, and it was also

much the commonest species in the neighbourhood of the Great

Lake, which some of us subsequently visited.

The following notes were made on living specimens fiom

Maria Island:—‘ Kyes as usual, When crawling long and

‘“narrow, but dorsal surface somewhat flattened as well as

“ventral. Dorsal surface very pale yellow with very narrow

“median stripe of dark grey or brown and two wider stripes of

“the same colour but less intense. Ground-colour between

“median and wide stripes minutely flecked with brown, also

“fine marginal mottling of brown separated from wide dorsal

‘* stripe by band of clear yellowish ground: colour, Colour of

stripes varies from olive-brown to reddish.’

The marginal band of mottling may develop into a more or

Jess distinct stripe and in spirit it becomes submarginal, visible

from the ventral but not from the dorsal surface, so that there

appear to be only three stripes on the dorsal surface (one narrow

and two broad) and a narrow yellow margin.

Specimens when crawling measure about 45 by 3 mm. The

largest specimens in spirif measure about 34 by 3°5 mm.

The shape of the body in transverse section, in spirit-specimens,

is somewhat flattened both above and balets and with slightly

pronounced marginal ridges.

A colour variety, repr eceated by two specimens from the neigh.

bourhood of the Great Lake, is characterized by the intensification

[The parentheses around the names of authors placed after scientific names in

Fe ‘paper are used in accordance with Article 23 of the International Rules of

Nomenclature (Proc. 7th Int. Cong. Boston, 1907, p. 44 (1912)),—Epit0r._

698 PROF. ARTHUR DENDY ON

of the dorsal stripes, the darkening of the ground-colour between

the median and paired stripes, and the disappearance of the

marginal mottling. The following notes were made on these

specimens during life :—‘‘ Dorsal surface dark brown, nearly

“black, with narrow median black line and a broader supra-

‘marginal black band, separated from the median stripe {line |

“by an intervening zone of dark brown. All these dark parts

‘appear at first sight as one very broad dark brown band, with a

“narrow white margin on either side.” In spirit the margins

of. the body ‘are very pale brown, not white lke the ventral

surface.

In another slight variety the intensification of the finely

mottled ground-colour of the dorsal surface to form a pair of

broad dorsal stripes does not occur.

Mr. Steel [1901 dis] has already pointed out that Professor

von Graff [1899| was probably mistaken in refusing to accept

my identification of this species with Darwin's Planaria tas-

maniana and in proposing for it the new name G'eoplana

balfourt.

GEOPLANA QUINQUELINEATA Fletcher & Hamilton.

Geoplana quinquelineata Fletcher & Hamilton [1887].

Geoplana quinquelineata Dendy [1890, 1891, 1895, 1896].

(reoplana quinquelineata von Graft [1899 |}.

Geoplana quinquelineata Steel {1901}.

More than a dozen specimens were obtained from the neigh-

bourhood of the Great Lake which seem to belong to this

common and widely distributed Australian species. I at first

considered them as representing a variety of Geoplana tasman-

iana, in which the broad, inner, paired stripe on each side was

broken up into two and the outer (marginal or submarginal)

stripe obsolete, giving, together with the median line, five narrow

longitudinal stripes, all on the true dorsal surface. The fact,

however, that the animal (in spirit) is a good deal narrower, and

at the same time less flattened, than G. tasmanana has induced

me to abandon this interpretation.

The specimens do not strike meas being quite typical examples

of G. quinquelineata, for the inner paired stripe is about twice

the width of either the median stripe or the outer paired stripe,

but it is well known that G. quinquelineata is subject to a good

deal of variation as regards the width of the stripes.

My notes on the living worm are very brief, as follows :—

“Dorsal surface yellowish, with dark grey or brown stripes.

“ Ventral surface pale grey, without markings. Anterior tip

“brown. Microscopic bluish or whitish specks appear under lens

‘on dorsal surface (?always). Eyes as usual.”

In spirit the ground-colour of the continuous dorsal and lateral

surfaces 1s light. brown, and the lateral surfaces appear on either

side of the narrow, white, creeping sole as a fairly broad band of

LAND-PLANARIANS FROM WEST AUSTRALIA, 699

this colour. All the dark stripes are visible from the dorsal

surface only, the outer paired stripe being supramarginal. The

body in section is oval, but somewhat flattened below. The

largest specimen now measures 35 mm. in length by barely 3 mm.

in greatest breadth.

This species has not previously been recorded from Tasmania.

GEOPLANA NICHOLLSI sp. nh.

? Geoplana diemenensis, pars, Dendy [1894].

¢ Artioposthia dicmenensis, pars, von Graft | 1899].

? Geoplana diemenensis Steel [1901 bis].

This species was obtained on Maria Island by Dr. G. E.

Nicholls and again at the Great Lake. The Maria Island

specimens may be regarded as the types of the species. ‘he

Great Lake specimens differ but slightly.

The Maria Island specimens, in life, were considerably

flattened even when crawling, but slightly convex above. Body

tapering gradually in front to narrow, horseshoe-shaped tip.

Posterior end bluntly pointed. Eyes as usual. Dorsal surface

pale grey, finely mottled with dark brown, and with fine median

dark brown stripe. Ventral surface pale grey, nearly white,

without markings. Anterior tip ight brown on both surfaces,

or pinkish. A specimen, when crawling, measured 50 mm. in

length by 4°5mm.in greatest breadth (not far from the posterior

end). When put in spirit the lateral margins turn in and

become visible from the ventral surface as two narrow, mottled

bands, one on either side of the broad creeping sole, forming two

narrow lateral surfaces sloping downwards and inwards from the

much flattened dorsal surface, which they join at an acute angle.

