by ers). 8

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL

SOCIETY OF LONDON.

1913.

COUNCIL.

His Grace Tue DuKke or Beprorpd, K.G., F.R.S., President.

Str JoHN Rose BRADFORD,

TEC KOR MICE WEIDER ID se..

F.R.S., Vice-President.

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

Lt.-Col. Sir R. Havetock

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

THe Rr. Hon. tHe Ear oF

@ROMER, PIC. 7G: @2Be

CC MEG KeEC Sale PRess

Vice-President.

F. G. Dawtrey Drewirtt, Esq.,

M.A., M.D.

CHARLES DRUMMOND,

Treasurer.

Str Epwarp Duranp, Bt., C.B.

FREDERICK GILLETT, Esq.

THE Lorp GLENCONNER.

F. Du Cane Gopmay,

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

Ksq.,

Sir Water Roper LAWRENCE,

13g, (CX CUIL IDL,

Str Epmunp G. Loprr, Bt.,

Vice- President.

Prof. Ernest W. MacBripe,

M.A., D.Se., F.R.S., Vice-

President.

Prof. Epwarp A. MINcHIN,

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

P. Cuatmers MircHett, Esq.,

M.A. DSc, LED. ass

Secretary.

W. R. Ocitviz-Grant, Esq.

ApriAn D. W. Pottocrk, Esq.

OLDFIELD THomas, Esq., F.R.S.

AntHony H. WINGFIELD,

Esq.

Henry Woopwarb, Ksq., LL.D.,

F.R.S., Vice-President.

PRINCIPAL OFFICERS.

P. Coatmers Mircuett, M.A., D.Sc., LL.D., F.RS.,

Secretary.

FrAnK EK. Bepparp, M.A., D.Sc., F.R.S., Prosector.

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

Resident Superintendent of the Gardens.

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

Epwarp G. BouLencer, Curator of Reptiles.

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

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

JoHN Barrow. Accountant.

W. H. Coxe, Chief Clerk.

LIST OF CONTENTS.

1913, pp. 1-337.

EXHIBITIONS AND NOTICES.

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

Menagerie during the month of October 1912 .........

The Secretary. Exhibition of a photograph of the Hainan

Gibbon (Hylobates hainanus) taken in natural colours.

Mr. G. A. Bounencser, F.R.S., F.Z.S. Exhibition of a

specimen of the African Cichlid Perch (Hemichromis

Dinnceilort ses Galan he cee auleeealsshev aan san ote ed lao Ns

Mr. Epwarp GerraArD. Exhibition of the Skull of an

Indian Rhinoceros (Rhinoceros unicornis) which had

recently died in the Society’s Gardens.....................

Mr. HK. G. Bouuencer, F.Z.8., Curator of Reptiles. Exhi-

bition of a living specimen of the Salamander,

Abi ypsbonmve tng en une ssi sain tersaeiessat ee vet seas os eto,

Mr. D. Sers-Smiry, F.Z.S., Curator of Birds. | Lantern

demonstration of photographs of young birds hatched

in the Society’s Gardens during 1912, and of the nest

made by the Sumatran Orang-utan. (PI. [X.).........

Page

Mr. C. Tate Reaan, M.A., F.Z.8. Exhibition of male and

female examples of the remarkable Cyprinodont Fish,

Cymolebras Delotta... ste. cen Makan oer ner re eee te

Mr. Oxprietp THomas, F.R.S., F.Z.S. Exhibition of a

specimen of the Amazonian Monkey, Callimico sneth-

UCL [2 ga Se BORE PIPES AS OEED tay Ss WARD AY Bac csocn nan 3

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

Menagerie during the months of November and

December Vou 2 i scenic sce gn evinesebelen wae emanate 150

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

a female Carolina Duck (Lampronessa sponsa) which-

had partially assumed male plumage ..................... 151

Mr. E. G. Boutencer, F.Z.S., Curator of Reptiles. Exhi-

bition of a blue specimen of the Edible Frog (Rana

esculenta), and of the remarkable Australian Lizard,

PUGOPUSNEDULOPUS, Baata rena a-ensne ote Ree Nese ce ee ene 151

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

Menagerie during the month of January 1913 ......... 151

Mr. KE. G. Bouteneer, F.Z.S., Curator of Reptiles. Exhi-

bition of the spines of an Insectivore found in the

EXCTeMEeNb O18 MOOR 5) dec ate e ee nee Ee eee eee 152

The Secretary. Exhibition of lantern-slides of the Tree-

Elvirax(Pendsolyn de COrsGlis) esis eee oe eee ate 152

The Secretary. Exhibition of a lantern-slide of an

abnormally-marked Domestic Donkey (Hquus asinus). 241

Mr. E. G. Boutencer, F.Z.8., Curator of Reptiles. Remarks

on the Central-African Lung-Fish (Protopterus ethi-

opicus) recently presented to the Society ............... 241

Dr. S. F. Harmer, M.A., F.R.S. Exhibition of a Hair-ball

Prom, Madagascar, go ccus-) cers molto seprecticklejctoeer eee 241

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

Menagerie during the month of February 1913......... 242

PAPERS.

. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.— VII. On Six Species of Tape-

worms from Reptiles belonging to the Genus /chthyo-

tenia (s. l.). By Frank EK. Bepparp, M.A., D.Sc.,

F.B.S., F.Z.8., Prosector to the Society. (Text-figs.

DOE) fo Sarsiay svat epicition stale egies ne aac SEEM coh ieda setae Se etaece

. Some Falkland Island Spiders. By H. R. Hoae, M.A.,

BAS iy. (Ris. Lge sli) i cose tec eisapeten! acl) basis actaog

. Descriptions of Three new Fishes discovered in the Gold

Coast by Dr. H.G. F. Spurrell, M.A., F.Z.8. By

Ga A] BoOUrENGHR gH SiS sy b)-Zecem (PIU \ar eae aie,

. On some Parasites of the Scoter Duck (Hdemia nigra),

and their Relation to the Pearl-inducing Trematode

in the Edible Mussel (Mytilus edulis). By H. LystEr

JamEsOoN, M.A., D.Sc., Ph.D, and Wiiiiam Nico.t,

NE AC ED See avis Dy a(Mextaitocemllnda lt) aime emacs nea

. Contributions to a Study of the Dragonfly Fauna of

Borneo.—Part J. The Corduliine : The Genus Amphi-

cnemis: The Legion Protoneura. By F. F. Laipiaw,

IME SAO EE Zip S eae CPIM ES Se ici cS ae ote ovis eae aagin snl eron van

. On the Structure of Bone in Fishes: a Contribution to

Paleohistology. By Epwiy 8. Goopricn, M.A.,

F.R.S., F.Z8., Fellow of Merton College, Oxford.

(Wextalies ml SOs ric sueeu ae ssa steis sages ecgie sh Nation oem

. Report on the Myzostomida collected by Mr. Cyril

Crossland in the Red Sea in 1905. By Cuaruzs L.

Boutencer, M.A., D.Sc., F.Z.S., Lecturer on Zoology

in the University of Birmingham. (Pls. V.-—VIII.

anid’ Next hosel OSa) i ve dsc tthe sols Sots Houeantnecaastenersst

. Description of an Amphipod belonging to the Family

Talitride, from the Woodbush, Transvaal. By Pau

Ay. MerauEny WAS. ..(Pis: XG Xs)! Menai ees.

Page

10.

13.

14.

15.

16.

18.

. The Genus Hngeus, or the Land Crayfishes of Australia.

By G. W. Smirn, M.A., Fellow of New College,

Oxford, and E. H. J. Scuuster, M.A., D.Sc., F.Z.S.,

Fellow of New College, Oxford. (Pls. XII.-XXYV.) .

On some Points in the Anatomy of the Mouth-parts of

the Mallophaga. By Bruce F. Cummines. (Text-figs.

24-32)

eee ecr ess oes eres s ease esossr es osoeeser%eOSesoeeorseresseoesene

Report on the Deaths which occurred in the Zoological

Gardens during 1912, together with the Blood-

Parasites found during the Year. By H.G. PimvMer,

FE.R.S., F.Z.S., Pathologist to the Society

eeccecceesevcese

. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.— VIII. On some Species of

Ichthyotenia and Ophidotenia from Ophidia. By

Frank E. Bepparp, M.A., D.8c., F.R.S., F.Z58.,

Prosector to the Society. (Text-figs. 33-38.)

The Anterior Ambulacrum of Hchinocardium cordatum

Penn., and the Origin of Compound Plates in Kchinoids.

By Herpert L. Hawkins, M.8c., F.G.S., Lecturer in

Geology, University College, Reading. (Pl. XXVI.

and Text-figs. 39-41.)

Coc eescee cere eresecesasecesaossesesssoes

Plankton from Christmas Island, Indian Ocean.—II. On

Copepoda of the Genera Oithona and Paroithona. By

CP SPARRAN. | (Es. XOX VI XOXO) erect neraccene

Diagnoses of New Species and Varieties of Agnathous

Mollusca from Equatorial Africa. By H. B. Preston,

WZ.) (PUSAN NON NEN cre teres eran acta aiera see

Notes on the Habits of Certain Reptiles in the Lagos

District. By W. A. Lamporn, M.R.CS. ...............

On the Gorgonopsia, a Suborder of the Mammal-like

Reptiles. By R. Broom, M.D., D.Sc, O.M.Z.8.

(Pls. XXXVI. & XXXVII.)

@esaseacesereaeeoesaesacesaersose

On two British Entomostraca belonging to the Orders

Copepoda and Ostracoda. By G. Strewarrson Brapy,

M.D...) LL.D. DSc.) 9H ss, 6 CMAs ae Cens:

XXXVIII.-XL.)

see PPP a er Cees et eesoeeesseseoseaesassorsaeragee

Page

128

141

153

169

181

194

218

1),

20.

ise)

vil

The Dwarf Buffalo of Southern Nigeria; with a

Revision of the Dwarf Buffaloes of Western Africa.

By R. Lyprexxer, F.R.S., F.Z.8S. (Text-figs. 42-44.) .

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—IX. On a new Genus of

Ichthyoteniids. By Frank EK. Bepparp, M.A., D.Sc.,

F.R.S., F.Z.S., Prosector to the Society. (Text-figs. 45—

DDe) Wait soins sia sa sionlblee eeaierna eee teuts mesg ena Sosa Gets

. oological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, 1904-1905.—

Report on the Branchiura. By Wiuiam A.

Cunnineton, M.A., Ph.D., F.Z.S. (Pls. XLI.-XLV.)

. Notes on Plankton collected across the mouth of the

St. Croix River opposite to the Biological Station at

St. Andrews, New Brunswick, in July and August

S12 By ARTHUR, Wilton MAD: Se: Hig.

F.Z.S., McGill University, Montreal. (Text-figs. 54 &

De ese tion alors Nas. Olan SMR Se ERs ira eteene cia raels Tatts oN bee ance

. Variations in the Skeleton of the Pectoral Fins of

Polypterus. By Epira E. Bamrorp, Newnham College,

Cambridge. (WVext-figs: 56-60.) 77... 2.2-nss--- neces

. A Descriptive Study of an Oligocheete Worm of the Family

Enchytreide ; with an Appendix on certain Commensal

Protozoa. By H. H. Stirrup, B.Sc. (Birm.), Lecturer

in Agricultural Biology, East Anglian Institute of

Agriculture, Chelmsford. (Pls. XLVI.-XLIX. and

JRepes thas (GUIO7s)s tanoseumnacaasascorecbeeneteconnnmoan awe onore

. The Relationship of the Big Game of Africa to the

Spread of Sleeping Sickness. By Dr. W. YorkKg,

Liverpool School of Tropical Medicine. With an

Appendix containing Remarks by Sir Jonn BLAND-

Sutton, F.R.C.S., F.Z.8.; Guy A. K. MArsHatt, F.Z.8. ;

Prof. E. A. Mincatn, M.A., F.R.S., V.P.Z.S.; The

Hon. L. Watter Roruscuitp, D.S8c., F.R.S., F.Z.S. ;

Sir Henry Seron-Karr, K.C.M.G., F.Z.S.; and Sir

ALFRED SHARPE, K.C.M.G., LL.D.; and Reply by

DB Sra COLT ZCD) Py as CE aie ea to ANE NG HE :

Page

234

243

262

283

300

MDT

wah 4

ete S. eet

¥ me rh etd

£4 LPR TE

Ave A Bye) DC 7AG lie ec Sue

of THE

CONTRIBUTORS,

With References to the several Articles contributed by each.

(1913, pp. 1-337.)

Page

Bamrorp, Miss Epira E., Newnham College, Cambridge.

Variations in the Skeleton of the Pectoral Fins of

olyprenu sty (hext=H9S.),. 06-00! iene cilees nits scissile

io)

is)

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

to the Society.

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—VII. On Six Species of Tape-

worms from Reptiles, belonging to the Genus /chthyo-

UCoOOG (Ge Uo)s (ARSE, TESINO)) he Gana qaase scans sae cnoneacaede 4

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—VIII. On some Species of Jchthyo-

tenia and Ophidotenia from Ophidia. (Text-figs. 33-38.), 153

Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—IX. On a new Genus of Ichthyo-

teniids. (Text-figs. 45-53.)..........00.. a chBPi Rona AARP HAN 243

Buanp-Surton, Sir Joun, F.R.C.S., F.Z.8.

Remarks on the Relationship of the Big Game of

Africa to the Spread of Sleeping Sickness. See YoRKE,

eeeececceeceor esc eo coe oe ee H eee o eee EEESHSEEESE Eoeosos EB oeeD ES ORS OEE

Bou.encer, Coartes L., M.A., D.Sc., F.Z.S., Lecturer on

Zoology in the University of Birmingham.

Report on the Myzostomida collected by Mr. Cyril

Crossland in the Red Sea in 1905. (Pls. V.-VIII. and

Text-figs. 17-23.)

Cee eer oee ses oes esse rs oseeesoseeeoesereeeesesosed

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

Exhibition of a living specimen of the Salamander,

Amblystoma tigrinum

ecw ecoeeaeso ne, OHH GSHFTHHF OEE OSES EHO TEO SEO EE OES

Exhibition of a blue specimen of the Edible Frog

(Rana esculenta), and of the remarkable Australian

Lizard, Pygopus lepidopus

eeeeereece sees eee esse esr ose oeeesteeDose

Exhibition of the spines of an Insectivore found in the

excrement of a Boa

eck ocr coerce c eer oee esse esesse ee resessesseoeesee

Remarks on the Central-African Lung-Fish (Proto-

pterus cethiopicus) recently presented to the Society ......

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

Exhibition of a specimen of the African Cichlid Perch

(Hemichromis bimaculatus Gill)

eecoeesece cores ecse see ose ceo ese

Descriptions of Three new Fishes discovered in the

Gold Coast by Dr. H. G. F. Spurrell, M.A., F.ZS.

CPT) oot ionic aiouti hs boa duct ebiaat Ma ugtt eck sotari eee tare ee

Brapy, G. Srewarpson, M.D., LL.D., D.Sc., F.R.S.,

C.M.ZS.

On Two British Entomostraca belonging to the Orders

Copepoda and Ostracoda. (Pls. XXXVIIL-XL.) ......

Page

151

152

241

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

On the Gorgonopsia, a Suborder of the Mammal-like

Reptiless-s (Piss XOX X Vis ds XOXO ae siccss seeds oleate

Cummines, Bruce F.

On some Points in the Anatomy of the Mouth-parts

of the Mallophaga. (Text-figs. 24-32.)

eceeecsccceoscoesseseree

Cunnineton, WittiAM A., M.A., Ph.D., F.Z.S.

Zoological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, 1904—1905.—

Report on the Branchiura. (Pls. XLI.-XLYV.)............

Farran, G. P.

Plankton from Christmas Island, Indian Ocean.—II.

On Copepoda of the Genera Oithona and Paroithona.

(TEI, OAH O- O70 Ih) cee sb noabo: soe cusgbasosanascomocnsad.ee5

GERRARD, EDWARD.

Exhibition of the Skull of an Indian Rhinoceros

(Rhinoceros unicornis) which had recently died in the

Society’s Gardens .........cccesececcrscecenscerenseeceeseesensoes

Goopricu, Epwin S., M.A., F.R.S., F.Z.8., Fellow of Merton

College, Oxford.