In spirit the peripharyngeal aperture is situated not far from

the middle of the body and the genital aperture rather nearer to

it than to the posterior end. -

In the spirit-specimens the mottlings have a distinct purple

tinge and show a slight tendency to arrange themselves in six

longitudinal bands on the true dorsal surface, three on each side

of the narrow median dark stripe, the outer band on each side

being close to the dorso-lateral margin. ‘The largest spivit-

specimen measures 32 mm. in length by 4 mm. in greatest

breadth.

One of the Maria Island specimens was specially noted during

life as being very dark brown dorsally. Narrow median stripe

and mottlings still visible. In addition also microscopic specks

of bluish white thickly dusted over dorsal surface.

The following notes were made on living specimens collected

in the neighbourhood of the Great Lake :—‘‘ When at rest very

“broad and much flattened on both surfaces. When crawling

‘long and narrow, with horseshoe-shaped anterior tip and pointed

‘“‘ posterior extremity. Hyes as usual. Dorsal surface pale grey

“with narrow longitudinal stripe of dark grey-brown down the

“middle, and thickly marbled with olive-brown, the markings

700 PROF. ARTHUR DENDY ON

“showing a slight tendency to longitudinal arrangement. Ventral

“ surface pale grey without markings, nearly white. Anterior tip

“pinkish brown. Colour of dorsal markings reddish brown,

“almost chestnut, in some specimens.” In these specimens the

tendency of the markings on‘ the dorsal surface to arrange them-

selves in three longitudinal bands on each side of the narrow,

mid-dorsal, dark stripe, is more marked than in the Maria

Island specimens. These specimens also do not attain so large a

size as those from Maria Island, measuring in spirit about 25 by

4 mm.

This species is evidently closely related to the Australian

G. quadrangulata, especially to the mottled Mount Wellington

(Victoria) variety of that species. It is, however, a very much

more robust form *, being intermediate in this respect between

G. quadrangulata and the Tasmanian G. diemenensis.

The relationship between these three species requires careful

investigation. G. diemenensis, it may be remembered, is charac-

terized by the presence of remarkable comb-like copulatory

organs, and is accordingly included by von Graff [1899] in his

genus Artioposthia. Such organs have not been observed in

G. quadrangulata or in G. nichollsit, but we must not overlook the

possibility that all the specimens of these species hitherto found

have been immature and that the peculiar copulatory organs may

be developed only in full-grown individuals. The specimens of

G. diemenensis in which I observed these organs were collected in

February { and March, while those of G. nichollsi described above

were collected in September. The size of the specimens may also

depend largely on the time of year. Unfortunately, we do not

yet know nearly enough about the life-history of land-planarians

to enable us to settle these points. :

Steel has already expressed a doubt as to whether all the

specimens originally referred by me to G. diemenensis, and

accepted as such by von Graff, really belong to the same species.

He himself gives a figure of a specimen of “ G. diemenensis,” one

of a small series of examples from 'Trevallyn Hills and Table

Cape, but lam doubtful whether it really belongs to that species—

or yet to G. nichollst, which it seems to resemble fairly closely in

pattern—because the transverse section appears to be of quite

different shape, not at all quadrangular. Possibly, however, the

outline given represents the transverse section in life, which is

very different from what it is in spirit-specimens.

In the type-specimens of 6. diemenensis from Mount Wellington

(Tasmania) there was no narrow median longitudinal stripe on the

dorsal surface, and I am inclined to think that the presence of

such a stripe in G. nichollst may serve as a valid specific distinction.

* T have, however [1895], described a robust variety of G. quadrangulata from

the Blue Mountains, N.S.W.

+ See, however, foctnote below. C

* This was one of the Parattah specimens, and as it shows a narrow, mid-dorsal,

dark stripe, I strongly suspect that it may be a specimen of G. xichollsi.

LAND-PLANARIANS FROM WEST AUSTRALIA. 701

G. nichollsi may approach pretty closely to the common

G. tasmaniana in general appearance, but may be distinguished

by the sharper dorso-lateral margins, the sides of the body being

rounded off (in spirit) in G. tusmaniana.

GEOPLANA MORTONI Dendy.

Geoplana mortoni Dendy [1894].

Geoplana mortoni von Graff [1899].

Geoplan« mortoni Steel [1901 bis].

We obtained a dozen good specimens of this beautiful and

well-characterized species in September 1914, in the neighbour-

hood of the Great Lake. It had been previously obtained by

the late Mr. Alexander Morton from an unknown locality in

Tasmania, and by Professor Baldwin Spencer from Parattah, and

my own acquaintance with it was limited to spirit-specimens.

The foilowing observations were made on a living specimen :—

‘“« At rest very broad and much flattened, no median dorsal ridge.

** Dorsal surface dull yellow, closely and finely mottled with chest-

‘nut-brown. Horseshoe-shaped anterior tip brown. Eyes as

“usual. Ventral surface white, much less closely mottled with

“small brown specks, with very narrow marginal band free from

“ mottling. When crawling, dorsal surface very strongly arched.”

A good-sized specimen when crawling measured about 50 mm. in

length by 4:5 mm. in greatest breadth, and one of my specimens

measures as much after preservation in spirit.

The shape of the body in spirit-specimens is characteristic ;

convex above and concave below, especially in the anterior

portion ; broader behind than in front; with prominent, narrow

margins at the junction of dorsal and ventral surfaces.

GEOPLANA TyPHLops Dendy.

Geoplana alba Dendy [1892].

Geoplana typhlops Dendy [1894].

Geoplana typhlops von Graft [1899 }.