On the Structure of Bone in Fishes: a Contribution

to Paleohistology. (Text-figs. 13-16.) ..........:.0.....08

Harmer, Sipney F., M.A., Sc.D., F.R.S., F.Z.8.

Exhibition of a Hair-ball from Madagascar ............

Page

181

im)

xil

Hawkins, Herpert L., M.Sc., F.G.S., Lecturer in Geology,

University College, Reading.

The Anterior Ambulacrum of Hchinocardium cordatum

Penn., and the Origin of Compound Plates in Echinoids.

(Pl. XX VI/ and Text-figs. See US) ian be ome p dwOneenn aaa gene oe

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

Some Falkland Island Spiders. (Pls. I. & IT.) .........

Jameson, H. Lysrrer, M.A., D.Sec., Ph.D., F.Z.S., and

Nicoitz, Wiuuiam, M.A., D.Sc., M.D., F.Z.S.

On some Parasites of the Scoter Duck (Hdemia nigra),

and their Relation to the Pearl-inducing Trematode in

the Edible Mussel (Mytilus edulis). (Text-figs. 11 & 12.)

Karr, Sir Henry Seron-. See Seron-Karr, Sir Henry.

Larpiaw, Frank F., M.A., F.Z.S8.

Contributions to a Study of the Dragonfly Fauna of

Borneo.—Part I. The Corduliine: The Genus Amphi-

enenis: ‘he Wesion Protoneura, ‘(PI Ve) jiecean-- spe:

Lamporn, W. A., M.R.C.S., F.Z.S.

Notes on the Habits of Certain Reptiles in the Lagos

TORS CITE US ih cts aieaiveie avian aie ROSE eS dans se aioe lataeants aeeenn cea vaca

LyprEkKER, RicHarp, F.R.S., F.Z.S.

The Dwarf Buffalo of Southern Nigeria; with a

Revision of the Dwarf Buffaloes of Western Africa.

(Mexprhiess 42442) coseciccusas sm agemcnetonnitc ot esie tess delice aeaanee

MarsHati, Guy A. K., F.Z.S.

Remarks on the Relationship of the Big Game of

Africa to the Spread of Sleeping Sickness. See Yor«KE,

Dr. W.

169

218

234

xiii

Meruuen, The Hon. Paut A., F.Z.S.

Description of an Amphipod belonging to the Family

Talitride, from the Woodbush, Transvaal. (Pls. X. &

DRT eet eral cfc hE tei Yaoi d ehees ese bags asta ote sens ben porotia hid

Mincuin, Prof. Epwarp A., M.A., F.R.S., V.P.Z.S.

Remarks on the Relationship of the Big Game of

Africa to the Spread of Sleeping Sickness. See YorkKE,

Dr. W.

MircHett, P. Caaumers, M.A., D.Sc., LL.D., F.R.S., F.L.S.,

Secretary of the Society.

Report on the Additions to the Society’s Menagerie

cunins theamonthot October 19V2 airs. 4ne sce nossa ee

Exhibition of a photograph of the Hainan Gibbon

(Hylobates hainanus) taken in natural colours ............

Report on the Additions made to the Society’s

Menagerie during the months of November and December

TSSILO YG aa Se Me Soa Ne ENS SMR CR in i Si Pe OI Ns 34 Nel Cea

Report on the Additions made to the Society’s

Menagerie during the month of January 1913 ............

Exhibition of lantern-slides of the Tree-Hyrax

(Dendrovypnaxc dorsalis) arte e eon es eae eevee

Exhibition of a lantern-slide of an abnormally-marked

Domestic Donkey (Hqwus asinis)............ cece cence nee ees

Report on the Additions to the Society’s Menagerie

duninagsthemmonthvot Mebruany ONG i ejseacae- acces srr

Nicout, Wituiam, M.A., D.Sc., M.D., F.Z.8., and JAMESON,

A lnysrer. Meas Disc. Ph.D, E:Z:s:

On Some Parasites of the Scoter Duck (Hdemia nigra),

and their Relation to the Pearl-inducing Trematode in

the Edible Mussel (M/ytilus edulis). (Text-figs. 11, 12.).

109

Xiv

Puimmer, Henry G., F.R.S., F.Z.S., Pathologist to the

Society.

Report on the Deaths which occurred in the Zoological

Gardens during 1912, together with the Blood-Parasites

found) durimesbhie Mea lg yee eee set eee ne wae creee tee eee tat

Preston, Hueu B., F.Z.S.

Diagnoses of New Species and Varieties of Agnathous

Mollusca from Equatorial Africa. (Pls. XXXIT.—

RV ae seacctnte ct itotasse taioneap saan ncleu painter maepee a ene

Rea@an, C. Tate, M.A., F.Z.S.

Exhibition of male and female examples of the

remarkable Cyprinodont Fish, Cynolebias bellottw ......

RoruscHiLD, The Hon. L. Wattser, D.Sc., F.R.S., F.Z.S.

Letter on the Relationship of the Big Game of Africa

to the Spread of Sleeping Sickness. See YorKe, Dr. W.

Scuuster, Epaar H. J., M.A., D.Sc., F.Z.8., and Smiru,

GroFrrrey W., M.A., Fellows of New College, Oxford.

The Genus Hngewus, or the lLand-Crayfishes of

WAnistrallaaisy, «(else Xeb EXON Vike netsh acca) ia eee cana

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

Lantern demonstration of photographs of young

birds hatched in the Society’s Gardens during 1912,

and of the nest made by the Sumatran Orang-utan.

GENID Xa) ea whesnsastootbGdasons 6 suc iocHesods0asbds sddaseoaabouesne seed

Exhibition of a female Carolina Duck (Lampronessa

sponse) which had partially assumed male plumage ......

Seron-Karr, Sir Henry, K.C.M.G., F.Z.S.

Letter on the Relationship of the Big Game of Africa

to the Spread of Sleeping Sickness. See YorKE, Dr. W.

Page

194

151

SsARPE, Sir AtFRED, K.C.M.G., LL.D.

Remarks on the Relationship of the Big Game of

Africa to the Spread of Sleeping Sickness. See YorkKE,

Dr. W.

Smiru, Davip Seru-. See Seru-Smiru, D.

SMItH, GEOFFREY W., M.A., and Scuusrer, Epear H. J.,

M.A., D.Sc., F.Z.8., Fellows of New College, Oxford.

The Genus Hngewus, or the lLand-Crayfishes of

Australias: (Ess Xalil XeXGVe) ers vaca ceic asic eectestdacensisiesiae 112

Stirrup, H. H., B.Sc., (Birm.), Lecturer on Agricultural

Biology, East Anglian Institute of Agriculture,

Chelmsford.

A Descriptive Study of an Oligochete Worm of the

Family Enchytreide; with an Appendix on certain

Commensal Protozoa. (Pls. XLVI.—XLIX. and Text-

fi sPe Oi Oil, Ve ccts ae aisle cord sin sinse cosa durlvawclensuasiioniel se\cuibrnsatetierdace 300

Surron, Sir JoHn Buanp-. See BLAND-SutTton, Sir JoHN.

Tomas, OLDFIELD, F.R.S., F.Z.S.

Exhibition of a specimen of the Amazonian Monkey,

GG UCORSH CURIA GNU accens neces nesta Seon hee este clos eee cokes 3

Wititey, ArtHur, M.A., D.Sc, F.R.S., F.Z.S., McGill

University, Montreal.

Notes on Plankton collected across the mouth of the

St. Croix River opposite to the Biological Station at

St. Andrews, New Brunswick, in July and August 1912.

(ext tie si 4 Wry ol0))) ss cemtaaasioncicncie slsclersesienvareiaetisietistcl eitoee 283

XVI

Page

Yorke, Dr. W., Liverpool School of Tropical Medicine.

The Relationship of the Big Game of Africa to the

Spread of Sleeping Sickness. With an Appendix

containing Remarks by Sir Joun Branp-Sutron,

F.R.C.8., F.Z.8.; Guy A. K. Marsnatn, F.ZS. ;

Prof. E. A. Mincuin, M.A., F.R.S., V.P.Z.:8.; The

Hon. L. Water Roruscaritp, D.S8c., F.R.S., F.Z.S.;

Sir Henry Seron-Karr, K.C.M.G., F.Z.S.; and Sir

ALFRED SHARPE, K.C.M.G., LL.D.; and Reply by

Di YO PURGE: Byers orc ca.u si erers ts ss sigssejne ale Bia alee Le Re eee 321

XVil

NEW GENERIC TERMS

PROPOSED IN THE PRESENT VOLUME (pp. 1-337).

Page | Page

Arunella (Crustacea) ...........- 232 | Solenotznia (Vermes) ......... 243

Ophidotznia (Vermes) ......... 25 | Talitriator (Crustacea) ......... 109

Scylacops (Reptilia) .......:.... 226 |

Proc. Zoou. Soc.—1913. h

aie. eit nk aapaioensy

INDEX

SCIENTIFIC NAMES.

Acanthotznia, 11, 34,

35, 154, 156, 160,

168, 244, 247.

shipleyt, 5, 25.

tidswelli, 16.

varia, 36.

Acartia, 285.

clausi, 284, 285.

tonsa, 284.

Acomus

erythrophthalmus, 147.

Acridotheres

tristis, 147.

Actinotrocha

browne, 290.

/Kolosoma, 316.

Agama

colonorum, 218.

Agelzeus

icterocephalus, 148.

Akidoproctus, 133,

Akodon

arenicola, |.

Alloneura, 74.

Amblystoma

tigrinum, 2.

Amia, 83.

calva, 82.

Amphicnemis, 63, 78.

sp., 1d.

ecornuta, TA.

Jurcata, 74.

gracilis, 74.

lestoides, 'T4.

louise, 71, 74, 79.

madelene, 71, 74, 79.

martini, 72, 74.

remiger, 72, 74.

wallacei, 70-74, 79.

Ancistrodon

contortrix, 149.

piscivorus, 149, 164.

Ancistrona, 132,

136.

133, |

Ancistrona

gigas, 129, 134. 139.

procellarie, 129, 180,

133, 134.

Antedon

savignyt, 88, 102.

serripinna, 88.

Anthobothrium, 256, 258.

musteli, 257.

Antirrhinum, 301.

Aphelocoma

woodhousei, 147.

Araneus

globiger, 39, 50.

productus, 39.

wallentini, 37, 50.

Arboricola

crudigularis, 2.

Archeeosuchus

cairnerossi, 230.

Arctoenathus, 230.

Argulus

africanus, 263-265,

267, 275, 277-282.

angusticeps, 263, 267,

273, 280, 282, 283.

exiguus, 263, 267, 272,

280, 282.

foliaceus, 264.

incisus, 268, 267, 276,

280-282.

megalops, 273.

niger, 266.

personatus, 263, 267,

271, 280, 282, 283.

pugettensis, 266.

rubescens, 263,

276, 280-283.

rubropunctatus, 263,

267, 269, 278, 280-

282.

striatus, 263, 267, 269,

274, 277, 278, 280-,

283.

267,

Arunella, gen. nov., 282,

subsalsa, 232, 234.

Aspidorhynchus, 83.

Astacopsis, 113-116, 118,

126.

Astacus

fossor, 119.

(Engzus) fossor, 119.

Asteracanthion

pallidus, 286.

Asterias

forbesti, 287.

glacialis, 288.

rubens, 287.

vulgaris, 286, 287.

Asteroceras

pergamena, 88.

Athene

noctua, 147, 148.

Auchenoglanis

occidentalis tangani-

canus, 269, 276,

281.

Bagrus

degent, 265, 278.

Bairdia

subdeltoidea, 233.

Balearica

pavonina, 152.

Barbus

ahblabes, 51.

spurrelli, 51, 53.

Barilius

loati, 52.

macrostoma, 51, 53.

senegalensis, D2.

Batagur

baska, 149.

Bathy bates

ferox, 272.

Biddulphia, 284.

Bison

americanus, 152.

Bitis

gabonica, 153.

Boa, 25.

constrictor, 148.

madagascariensis, 152.

Bolina

alata, 288.

Boopia, 130, 183, 138.

Bos.

centralis, 235.

caffer, 240.

— heddingtont, 241.

— brachyceros, 280,

236, 238, 240,

241.

— centralis, 240.

—cottom, 236, 287,

240.

—hunti, 237, 239,

241.

—nanus, 236, 239-

241.

— planiceros, 235, 238,

240, 241.

— simpsoni, 240, 241.

— thierry, 240.

nanus, 23d.

planiceros, 235.

(Bubalus) caffer bed-

dingtont, 240.

(—) — hunti, 240.

Bothriocephalus, 31.

Bubalus

brachyceros, 235.

caffer, 1.

Bungarus

candidus, 149.

Caconeura, 80.

(Alloneura), 75.

Calabaria

reinhardti, 149,

Calamaria, 70.

Calamoichthys, 298.

Calanus

jfinmarchicus, 284.

Cailicebus, 4.

Callimico

snethlageri, 4,

Calliobothrium, 257, 258,

261.

coronatum, 256.

leuckarti, 256.

Calophasis

mikado, 242.

Candona, 232.

Cardium, 55.

edule, 54, 62.

Catatropis

verrucosa, 57.

Caturus, 83.

Cavia

rufescens, 1.

Centetes, 242.

Cheraps, 113, 114-116,

TLS), 125),

quadricarinatus, 115.

Cheeropsis

liberiensis, 242.

Cheetoceras, 285.

Chiltonia, 109.

Chlamydoselachus, 297.

Chloropsis

aurifrons, 147.

Chonopeltis

inermis, 263, 280.

Chrysichthys

brachynema, 276, 277,

281.

Ciconia

alba, 38.

Cinosternum

cruentatwm, 149.

Clarias

anguillaris, 265, 278.

lazera, 275, 278.

robeechit, 275, 278.

Clypeaster, 178.

Ceereba

cyanea, 148.

Ceetoplana, 289.

Colius

erythromelon, 147.

Colpocephalum, 133.

Coluber

helena, 242.

Comatula

multiradiata, 87, 88.

Connocheetes

gnu, 150.

Copsychus

saularis, 147.

Corallebothrium, 34.

Cordulephya, 65.

Coronella

getula, 149.

Corvultur

crasstrostris, 2.

Corvus

corax, 148.

Coryceeus, 181.

Corycella, 181.

Corynura

bumpusti, 285.

Coscinodiseus, 284.

Crepidobothrium, 34, 36.

gerrardt, 12, 25, 153.

Crocodilus

acutus, 224.

INDEX OF SCIENTIFIC NAMES.

Crocodilus

madagascariensis, 223.

niloticus, 224.

Croszarchus

obscurus, 150.

Crotalus

atrox, 149.

Cryptoprocta

JSeroz, 1.

Crypturus

cinnamomeus, 242.

Ctenoplana, 290.

korotneffi, 289.

Cyanops

Jlavifrons, 2.

Cyclemys

amboinensis, 149.

Cygnus

melanocoryphus, 3.

Oynolebias

bellottii, 3.

Cynosuchus, 230.

Dapedius, 83.

Dasyurotenia

robusta, 9.

Davainea, 10.

Dendraspis

viridis, 149.

Dendrobrna

subrubicunda, 308.

Dendrocitta

vagabunda, 147, 148.

Dendrohyrax

dorsalis, 152.

Diaptomus

pusillus, 231, 233.

Dinopterus

cunningtont, 275.

Disparoneura, 70, 74,

aurantiaca, 76.

collaris, (6.

delia, 75.

dorsalis, 79.

hosei, 76, 78.

humeralis, 76, 78, 80.

— nigra, 76.

hyperythra, 76.

wmtegra, 75,

interrupta, 76.

lansbergi, 76.

moultoni, 76.

obsoleta, 715.

peramena, 76, 79.

verticalis, 75-17.

Dissemurus

paradiseus, 147.

Dochophorus, 133,

icterodes, 133.

INDEX OF SCIENTIFIC NAMES.

Dochophorus

sphenophorus, 130, 133-

135, 140.

— Dolops

ranarum,

278-282.

(Gyropeltis) ranarum,

278.

263,

Duthiersia, 5.

Echeneibothrium, 257.

Eehinoeardium

cordatum, 169-173,

175-177, 180, 181.

flavescens, 173, 176.

Echinostomum

revolutum, 57.

Echinus, 171, 178.

Kehis

carinatus, 149.