This curious eyeless species appears to be one of the commonest

of the Tasmanian land-planarians. It has previously been re-

corded from Mount Wellington, Hobart and Parattah, and in

September 1914 we obtained it near the Great Lake and on Maria

Island (one small specimen). Some of the specimens in spirit

develop a distinct narrow median ventral band, due to darkening

of the rest of the ventral surface. I have already noticed this

feature In my previous descriptions.

GEOPLANA FLYNNI Sp. nh.

I propose this species for four specimens collected by Dr.

Nicholls on Maria Island, and have much pleasure in naming it

after Professor Flynn of Hobart, who so successfully organized

the zoological excursions for members of the British Association

in Tasmania.

Proc. Zoot. Soc.—1915, No. XLVIII, 48

702 PROF, ARTHUR DENDY ON

The worm when crawling is long and narrow, convex above

and flat below. The ground-colour of the dorsal surface in life

ranges from grey to “allow, but even in the grey specimens it

becomes fairly bright yellow in spirit. Minute brown specks

are sparsely scatter ed over the dorsal surface and tend to arrange

themselves in a very narrow median stripe. The horseshoe-

shaped anterior tip 1s brown or brownish pink, and the ventral

surface nearly white, without markings. The eyes are arranged

as usual. In spirit the body is narrow, more or less flattened,

especially ventrally, and with fairly well pronounced lateral

margins.

Im one specimen (in spirit) the peripharyngeal aperture is

cated well in front of the middle of the body, and the genital

aperture rather nearer to it than to the posterior end. This is

the only one in which both apertures are visible, but in another

the peripharyngeal aperture seems to be about central; but of

course the position of the apertures is influenced by the ‘telative

state of contraction of the two ends of the body.

One of the four specimens is much larger than the other three,

measuring in spirit about 32 mm. in Jength by a shade over

3 mm. in greatest breadth. dine sinaller ones in spirit measure

about 19 mm. in length by 2°5 mm. in breadth. In life one of

them measured alone 32 by 2 mm.

I think that the specimen which I described [1892] under the

name Geoplana sp., from near Hobart, probably belongs to this

species. Von Graff [1899] referred this specimen to the

Victorian G. guadrangulata var. wellingtoni, but there was very

little justification for so doing, or, I fear, for my own comparison

of it with G. quadrangulata and G’. ventropunctata.

List of Literature referred to.

1844. Darwin, C.— Brief Descriptions of several terrestrial

Planarie, &e. (Ann. & Mag. Nat. Hist. vol. xiv.)

1890. Denby, A.—On the Victoria Land-Planarians. (Trans.

Royal Soc. Victoria for 1890.)

1891. /bid.— Additional Observations on the Victorian Land-

Planarians. (Trans. Roy. Soc. Victoria for 1891.)

1892. /bid.—Notes on some Land-Planarians from Tasmania and

South Australia. (Proc. Aust. Assoc. Adv. Sci. Hobart,

1892.)

1894. /bid.—Notes on some new or little-known Land-Planarians

from Tasmania and South Australia. (Proc. Royal Soe.

Victoria for 1893.)

1895. /bid.—Notes on some Land-Planarians collected by Thomas

Steel, Esq., F.C.S., in the Blue Mountains, N.S.W.

(Proc, Linn. Soc, N.8.W., Nov. 1894.)

LAND-PLANARIANS FROM WEST AUSTRALIA. 703

1895 bis. Zbid—The Cryptozoic Fauna of Australasia. (Presi-

1896.

1887.

1899.

LSOT.

LOO

dential Address, Section D, Proc. Aust. Assoc. Adv. Sei.

Brisbane, 1895.)

Ibid.—Notes on New Zealand Land-Planarians, Part 2.

(Trans. N.Z. Inst. for 1895, vol. xxviii.)

FiercHer, J. J., and Hamriron, A. G.—Notes on Austra-

lian Land-Planarians, with Descriptions of some new

Species. (Proc. Linn. Soc. N.S.W., June 1887.)

GraFF, L. von.—Monographie der Turbellarien. II. Tricla-

dida Terricola (Landplanarien). (Leipzig.)

Sree, T.— Austraiian Land-Planarians, &e. (Pxoc. Linn.

Soc. N.S.W., April 1897.)

/bid.—Australian Land-Planarians, &ce. (Proc. Linn. Soe.

N.S.W., Sept. 1900.)

1901 bis. 7bid.—Tasmanian Land-Planarians, &e. (Proc. Linn.

Soe. N.S. W., Sept. 1900.)

48%

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 70

EXHIBITIONS AND NOTICES.

October 26, 1915.

Prof. E. W. MacBripg, D.Sc., F.R.S., Vice-President,

in the Chair.

The Secrerary read the following report on the Additions

made to the Society’s Menagerie during the months of June,

July, August, and September, 1915:

JUNE.

The registered additions to the Society's Menagerie during the

month of June were 249 in number. Of these 61 were acquired

by presentation, 87 by purchase, 36 were received on deposit, 9 in

exchange, and 56 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 161.

Amongst the additions special attention may be directed

UO) 2

1 Pallas’s Cat (Felis manul) 3, from Tibet, and 2 Burrhel

Sheep (Pseudois nahura) 2 2, from the Himalayas, deposited on

June 19th.

1 Hybrid Chapman’s and Mountain Zebra (Hquus chapmanni

x H. zebra) 3, born in the Menagerie on June 17th.

1 White-browed Wood-Swallow (Artamus superciliosus), hatched

in the Menagerie on June 17th.

1 Lesser Double-collared Sunbird (Cinnyris chalybeus) from

South Africa, presented by Alfred Ezra, F.Z.8., on June 12th.

2 White Storks (Ciconia ciconia), hatched in the Menagerie on

June 8th.

A collection of Pheasants, including 3 Vieillot’s Firebacks

(Lophura rufa), 4 Himalayan Monauls (Lophophorus impeyanus),

and others, purchased on June 5th.

2 Ocellated Turkeys (Meleagris ocellata), from British Honduras,

deposited on June 21st.

JULY.

The registered additions to the Society's Menagerie during the

month of July were 382 in number. Of these 260 were acquired

by presentation, 34 were received on deposit, 45 in exchange, and

43 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 241.

706 THE SECRETARY ON ADDITIONS TO THE MENAGERIE,

Amongst the additions special attention may be directed

10)

2 Puma cubs (Melis concolor) and 1 Kyra Cat (Melis eyra), from

Cordova, Argentina, presented by Wilfred A. Smithers, C.M.Z.5.,

on July 28th.

2 Californian Sea-Lions (Otaria californiana), from California,

received in exchange on July 19th.

1 Eland (Laurotragus oryx), born in the Gardens on. July 1dth.

A large collection of Birds, mostly European, presented by

Major Frank Johnson, F.Z.8., on July 19th.

A collection of North American Birds, received in exchange

from the Zoological Society of New York on July 19th.

2 Buff-breasted Partridges (Ptilopachys fuscus), 9 West-Afvican

Quail-Finches (Ortygospiza atricollis), and other birds from

Gambia, presented by Dr. E. Hopkinson, D.8.0., F.Z.5., on

July 21st.

2 Chinese Starlings (Stwrnia sinensis), new to the Collection,

received in exchange on July 28th. ’

10 Common Rheas (Rhea wmericana), hatched in the Menagerie

on July 18th.

1 Blackish Tortoise (Zestudo nigrita), from Indefatigable

Island, and 1 Baur’s Tortoise (7. galapagoensis), new to the

Collection, from Charles Island, deposited on July 29th.

1 Royal Python (Python regius), from West Africa, presented

by Robert R. Jones on July 19th.

AUGUST.

The registered additions to the Society’s Menagerie during the

month of August were 129 in number. Of these 89 were acquired

by presentation, 21 were received on deposit, 1 in exchange, | by

purchase, and 17 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 115.

Amongst the additions special attention may be directed

2 Orang-Utans (Simia satyrus), from British North Borneo,

deposited on August 7th.

| Rosy Minivet (Pericrocotws roseus), 1 Rufous-backed Shrike

(Lnneoctonus erythronotus), 1 Rufous-bellied Babbler (Dumetia

hyperythra), 1 Green Lora (githina viridissima), from India,

all new to the Collection, presented by E. W. Harper, F.Z.8., on

August 4th.

A collection of Mammals, Birds, and Reptiles, including a

Korin Gazelle (Gazella rufifrons), a Crowned Duiker (Sylvicapra

coronata), a Waterhouses Genet (Genetta poensis), 2 White-

necked Crows (Corvus scapulatus), a Long-nosed Crocodile

(Crocodilus cataphractus), and a Moebius’s Snake (“lapechis

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. (07

moebii), the last-named new to the Collection. from the

Northern Territories of the Gold Coast, presented by Capt. C.

H. Armitage, C.M.G., D.8.0., F.Z.S., on August 18th.

SEPTEMBER.

The registered additions to the Society’s Menagerie during the

month of September were 115 in number. Of these 65 were

acquired by presentation, 32 were received on deposit, 7 in

exchange, 8 by purchase, and 3 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 123.

Amongst the additions special attention may be directed

1 Ocelot (4elis pardalis), 1 Kinkajou (Potos caudivolvulus),

1 Fraser’s Squirrel (Scvwrus stramineus), 1 Spiny Tree-Porcupine

(Coendu prehensilis), | Harris’s Owl (Gisella harrisi), 1 Salmon’s

Tiger-Bittern (TLigrisoma salmoni), and 4 Peruvian Ground-

Doves (Chamepelia cruziana), from HEeuador, presented by EH. J.

Brook, F.Z.S., on September 25th and 29th. The three species

of birds are new to the Collection.

10 Wilson’s Birds of Paradise (Schlegelia wilson), from Waigiou,

new to the Collection, deposited, and 2 of the same presented, by

A. Hi. Pratt on September Ist and 8th.

1 Black Manucode (Manucodia atra), and 7 Red Birds of

Paradise (Paradisea rubra), deposited on September 13th.

2 Red Birds of Paradise (Paradisea rubra), and 4 Rufous-

erowned Ground-Doves (Phlogenas rufigula), the latter new to

the Collection, from Waigiou, purchased on September 16th.

2 Chestnut-backed Finches (Spermestes nigriceps), new to the

Collection, received in exchange on September 8th.

Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals, ex-

hibited a series of burrows of trapdoor spiders from South Africa,

belonging to types never previously brought to England. They

were collected at Alicedale, near Grahamstown, hy Messrs. F.

Thurston and F. Cruden, and were given to Mr. Pocock by the

former gentleman, who at the same time presented to the Society

living examples of Stasimopus and Gorgyrella.

Mr. D. Sers-Smiru, F.Z.8., Curator of Birds, exhibited a ving

hybrid Swinhoe’s and Silver Pheasant (Genneus nycthemerus

x G. swinhow), which had been presented to the Scciety by

Mr. H. J. Elwes, F.R.S. It was a male, and a very handsome

bird, but, although somewhat resembling Swinhoe’s Pheasant as

regards the head and neck, the back, rump, and tail were of a

chestnut-brown colour, differing from the males of both parent

708 MR. D. SETH-SMITH ON AN ABNORMAL DUCK,

species. Mr. Elwes had also presented to the Society a young

bird which he had bred from the above-mentioned hybrid, paired

to a pure hen Swinhoe Pheasant, but this bird was not yet in

adult plumage.