Emberiza

fucata, 148.

Eimmenomma

talklandica, 45, 50.

Emys

orbicularis, 149.

Enchytreeus, 301.

albidus, 301, 302, 310, |

313, 320.

argeniteus, 301, 315.

buchholzii, 303.

hortensis, 308, 310.

mobi, 319.

pellucidus, 301, 302,

306-311, 313-316,

318, 320.

vermicularis, 316.

Engeeus, 112-116.

affinis, 117-122, 127.

cunicularius, 118, 119,

124-127.

fossor, 117-122, 126,

127.

fultoni, 117-119, 126.

hemicirratulus, 117—-

119, 123, 124. 127.

phyllocercus, 117-119,

122, 126, 127.

victoriensis, 117-119,

121, 122; 127.

Ennea

aberdarensis. 207, 217.

adelpha, 199, 217.

—, yar., 217.

adjacens, 214, 217.

arthuri, 198, 217.

haccata, 198, 217.

buccina, 196, 217.

burungaensis, 206, 217.

carea, 200, 217.

claustrum, 208, 217,

264, |

| Ennea

coent, 199, 217.

| commoda, 210.

consobrina, 216.

copiosa, 200, 217.

— robusta, 201.

curvicolumella, 201,

217.

decussatula, 201, 217.

disseminata, 202, 217.

elgonensis, 197, 217.

eussoensis, 202, 217.

fortidentata, 199, 200.

foveolata, 208, 217.

funerea, 203, 217.

—- levis, 203.

hector, 203, 217.

ingeziensis, 204, 217.

— mhararaensis, 205.

— pusilla, 204.

ingloria, 197, 217.

tunocens, 197, 217.

insula, 205, 217.

intradentata, 205, 217.

— curta, 206.

gombeneénsis, 213, 217.

keniana, 207.

— intermedia, 206.

— parvula, 206.

kigeziensis, 207, 217.

kivuensis, 197, 217.

laqueus, 207, 208, 217.

— hercules, 208.

lima, 208, 217.

malasangiensis, 209.

margarita, 196, 197,

217.

masakaensis, 209, 217.

mikenoensis, 209, 217.

mirifica, 214, 217.

monticola, 210, 217.

mutandaensis, 211, 217.

mweruensis, 210, 217.

nairashaensis, 211.

— elgonensis, 211.

nyikaensis, 199, 200.

nyiroensis, 211, 218.

optata majuscula, 212,

218

— ohesa, 211, 212, 218.

papyracea, 212, 218.

percivali, 216, 218.

pergrata, 212, 218.

perturbata, 199, 218.

pervitrea, 213, 218.

pilula, 213.

pollonere, 213, 218.

pretiosa, 214.

— nyiroensis, 214.

tridescens, 205, 206, 217.

xxi

Hnnea

rectangularis, 197, 218.

reniformis, 215. 218.

spatium, 214, 218.

suavissina, 215, 216,

218

syngenes, 216.

viatoris, 216, 218.

vieina, 199.

woodhousei, 199, 218.

(Ptychotrema), —om-

haensis, 216, 217.

Emnneacanthus

gloriosus, 242.

Epicrates

cenchris, 149,

Epophthalmia

australis, 64, 69.

vittigera, 64, 70.

Equus

asinus, 241.

Kriculus

setosus, 152.

Hrigonocephalus

pisctvorus, 166.

Eriphostoma, 230.

Hryx

johni, 149.

Kugnathus, 83.

Kureum, 133.

Eurymetopus, 133.

Kurytemora

americana, 285.

herdmani, 284, 285.

hirundoides, 285.

Kutrichophilus, 183.

Evadne

nordmanni, 285.

| Felis

nehulosa, 146.

pajeros, 1.

viverrina, 242.

Foudia

madagascariensis, 147,

148.

Fridericia

bisetosa, 301.

Fritillaria

borealis, 284, 285.

Fuligula

affinis, 242.

collaris, 242.

terina, 56.

Fundulus

gardneri, 53.

spurrelli, 52.

Galepus

Joubertt, 226.

Gallus

gallus, 147, 148.

Garrulax

albigularis, 146, 148.

lanceolatus, 146.

Gazella

isabella, 242.

subgutturosa, 242.

Gerrhosaurus

nigrolineatus, 149.

Giebelia, 133.

Glareola

pratincola, 148.

Gliricola, 133.

Glossina

morsitans, 322-3026,

332, 334, 386, 337.

palpalis, 321, 322, 324,

331, 332, 334.

Gomphomacromia

paradoxa, 66.

Goniocotes, 133.

Goniodes, 133.

dissimilis, 139.

falcicornis, 128, 129.

tetraonis, 128.

Gorgonocephalus

eucnemis, 92,

Gorgonops

torvus, 225-227, 230.

Gubernatrix

cristata, 147.

Gymnophallus, 56.

affinis, 57-62.

bursicola, 55, 57.

dapsilis, 55, 57.

macroporus, 57, 60, 61.

edemie, 55, 57, 58,

62.

ovoplenus, 57, 62.

somaterte, Dd.

(Lecithodendrium) so-

materia, do.

Gymnorhis

flavicollis, 148.

Gyrodus, 83.

Gyropeltis

ranarum, 263, 264.

Gyropus, 130, 133.

Hemoproteus

danilewskyi, 146, 148.

Haplochilus

danganicanus, 273.

Hemiaster, 175.

Hemichrowmis

bimaculatus, 2.

Hemicirratulus

hystrix, 124.

INDEX OF SCIENTIFIC NAMES.

Hemicordulia

assimilis, 64.

Hemipedina, 177.

Hesperocordulia, 65.

Heteraster

oblongus, 175, 180,

181.

Heterobranchus

bidorsalis, 265, 279.

Heterodoxus, 130, 133.

macropus, 138.

Heterolepidotus, 83.

Hierofalco

islandus, 152.

Huhua

nipalensis, 151.

Hyalella, 109.

Hyla

arborea, 151.

Hylobates

hainanus, 2.

Hylocichla

guttata pallasi, 147.

Hyphantornis

textor, 147, 148.

Ichthyotenia, 4-6, 10,

36, 249, 253, 257.

sp., 26, 164, 166,

167.

abscisa, 249.

birot, 11, 138-16, 19,

25.

calmettei, 17, 27, 168,

247.

fossata, 245.

gabonica.

159-162,

254, 255.

gracilis, \1, 14, BBD.

lonnbergi, 5.

marenzelleri, 14, 166,

168.

natterert, 14, 19, 26,

27, 29, 35, 160, 249

nilotica, 11, 12, 15, 16,

20, 22, 26, 31.

ocellata, 245.

racemosa, 160.

saccifera, 22, 25.

schulizet, d.

didswelli, 7, 9, 11, 24,

153 - 157,

168, 243,

varia, 11, 15, 17, 20,

22-95, 31, 33, 34.

(Acanthotznia), 5.

(—) sp., 8, 9.

(—) nilotica, 138.

(—) tidswelli, 5, 7.

(—) varia, 18, 20, 21,

»)

Icterus

gularis, 146.

melanocephalus, 2.

Tetido-aurus, 230.

Idionyx

claudia, 80.

dohrii, 66-68, 79.

— horneensis, 64, 67,

68, 80.

Iguana

tuberculata, 149.

Inostransewia, 230.

Kelloggia, 133.

Lacerta

ocellata, 149.

Lachesis

alternans, 248.

mutus, 151.

Lemobothrium, 132, 133,

141.

gypsis, 129, 130, 139.

titan, 129, 130, 1389,

140.

Lampronessa

sponsa, 151.

Larus

hemprichi, 3.

Lates

microlepts,

278, 281.

Latumcephalum,

133.

Lecithodendrium

somaterié, 57.

Lemur

varius, 150.

Leontocebus

adipus, 146.

Lepidosteus, 81, 83, 85.

osseus, 82.

Lepidotus, 83.

Leptolepis, 83.

Levinseniella

brachysoma, 57.

Lipewus, 132, 133.

Jerox, 129, 131.

longipilus, 133.

picturatus, 133.

Loriculus

galgulus, 147, 148.

Loxia

curvirostra, 147, 148.

Lycosaurus, 250.

Lycosuchus, 226.

264, 271,

129,

Macacus

inuus, 152.