Mr. Serx-Smiry also exhibited a male Mandarin Duck (We

gulericulata) bred in the Society’s Gardens during the current

year, in which an extra digit had grown from the inner side

of the right tibiatarsal joint (text-fig. 1). He remarked that

Text-figure 1.

Right leg of male Mandarin Duck, with supernumerary digit.

similar cases had been recorded by Bateson (‘Materials for

the Study of Variation,’ page 394), but he believed that such

abnormalities were rare, especially in birds other than those

long domesticated.

NOTES FROM THE CAIRD INSECT HOUSE. 709

November 9, 1915. :

Dr. 8S. F. Harmer, M.A., F.R.S., Vice-President,

in the Chair.

The Secretary read the following report on the Additions to

the Society’s Menagerie during the month of October 1915 :—

The number of repister ed ailditions to the Society's Menagerie

during the month of October was 162. Of these 98 were acquired

by presentation, 24 were received on deposit, 15 in exchange, ih

by purchase, and 14 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 206.

Amongst the additions special attention may be directed to :—

1 Hairy Tree-Porcupine (Coendu insidiosus), from Venezuela,

presented by Hugo Pam, C.M.Z.S., on October 4th.

2 Larger-streaked Spider-hunters (Arachnothera magna), from

the Himalayas, new to the Collection, presented by Alfred Ezra,

F.Z.S., on October Ist.

1 Red-faced Crake (Neocrex erythrops), new to the Collection,

presented, with a number of other birds from Venezuela, by

Hugo Pam, 0.M.Z.8., on October 4th.

1 Japanese Robin (Hrithacus akahige), from Japan, and

1 Yellow-collared Ixulus (Jvulus flavicollis), from the Hima-

layas, both new to the Collection, deposited on October 13th.

1 Greater Amethyst Sunbird (Chalcomitra amethystina), from

South Africa, new to the Collection, presented by Alfred Ezra,

F.Z.8., on October 20th.

1 Sauvage’s Tree-Frog (Phyllomedusa sauvagii), from Cordova,

Argentina, new to the Collection, presented by Wilfred A.

Smithers, C.M-Z.S., on October 20th.

Prof. H. Maxwent Lerroy, M.A., F.Z.S., Curator of Insects,

read a report on the House-Fly Investigations carried out during

the present year in the Society's Gardens.

It is hoped that this report will be published in full in the

‘ Proceedings’ for 1916.

Notes from the Caird Insect House.

Mr. C. J. C. Poot, Assistant Curator of Insects, read the

following notes upon species recently bred and exhibited :—

ORTHOPTERA.

MANTID 4.

Sphodromantis guttata.

The female Mantid deposits her eggs in a case made of a

gummy secretion. The eggs are in regular rows inside the case,

which is attached to a twig or to the bark of a tree. There is an

710 MR. CG. J. C. POOL:

opening along the centre of the outer surface, each egg-cell being

protected by a flap or door, ‘The young emerge in a cluster and

on reaching the exterior they remain suspended by threads

> tubes for from 3 to 5 days. During this period they are

eae of seizing other insects or of feeding i in the ordinary

way, and yet they obtain sufficient nourishment to increase their

size very consider ably within a week. I have observed this de-

velopment upon several occasions, and it is my opinion that we

must regard the egg-case as a kind of feeding-bottle capable of

supplying the young Mantids with liquid food, absorbed through

the threads or tubes by which they are suspended until the first

moult. After this important event they are active and are at

last capable of an independent existence. To test my “ feeding-

bottle” theory I have on several occasions severed the con-

nection between the young larva and the egg-case. The result

was the same every time, and the creature died without further

development.

I have examined several cases from which the young had

emerged, and found that they all contained fluid matter which

might form the necessary food, probably with the assistance of

rain or dew.

The specimen referred to in a previous paper (P. 4.8. 1915,

p- 289) lived from 6th July, 1914, until 24th September, 1915.

On 19th August, 1915, it imagined a front leg, a mishap which

reduced its chances of seizing a meal. ljaen the third day

following the accident, the Mantid surprised me by devouring

the offending and sill unsevered limb, after which it appeared

to revive its. interest in life by depositing an ege-mass, the

eighth in its lifetime in the Society’s Gardens. It “died shortly

after this ev ent, having completed nearly fifteen months of

active life.

Sphodromantis gastrica.

An egg-mass from Pretoria produced some young Mantids on

28th I May, 1915. The majority of these died in the earlier

stages. One fine female specimen, having devoured a number

of her own kind and an abundance of other insect- food, arrived

at maturity and developed her wings on October 18th, 1915

having occupied nearly five months in the process. This speci-

men is still alive and healthy, her food consisting principally of

stick-insects. A few egg-masses produced numerous small

Parasitic Hymenoptera (Chalcids) new to science, which when

described will form the subject of another paper.

COLEOPTERA.

DERMESTIDA.,

Thaumaglossa bimaculata Arrow.

Some Mantid egg-masses received from Pretoria in May 1915

were found to be inhabited by Coleopterous larvee much resembling

NOTES FROM THE CAIRD INSECT HOUSE. men

the museum pest Anthrenus musearum. The infested cases

were isolated and kept under observation, he larvee fed upon

the contents of the egg-masses, and duly pupated within their

own skins. ‘The beetles have been emerging during September

and October.

Lhe genus Thaumaglossa is represented in the National Col-

lection by eight species known to inhabit egg-masses of Mantids,

their distribution being China, S. Australia, Nigeria, Ronde,

and the Malay Peninsula. The insect now exhibited was ex-

amined by Mr. G. J. Arrow at the British Museum (Nat. Hist.)

and has been described by him under the above name.

The type-specimen will be deposited in the National Collection

and the co-type in the general reference collection now in

course of formation in the Caird Insect House.

PTINID#.

Ptinus pusillus Starm.

This active little beetle is a well-known granary pest in France

and Germany, but was not known to inhabit this country until

1906, when I discovered it in a corn-shop at Edmonton.

The female, which is more robust than the male and with

shorter antenne, has for some years been confounded with

another species, P. testacews Boield., in British works and

collections.

I have taken both sexes in profusion in the Society’s Gardens,

and it is due to the study of the specimens collected there and at

Edmonton that I have been able to clear up this old error,

Ptinus tectus Boield.

This beetle is another cosmopolitan granary pest, and during

the last few years has become exceedingly common throughout

this country. During last May a so-called insect-destroyer v

recommended and submitted for test upon some Blow-fly maggots.

It was not a success, its intended victims proving its ineffective-

ness by simply pupating in a dish of the supposed deadly powder.

The paper-bag containing the remainder was put aside in a cup-

board in the Fly-room, and was forgotten until after the closing

of the Fly Exhibition in September, when upon clearing out an

accumulation of samples, etc., I noticed numerous small round

perforations in the bag which had become slightly mouldy, and,

upon peeling back the paper near the perforations, I discovered

P. tectus, larvee, pupe, and imagos, perfectly healthy and ap-

parently enjoying the novelty of breeding in an insect-killing

powder, which actually appears to be a vegetable-meal upon

which a corn-shop pest might be expected to thrive.

In conclusion, I must confess to a feeling of pleasure in

recording the successful rearing of an African beetle new to

Tlie NOTES FROM THE CAIRD INSECT HOUSE.

science in the Caird Insect House, as well as the correction of

an error which has caused considerable confusion in connection

with the species of Ptinus. We have been handicapped by

reduction of staff, our two helpers having joined the colours,

otherwise we might have accomplished some other useful

research-work with the aid of valuable living material received

from various sources. Some day, under happier circumstances,

we hope to be able to devote more attention to breeding, etc.,

thereby adding to the scientific value as well as to the popular

interest of the Caird Insect House.

November 23, 1915.

Dr. A. Smira Woopwarp, F.R.S., Vice-President,

in the Chair.

Mr. R. W. Harotp Row, B.Sc., F.Z.S., exhibited photographs

of an interesting abnormal Frog (Rana temporaria), in which both

posterior limbs lacked the hallux, though the calear, or prehallux,

was present on each side. he characteristic structure of the

digits still existing clearly proved that the missing toe was the

hallux, and dissection showed that there was no trace of it

internally. All the other four digits were perfectly normal.

A complete description of this abnormality, with illustrations,

will be published in the next part of the ‘ Proceedings.’

No. 148.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON,

October 26th, 1915.

Prof. E. W. MacBrrpg, D.8c., F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Srcrerary read a Report on the Additions to the Society’s

Menagerie during the months of June to September, 1915.

Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals,

exhibited a series of burrows of trapdoor spiders from South

Africa, belonging to types never previously brought to England.

They were collected at Alicedale, nr. Grahamstown, by Messrs.

F. Thurston and F. Cruden, and were given to Mr. Pocock by

the former gentleman, who at the same time presented to the

Society living examples of Stasimopus and Gorgyrella.

Mr. D. Sera-Surra, F.Z.8S., Curator of Birds, exhibited a

Mandarin Duck (#« galericulata) in which an extra digit had

grown from the inner side of the right intertarsal joint. He

remarked that similar cases had been recorded by Bateson

(‘Materials for the Study of Variation,’ page 394), but he

believed that such abnormalities were rare, especially in birds

other than those long domesticated.

Mr. Sera-Smiru also showed a living hybrid Swinhoe’s and

Silver Pheasant, which had been presented to the Society by

Mr. H. J. Elwes, F.R.S. It was a male, and a very handsome

bird, but, although somewhat resembling Swinhoe’s Pheasant as

regards the head and neck, the back, rump, and tail were of a

* This Abstract is published by the Society at its offices, Zoological Gardens,

Regent's Park, N.W., on the Tuesday following the date of Meeting to which

it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,

to all Fellows who subscribe to the Publications; but it may be obtained on the

day of publication at the price of Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in adyance,

chestnut-brown colour, differing from the males of both parent

species. Mr. Elwes had also presented to the Society a young

bird which he had bred from the above-mentioned hybrid, paired

to a pure hen Swinhoe Pheasant, but this bird was not yet in

adult plumage.

Mr. E. G. Boutencsr, F.Z.8., Curator of Reptiles, read a

paper on the feeding of Snakes in captivity, based on observations

made upon specimens in the Society’s Collection. The results

showed that, with rare exceptions, snakes that refused to feed on

dead animals were not more likely to accept them if alive.

Dr. 8S. F. Harmer, F.R.S., F.Z.8., read a preliminary paper

“On Specimens of Cuvier’s Whale (Ziphius cavirostris) from the

Trish Coast.”

The inclusion of Z. cavirostris in lists of the Cetacea of the

British Seas appears to rest on the evidence of a single skull

obtained by Sir William Turner from Shetland. By an arrange-

ment made with the Board of Trade in 1912, the British Museum

receives telegraphic reports of the stranding of Cetacea on the

British Coasts. Two of the specimens thus reported have proved

to belong to this rare species, and their skeletons have been

secured for the Museum. One of these individuals, stranded in

Co. Wexford on July 18, 1915, was ascertained to be a male, and

it possessed a pair of massive teeth, about 14 inches in diameter

and 13 inches in length, at the anterior end of the lower jaw.