Macromia, 66.

sp., 64, 68.

borneensis, 64, 68, 69.

cincta, 64, 68, 69.

cingulata, 64, 68, 69.

gerstaeckeri, 64, 68, 69.

westwoodi, 64, 68, 69.

Macrosemius, 83.

Melophus

. melanicterus, 147, 148.

Melopyrrha

- nigra, 152.

Menopon, 133, 140.

distinctum, 188.

malleus, 133.

pallidum, 141.

persignatum, 133.

precursor, 133.

rediculosum, 133.

robustum, 133.

titan, 133, 186, 140.

tridens, 133, 136, 140.

Merula

castanea, 147.

. infuscata, 242.

Mesenchytreus,

315.

Mesia

- argentauris, 148.

Mesnilella, 317, 318.

fastigiata, 316, 320.

Mesocestoides, 36.

litterata, 33.

Mesturus, 83.

Metaphya, 66, 67.

micans, 64, 65, 79.

Metorchis

xanthosomus, 56, 57.

Mico, 4.

Micraster, 175.

Microceccus

melitensis, 328.

Midas

goeldit, 4.

Mimus

modulator, 1.

Misumena, 48.

Monostomum

sp., 07.

Mungos

ichneumon, 152.

Mytilus, 54-56.

edulis, 53.

Myzostoma

asterie@, 92, 94, 96, 98,

108.

belli, 101.

carpenteri, 100.

circinatum, 106.

cirriferum, 87, 90, 99,

306,

- INDEX OF SCIENTIFIC NAMES.

Myzostoma

costatum, 86-100, 102,

103, 105, 106, 108.

crosslandi, 102-104,

106-108.

eryptopodium, 101i.

elongatum, 105.

Soliwm, 102, 103, 107.

giganteum, 95.

glabrum, 87, 90, 98,

100.

graffi, 95.

japonicum, 88.

mebianum, 96.

nansent, 102, 108.

rubrofasciatum,

101, 102, 108.

86,

Naia

bungarus, 149.

tripudians, 25, 149.

Necturus, 5.

Nirmus, 133.

cameratus, 128.

signatus, 133.

Nitzschia, 1383, 141.

pulicaris, 129, 137.

Nyctomys

sp., 146.

Octochextus

mulliporus, 303.

Qidemia

migra, 03, 56, 57.

(dura

ocellata, 242.

(noscopus, 83, 84.

Oithona, 181.

atlauntica, 191.

attenuata, 182, 187,

191, 193.

brevicornis, 191.

challengeri, 192.

decipiens, 182, 184,

186, 191, 193.

fallax, 182, 185, 191,

193.

Frigtda, 191.

hebes, 190, 191.

helgolandica, 192.

linearis, 182, 183, 191.

minuta, 192.

nana, 182, 186, 187,

189, 191.

oculatu, 182, 188, 189,

192, 193

pelagica, 182, 183.

plumifera, 182,

189, 191.

183,

XX1ll

Oithona

pygmea, 192.

rigida, 189, 192.

robusta, 182,

191.

setigera, 182, 183, 189,

191,

184,

similis, 185; 186, 191.

simplex, 182, 187, 191,

spinifrons, 192.

spinirostris, 192.

tropica, 182, 183.

vivida, 182-184, 191,

193.

Oligopleurus, 81, 83.

esocinus, 84.

vectensis, 84.

Oachobothrium

uncinatum, 256.

Onchophorus, 133.

Oochoristica, 6, 153.

Ophiacantha

vivipara, 88.

Ophidotzxnia, gen. nov.,

25, 35, 36, 153, 156,

243, 247, 252, 254,

255, 257, 259, 261.

nate, 25, 28, 30, 32,

153, 154, 558, 159,

161-164, 253.

russelli, 160, 161, 163, °

165, 168, 258.

Ophiopsis, 83.

Opisthostoma, 70.

Ornicholax, 133.

Ornithobius, 133.

Ornithorhynchus, 227,

Osteolepis, 84.

‘Oxygastra, 65.

Pachycormus, 83.

Pachydrilus, 316,

319.

Palzoniscus, 84.

Palzornis

Jasciatus, 148.

Palaia

varami, §, 6, 156.

Panceria, 6.

Pancerina, 3, 6.

Papio

anuhis, 152.

hamadryas, 152.

maimon, 152.

Parachzraps, 114, 118,

126.

bicarinatus, 118, 115,

116.

XX1V

Paramonostomum

alveatum, 57.

Paroithona, 181.

parvula, 190-192.

pulla, 182, 190, 192,

193.

Parotia

lawesi, 146.

Passer

arcuata, 148.

luteus, 147.

Penthetria

laticauda, 148.

Peridinium

divergens, 285,

286.

Petricus

niveus, 48.

signatus, 46, 50.

Philisea

colulata, 42, 50.

navarinensis, 44.

oculatum, 46.

Philodryas

schotti, 149.

Phlogeenas

helviveniris, 2.

Pholidophorus, 83.

Phoronis, 290-292.

brownet, 292.

sabatiert, 291.

Phryniscus

negricans, 242.

Phyllobothrium

gracile, 257.

Physostomum, 133.

sp., 139.

mystax, 139.

Pithecia

monachus, 150.

Pituophis

sayt, 147, 149.

Plasmodiuin

precox, 146, 148.

Platalea

leucorodia, 133.

Platycnemis, 7V.

Platysticta

quadrata, 79.

rufostigma, 79.

Pleurobrachia, 289.

Plocepasser

mahali, 150.

Podon

polyphemoides, 285.

Poéphila

gouldie, 147.

Polypterus, 85, 292, 294,

295, 297-300.

lapradii, 293.

senegalus, 298.

Pratincola

caprata, 148.

Procellaria

capensis, 133.

Propasser

rhodochrous. 150.

Proteocephalus, 258.

singularis, 259.

Protomyzostomum

polynephris, 92, 96,

101, 107.

Protopterus

ethiopicus, 241, 265,

278. .

Protosticta

versicolor, 78.

Psammophis

leithii, 2.

Pseudomenopon, 133.

tridens, 136, 140.

Psilochasmus

oxyurus, 57.

Psilostomum

brevicolle, 57.

Pternistes

swainsont, 148,

Ptilopus

tozonus, 2.

wallacet, 2.

zonurus, 2.

Ptychobothrium

belones, 251.

Pygaster, 178.

semisulcatus, 179.

Pygopus

lepidopus, 181.

Pyrodinium

bahamense, 286.

Python

molurus, 148.

Quiscalus®

purpureus, 147.

Ramphastos

dicotlorus, 146.

Rana

adspersa, 5.

esculenta, 147, 151.

Reithrodon

typicus, 1.

Rhinoceros

unicornis, 2.

Rhizosolenia, 285.

Rhyothemis, 66.

Sadocus

vallentini, 48, 50:

vitellinosulcatus, 50.

INDEX OF SCIENTIFIC NAMES.

Saxicava

rugosa, 59.

Scapanodon

duplessisi, 230.

Schizaster, 175, 181.

Scops

giu, 147, 148.

Scylacognathus, 228.

parvus, 227, 230.

Seylacops, gen. nov., 226,

228, 229.

capensis, 226, 227, 230.

Scylacosaurus, 226.

Seymnognathus, 228.

tigriceps, 227, 230.

whattsi, 225, 230.

Scyphocephalus, 5.

Sialia

sialia, 146, 147.

Simia

satyrus, 3.

Simochroinis

diagramma, 273.

| Solenophorus, 153.

Solenotenia, gen. nov.,

243, 247, 249, 251,

254, 257-259.

viperis, 244-246, 248,

250, 252, 253, 255,

260, 261.

Somateria

moallissima, 5b.

Somatochlora, 65.

| Spathiurus, 83, 84.

| Spelotrema

pygmeum, 57.

Speotyto

cunicularia hypogea,

147.

Sporophila

superciliaris, 242.

| Streptazis

desiderata, 195, 217.

kirkii, 194.

marsabitensis, 195, 217.

percivali, 194, 217.

urguessensis, 195, 217.

woodhousei, 194, 217.

Strigea

tarda, 57.

Struthio

australis, 152,

Sturnella

defilippi, 148.

Sycalis

pelzeini, 1.

| Syncheeta, 285.

Teenia

filicollis, 249, 257.

Tsenia

ocellata, 257.

racemosa, 160.

torelosa, 257.

varant, ©.

Talitriater,

109.

eastwoode, 110, 112.

Talitrus, 118.

locusta, 110,

sylvaticus, 109.

Tapes, 55.

decussatus, 54.

Tarbophis

fallax, 149.

Teinobasis

rajah, 73.

Testudo

angulata, 149,

emys, 149.

Tetrabothrium, 12, 36.

trionychium, 5, 34.

Tetragnatha

ensulata, 41, 50.

Tetrophtalmus, 153.

titan, 135, 136, 139,

140.

Tharrhaleus

jerdont, 148.

Thereiceryx

zeylonicus, 152.

Thrissops, 83.

Titanosuchus, 225.

cloetet, 230.

ferox, 230.

gen. noy.,

INDEX OF SCIENTIFIC NAMES.

Tjalfiella

tristoma, 289.

Tocotrema

concavum, 56, 57.

Tortanus

discaudatus, 284, 235.

setacaudatus, 285.

Toxostoma

cinereum, 147, 148.

Trichodectes, 133. |

Trigonocephalus

pescivorus, 166.

Trimenopon, 133.

Trinoton, 129, 130, 153,

136, 141,

luridum, 136, 139.

Trionyx

ferox, 36.

Tropidonotus

piseator, 149.

Trypanosoma

brucei, 329, 331.

cazalboni, 329.

cruzi, 331.

gambiense, 329-331.

nanum, 32.

pecorum, 325.

rhodesiense, 322,

331, 335.

theilert, 331.

vivax, 325,

Tupinambis

tequexin, 149,

Turdus

albiventris, 147.

329-

Proc. Zoou, Soc.—1913, No. XXIII.

XXV

Uvobrachya

dbonotata, 146, 147.

Urocissa

oecipitalis, 146.

Varanus, 8, 26, 27, 30,

35, 36, 168.

arenarius, 6.

indices, 5.

niloticus, 6,

salvator, 25.

varius, 5, 7, 10, 16, 24,

34.

Vipera

ammodytes, 149.

russelli, 149, 160.

li,

Xanthura

luxuosus, 147.

Zaglossus

nigroaculeatus?, LA0.

Zamenis

gemonensis, 149,

mucosus, 149.

viridiflavus, 153.

5! f4

i ocr gia é

Ale WER

OA aula

DEX

~[LLUSTRATIONS.

Acanthotenia sp., Figs. 1, 2, pp. 8, 9.

nilotica, Vig. 3, p. 13.

varia, Figs. 4—7, pp. 20-22.

Amia calva, Fig. 15, p. 82.

Amphicnemis louise, Pl. 1V. p. 63.

madelene, Pl. LV. p. 63.

-—— wallacei, Pl. LV. p. 63.

Ancistrona procellarie, Figs. 25, 26,

pp- 182, 134.

Araneus globiger, Pl. I. p. 87.

vallentint, Pl. I. p. 37.

Argulus angusticeps, P\s. XLII., XLIV.,

p.- 262.

-—— exiguus, Pls. XLII., XUIIL.,

p- 262.

imceisus, Pls. XLI., XUIIT., p. 262.

—— wpersonatus, Pls. XLI., XLIV.,

p. 262.

rubescens, Pls. XLII., XLV.,

p. 262.

rubropunctatus, Pls. XLI., XLILI.,

p- 262.

Surdcnos, ells OSI0INIL,, = RDA e,

p. 262.

Arunella subsalsa, Pls. XXXIX., XL.,

p. 231.

Astacopsis, Pl. XII. p. 112.

Barbus spurrelli, Pl, 111. p. 51.

Barilius macrostoma, Pl. 111. p. 51.

Bos caffer cottoni, Fig. 42, p. 287.

hunti, Figs. 43, 44, pp. 237,

239.

Brachiolaria attached to seaweed.

Fig. 54, p. 287.

Diaptomus pusilius, Pl. XXXVIII.

p. 281.

Disparoneura peramena, P\. LV. p. 63.

Docophorus sphenophorus, Fig. 27,

p. 135.

Echinocardium cordatum, Pl. XXVI.

p. 169; Figs. 39, 40, pp. 172, 173.

Enmenomma falklandica, Pl. I. p. &7.

Enchytreus albidus, Pl. XLVI. p. 300.

— argenteus, Fig. 67, p. 313.

pelluctdus, Pls. XLVI.—XLIX.,

p- 800; Figs. 61-66, pp. 806-311.

Engeus affinis, Pls. XVII., XVIIL.,

p- 112.

cumcwlarius, Pls. XIII., XXII.-

2.OMon JO LIZ

-—— fossor, Pls. XIV.-XVIL., p. 112.

hemicirratulus, Pls. XX.—XXIT.,

p. 112.

-—— phyllocercus, Pls. XI1X., XXI.,

p- 112.

victoriensis, Pls. XVIII.-XX.,

p. 112.

Ennea aberdarensis, Pl, XXXII. p. 194.

adelpha, P\, XXXII. p. 194.

, var., Pl. XXXIV. p. 194.

adjacens, Pl, XXXII. p. 194.

arthurt, Pl. XXXII. p. 194.

haccata, Pl. XXXII. p. 194.

buccina, Pl. XXXII. p. 194.

—— burungaensis, Pl. XXXII. p. 194.

carea, Pl. XXXIT. p. 194.

claustrum, Pl. XXXII. p. 194.

coent, Pl. XXXII. p. 194.

—— copiosa, Pl. XXXII. p. 194.

curvicolumella, Pl. XXXII. p. 194.

decussatula, Pl. XXXIIT. p. 194.

—— disseminata, Pl. XXXIII. p. 194.

—— elgonensis, Pl. XX XIII. p. 194.

eussoensis, Pl. XXXIII. p. 194.

Joveolata, Pl. XXXIII. p. 194.

Sunerea, Pl. XXXIIL. p. 194.

hector, Pl. XX XIII. p. 194.

ingeziensis, Pl. XXXIII. p. 194,

ingloria, Pl. XXXIIT. p. 194.

tnnocens, Pl. XXXIITI. p. 194.

insulsa, Pl. XXXTIT. p. 194.

intradentata, Pl. XXXIII. p. 194.

iridescens, Pl. XXXITII. p. 194.

jombeneénsis, Pl. XXXIIT. p. 194,

—— kigeziensis, Pl. XXXIV. p. 194.

XXVill

Ennea kivuensis, Pl. XXXIV. p. 194.

laqueus, Pl. XXXIV. p. 194.

— lima, Pl. XXXIV. p. 194.

malasangiensis, Pl. XXXIV.

p- 194.

margarita, Pl, XXXIV. p. 194.

——— masakaensis, Pl. XXXIV. p. 194.

mikenoensis, Pl, XXXIV. p. 194.

mirifica, Pl. XXXIV. p. 194.

monticola, Pl. XXXIV. p. 194.

mutandaensis, Pl. XXXIV. p. 194.

mweruensis, Pl, XXXIV. p. 194.

—— nyiroensis, Pl. XXXV. p. 194.

p. 194.

——, var. obesa, Pl. XXXV.

p. 194.

papyracea, Pl. XXXYV. p. 194.

percivali, Pl. XXXV. p. 194.

pergrata, Pl. XXXV., p. 194.

perturbata, Pl. XXXV. p. 194.

—— pervitrea, Pl. XXXV. p. 194.

—— pollonere, Pl. XXXYV. p. 194.

rectangularis, Pl. XXXY. p. 194.

—— renifornvis, Pl. XXXY. p. i94.

—— spatium, Pl. XXXV. p. 194.

—— suavissima, Pl. XXXYV. p. 194.

—- viatoris, Pl. XX XV. p. 194.

—— woodhousei, Pl. XXXV. p. 194.

(Ptychotrema) kombaensis, Pl.

XXXIV. p. 194.

Gymnophallus affinis, Fig. 11, p. 59.

macroporus, Fig. 12, p. 61.

Heterodoxus macropus, Fig. 31, p. 138.

Ichthyotenia sp., Figs. 37, 38, pp. 166,

167.

gabonica, Figs. 33, 34, pp. 155,

157.

— (Acanthotenia) sp., Figs. 1, 2,

pp. 8, 9.

( ) nilotica, Fig. 3, p. 13.