The sex of the other specimen, which was stranded in Co. Cork

on February 13, 1913, could not be ascertained, although its

cranial characters point to its having been a female. The teeth

were completely hidden beneath the gum, but dissection showed

that two were present, in the same position as those of the male,

but considerably smaller, their diameter only slightly exceeding

half an inch. Casts of the lower jaws were exhibited. It was

suggested that Cuvier’s Whale is likely to prove less rare than

has hitherto been: supposed, as both specimens here recorded

were at first believed to belong to Hyperoodon rostratus. It will

be desirable to scrutinise carefully future records of ‘ Bottle-

nosed Whales,” in the hope of ascertaining that some of them

really belong to Ziphius or to the allied genus Mesoplodon.

Dr. F. E. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the

Society, read a paper on Avian Cestodes entitled “On Tania

struthionis Parona and Allied Forms,” in which he defined a

probable new species of Davainea parasitic in the Ostrich

(Struthio masaicus).

A paper was read from Mrs. Mary G. Roserts, C.M.Z.S., on

the successful breeding and rearing of the young of a pair of

Tasmanian Devils (Sarcophilus harrisi), kept in captivity at

Hobart.

Professor 8. J. Hickson, M.A., F.R.S., F.Z.S., contributed a

paper on some Alcyonaria and a Stylaster from the west coast of

North America, Three new species of Aleyonaria were described

therein. ,

The next Meeting of the Society for Scientific Business will be

held on Tuesday, November 9th, 1915, at half-past Five o’clock

p.M., when the following communications will be made :—

EXHIBITIONS AND Novices.

G. E. Nicuotzs, D.Sc.

Some Notes upon the Anatomy of Rana tigrina.

. C. Morrram, M.B.(Lond.).

(1) The Distribution of Secondary Sexual Characters

amongst Birds, with relation to their liability to the Attack

of Enemies.

(2) Some Observations on Pattern-Blending with Reference

to Obliterative Shading and Concealment of Outline.

C. Boprn Kuioss, F.Z.S8., F.R.G.S.

On a Collection of Mammals from the Coast and Islands of

S.E. Siam, with an account of the Fruit-Bats by Dr. Knud

Andersen.

Prof. W. J. Dakin, D.Sc., F.Z.S8.

Fauna of West Australia.—III. A new Nemertean—Geo- -

nemertes dendyi, sp. n.—being the first recorded Land

Nemertean from Western Australia.

IV. Palemonetes australis, sp. n., being the first record of

the Genus in Australia.

The following papers have been received :—

G. A. Boutencer, F.R.S., F.Z.S.

(1) A List of the Snakes of Hast Africa, North of the

Zambesi and South of the Soudan and Somaliland, and of

Nyassaland.

(2) A List of the Snakes of North-Hast Africa, from the

Tropic to the Soudan and Somaliland, including Socotra.

(3) Deseriptions of a new Amphisbena and a new Snake

discovered by Dr. H. G. F. Spurrell in Southern Colombia.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,

Recent’s Park, Lonpon, N.W.

November 2nd, 1915.

No. 149.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON,.*

November 9th, 1915.

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

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secrerary read a Report on the Additions to the Society’s

Menagerie during the month of October, 1915.

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

read a report on the House-Fly Investigations carried out during

the present year in the Society’s Gardens.

Mr. C. J. C. Poon, Assistant Curator of Insects, read some

notes upon a number of species which had been bred and

exhibited in the Caird Insect House.

Dr. G. E. Nrcuoxts read a paper containing an account of the

anatomy of Rana tigrina, the so-called Bull-frog of India, and

drew attention to certain features in which this species differs

from its Huropean congeners,

Dr. J. C. Morrram read a paper on “ Pattern-blending with

reference to Obliterative Shading and Concealment of Outline.”

The paper recorded the results of laboratory experiments with

* 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 cn the

day of publication at the price of Simpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in adyance,

artificial patterns. The experiments showed that obliterative,

or counter-shading could be produced by blended black-and-

white pattern, and that beyond the blending distance, inter-

ruptions at the margin of a pattern, or similarly placed eye-spots,

blur the margins. The laboratory experiments were compared

with actual patterns of animals.

Dr. Morrram also read a paper “On the Distribution of

Secondary Sexual Characters amongst Birds, with relation to

their Liability to the Attack of Enemies.” The paper was based

upon a statistical enquiry into the possible existence of a

correlation between these factors.

Mr. C. Bopen Kuoss, F.Z.8., F.R.G.S., contributed a paper

on a collection of Mammals made by him on the coast and

islands of 8.H. Siam, over 500 specimens being obtained. One

species and twenty-two subspecies were described ag new.

Two papers, dealing with the Fauna of West Australia, were

received from Prof. W. J. Dakin, D.Se., F.Z.S. The first paper

contained the description of a new Land Nemertean, the first to

be recorded from West Australia. The second paper described a

new prawn-like Crustacean of the genus Palemonetes, which

genus had not hitherto been recorded in Australia.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, November 23rd, 1915, at half-past Five o’clock

P.M., when the following communications will be made :—

EXHIBITIONS AND NoTvICcEs.

ean

G. A. Boutencsr, F.R.S., F.Z.S.

ak SD

(1) A List of the Snakes of East Africa, North of the

Zambesi and South of the Soudan and Somaliland, and of

Nyassaland.

(2) A List of the Snakes of North-East Africa, from the

Tropic to the Soudan and Somaliland, including Socotra.

(3) Descriptions of a new Amphisbena and a vew Snake

discovered by Dr. H. G. F. Spurrell in Southern Colombia.

Prof. ArrHur Denpy, D.Sc., F.R.S., F.Z.S.

On some Land-Planarians collected in West Australia and

Tasmania by Members of the British Association for the

Advancement of Science.

C. Tare Reean, M.A., F.Z.S.

| ES CI

The Morphology of the Cyprinodont Fishes of the Subfamily

Phallostethine.

The following paper has been received :—

J. H. Lioyp, M.Sc.

Some Observations on the Structure and Life-history of the

common Nematode of the Dogfish.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,

Recent’s Park, Lonpon, N.W.

November 16th, 1915,

No. 150.

ABSTRACT OF THE PROCEHDINGS

ZOOLOGICAL SOCIETY OF LONDON.

November 23rd, 1915.

Dr. A. Smrra Woopwarb, F.R.S., Vice-President,

in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. BR. W. Haroxp Row, B.Sc., F.Z.8., exhibited photographs

of an interesting abnormal Frog, in which both posterior limbs

lacked the hallux, though the calcar, or prehallux, was present on

each side. The characteristic structure of the digits still existing

clearly proved that the missing toe was the hallux, and dissection

showed that there was no trace of it internally. All the other

four digits were perfectly normal.

Prof. ArrHur Denpy, D.Sc., F.R.S., F.Z.S., read a paper on a

collection of Land-Planarians made by members of the British

Association in West Australia and Tasmania. Three species were

obtained in West Australia, all of which proved to be new, and

of six species collected in Tasmania, two were described as new.

Mr. G. A. Boutencsr, F.B.S., F.Z.S., gave an account of two

papers containing lists of the Snakes of Hast Africa and Nyassa.

land, and of North-east Africa and Socotra, with keys to the

identification of the genera and 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 Sixpence, or, if desired, sent post-free for

the sum of Six Shillings per annum, payable in advance.

Mr. BovnencEr also read a paper in which he described a

new Amphisbena and a new Snake discovered by Dr. H. G. F.

Spurrell in Southern Colombia.

Mr. C. Tavs Reean, M.A., F.Z.S., read a paper on the morph-

ology of the Cyprinodont Fishes of the subfamily Phallostethinee.

He described the structure of these extraordinary little fishes

from Johore, and particularly the differences in the priapium of

the two genera he recognised.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, February 8th, 1916, at Half-past Five o’clock

p.M. The Agenda will be announced early in January.

The following paper has been received :—

Prof. E. B. Poutton, M.A., F.R.S., F.Z.S.

On a Collection of Moths made in Somaliland by Mr. W.

Feather. With Descriptions of new Species by Sir G. F.

Hampson and others.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,

Recent’s Park, Lonpon, N.W.

November 30th, 1915.

39.

AO.

42,

43.

44.

AG.

47,

49.

50.

Papers (continued).

Fauna of West Australia—III. A new Nemertean, Geonemertes dendy?, sp. n., being

the first recorded Land Nemertean from Western Australia, By W. J. Daxin, D.Sce.,

F.Z.8., Professor of Biology, University of W. Australia. (Text-figure 1.)

Fauna of West Australia.—lV. Palemonetes australis, sp. n., being the first record of

the genus in Australia. By W. J. Daxin, D.Sc., F.Z.S., Professor of Biology,

Wintiversityzol Wie anstralia. «(Plate da)iasere wes ate Veta os bean eee

The Keeping and Breeding of Tasmanian Devils (Sarcophilus harvist). By Mrs. Mary

Go oRERTssCMeZ.S., MR ACO Ute (Wextefioure deo... chsc eso ees ke

Notes on the Feeding of Snakes in Captivity. By HE. G. Bouruycur, F.Z.S., Curator

CMERE ILS Sista cere EC eure onan aac ELE ik co py BuGLS i AER IES.

Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.—

XVIII. On Tenia struthionis (Parona) and Allied Forms By Frank E. Bepparp,

M.A., D.Sc., F.R.S., F.Z.8., Prosector to the Society. (Text-figures 1-6.)..........

Some Notes upon the Anatomy of Rana tigrina. By Guo. E. Nicuorrs,7D.8e., late

Professor of Biology, Agra College, India. (Text-figures 1-3.) ....-........0000e

. A List of the Snakes of East Africa, North of the Zambesi and South of the Soudan

and Somaliland, and of Nyassaland. By G. A. Bounmnesr, F.R.S., F.Z.S. (Text-

HS TURES Slr cbe eam ares tet ae eine ieee ere er aca Pratap) Se oad ore Sa a i

A List of the Snakes of North-East Africa, from the Tropic to the Soudan and

Somaliland, including Socotra. By G. A. Boutenaur, F\R.S., FZS. ......2.....

Descriptions of a new Amphisbena and a new Snake discovered by Dr. H. G. F,

Spurrell in Southern Colombia. By G. A. Bounenaur, F.R.S., F.Z.S. (Text-

HUMES a aE ong. Sects cho cite OR ae REE AG eRe SRE etic a a

. The Distribution of Secondary Sexual Characters amongst Birds, with relation to their

Liability to the Attack of Knemies. By J. C. Morrram, M.B.(Lond.).........+.--

Some Observations on Pattern-Blending with reference to Obliterative Shading and

Concealment of Outline. By J. C. Morrram M.R. (Lond.), (Text-figures 1-5.)

On some Land-Planarians collected in Western Australia and Tasmania by Members

of the British Association for the Advancement of Science. By Arruur Denpy, D.Se.,

F.R.S., F.Z.8., Professor of Zoology in the University of London (King’s College) . ,

Titlepage CeO CIC Osc (RORee tGioheiy 0-1 CHOSE UAIO Oo. Se ue sienna a Reape men Nts Glin ena ME shale

Eistion C onmeiMandyOiicers puis weep etn oor SU 6 yh CLA UN aT bg

Index

Page ©

567

571

575

583

589

603

611

641

659

663

679

693