( ) varia, Figs. 4-7, pp. 20-22.

Idionyx dohrni, Pl. 1V. p. 63.

Lemobothrium sp., Fig. 25, p. 132.

Lepidosteoid bone, Diagram of struc-

ture and growth of. Fig. 13, p. 81.

Lepidosteus osseus, Fig. 14, p. 82.

Lipeurus ferox, Figs. 24, 25, pp. 131,

132.

Mesnilella fastigiata, Pls.

XLVIL., p. 500.

Metaphya micans, Pl. IV. p. 63.

XLVI,

optata, var. majuscula, Pl. XXXV.

INDEX OF ILLUSTRATIONS.

Myzostoma costatum, Pls. V., VIIL.,

p. 85; Figs. 17-22, pp. 89, 91, 94,

95, 97, 99.

crosslandi, Pls. VI.-VIIL., p. 89 ;

Fig. 23, p. 104.

rubrofasciatwm, P|, VI. p. 85.

Nest made by Orang-utan, Pl. TX. p. 3.

Nitzschia pulicaris, Fig. 80, p. 137.

Oithona attenuata, Pl, XXX. p. 181.

decipiens, P|. XXVIII. p. 181.

fallax, Pls. XXVII., XXVIIL,,

p. 181.

oculata, Pls. XXX., XXXLI.,

p. 181.

simplex, Pls. XXIX., XXX.,

p. 181

vivida, Pl. XXVIII. p. 181.

Ophidoteniu naie, Figs. 8-10, pp. 28,

30, 32.

russelli, Figs. 35, 36, pp. 163, 165.

Ophiopsis, Fig. 16, p. 83.

Paracheraps, Pl. XII. p. 112.

Paroithona pulla, Pls. XX1X., XXXT.,

p- L81.

Petricus signatus, Pl. I. p. 37.

Philisca colulata, Pl. U1. p. 37.

Phoronis (juy.), Fig. 55, p. 291.

Physostomum sp., Fig. 32, p. 139.

Platysticta rufostigma, Pl. LV. p. 65.

Polypterus, Figs. 56-60, pp. 294, 295,

297-299. $

Pygaster semisulcatus, Fig. 41, p. 179.

Sadocus vallentini, Pl. II. p. 37.

Scylacops capensis, Pl. XXXVI. }

Scymnognathus tigriceps, Pl. XXX VII.

p- 225.

Simia satyrus, Pl. IX. p. 3.

Solenotenia viperis, Figs. 45-58,

pp. 244-246, 248, 250, 252, 253, 250,

260.

Streptaxis

p. 194.

marsabitensis, P]. XXXII. p 194.

percivalt, Pl. XXXII. p. 194.

urguessensis, Pl, XXXIT. p. 194.

—— woodhousei, Pl. XXXII. p. 194.

desiderata, Pl. XXXII.

Talitriator

p- 109.

Tetragnatha insulata, Pl. 11. p. 37.

Tetrophthalmus titan, Fig. 28, p. 133.

Trinoton luridum, Fig. 29, p. 136.

eastwoodé, Pls. X., XI.,

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET,

PROCEEDINGS

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1913.:.

PART I.

CONTAINING Paces 1 To 152, witH 25 Puaves

AND 82 TEXT-FIGURES.

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LIST OF CONTENTS:

1913, Part I. (pp. 1-152).

EXHIBITIONS AND NOTICES.

Page

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

of October 1912 ...... reeves tee cite A SPU Para ACEP Mim sys Serie rei sigee

The Secretary. Exhibition of a natural colour photograph of the Hainan Gibbon (Aylobates

RQUIUCIUUS) evapo nstels eels (s helenche = Rr cackate ec aleust Reon elevates ae svesd late (sooo eve ReeOare MERON 2

Mr. G. A. Bovtencer, F.R.S., F.Z.8. Exhibition of a large specimen of the African

Cichlid Perch (Hemichromis bimaculatus Gill) ........+ alg gniaie eae

Mr. Epwarp Grrrarp. Exhibition of the skull of an Indian Rhinoceros (Ahinoceros

unicornis) which had recently died in the Society’s Gardens -..+.+.-eeeeceseseeees 2

Mr. E. G. Bovrrnerr, F.Z.S. Exhibition of a living specimen of the Salamander,

Amblystoma tigrimum vs sevecaccecccrsseeeecrnseterernctgs oan ae agehe eee

Mr. D. Szru-Sarta, F.Z.S. Lantern demonstration of photographs of young birds hatched

in the Society’s Gardens during 1912, and of the nest made by the Sumatran Orang- :

ThehBie alls: D-@)\>otosinn eng REE Toone Od GEOR AN aR a oes oko. co: +. ee

Mr. C. Tare Reean, M.A., F.Z.8. Exhibition of the remarkable Cyprinodont Fish,

Cynolebias bellottti .... +++ fest ers are Girne nin o\hetetteraiote sin tetaneiage ee + acne oPepeia eels wales e)

Mr. Oxpriztp Tuomas, F.R.S., F.Z.S8. Exhibition of an Amazonia:’ Monkey, Callimico :

SNCERIAGETE ..22-esereres ten eceeeres Sofeliens ioe aijaie ein ele fies sere cleinte seveaelenn's Coleeaente ee

The Secretary. Report on the Additions to the Society’s Menagerie during the months

~ of November and December 1912.......02. 0... scence eee cece eee cece eens és 150

Mr, D. Srrn-Saurn, F.Z.S. Exhibition of a female Carolina Duck (Lampronessa sponsa)

assuming male plumage ..++..eeeeee cere AUDDOOOODGUGHMAGCOUDENOnOm bo 0occkc 151

Mr. E. G. Bovrznerr, F.Z.S. Exhibition of a Blue specimen of the Edible Frog a

(Rana esculenta), and of the Australian Lizard (Pygopus lepidopus) .+...+0+.1++++s : 151

The Sucrutary. Report on the Additions to the Society’s Menagerie during the month of — :

January 1913 evceceesoeoee cece eevee ectrc tere eens ce serecese CPce te rn eter eeesetesece 151

Mr. E. G. Bourrnerr, F.Z.8. Exhibition of spines of the Hedgehog (Hriculus), taken from a

the excrement of a Boa madagascariensis

Contents continued on page 3 of Wrapper

“i

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

EXHIBITIONS AND NOTICES.

November 12, 1912.

Dr. A. Smita Woopwarp, F.R.S., Vice-President,

in the Chair.

The Secrerary read the following report on the Additions that

had been made to the Society’s Menagerie during the month of

October, 1°12.

The registered additions to the Society’s Menagerie during the

month of October were 259 in number. Of these, 120 were

acquired by presentation, 64 by purchase, 31 were received on

deposit, 33 in exchange, and 11 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 262.

Amongst the additions special attention may be directed to :—

1 African Buffalo (Bubalus caffer) 9 , from the Sezibwa Swamp,

Uganda, presented by H. M. Birch, Esq., on October 22nd.

1 Fossa (Cryptoprocta ferox) 3, from Madagascar, purchased

on October 23rd.

1 Pampas Cat (felis pajeros), 1 Pampas Cavy (Cavia rufescens),

3 Pampas Gerbilles (Retthrodon typicus), 2 Pampas Field-Mice

(Akodon arenicola), 1 Argentine Mocking-Bird (ddimus modu-

lator), and 3 Pelzeln’s Saffron-Finches (Sycalis pelzelni), from

- Argentina, the five last-named new to the Collection, presented

by Wilfred A, Smithers, Hsq., on October 16th.

Proc. Zoot. Soc.—1913, No. I. 1

i?)

MR. E. G@. BOULENGER ON A SALAMANDER.

1 Black-headed Hangnest (Jcterws melanocephalus), from.

Mexico, new to the Collection, presented by W. O. Danckwerts,

Esq., K.C., F.Z.8., on October 10th.

2 Great-billed Ravens (Corvultur crassirostris), from N.E.

Africa, new to the Collection, presented by the Marquess of

Tavistock, F.Z.S., on October 3rd.

1 Yellow-fronted Barbet (Cyanops flavifrons), from Ceylon,

new to the Collection, purchased on October 17th.

2 Orange-bellied Fruit-Pigeons (Ptilopus iozonus), 2 Pearl-

spotted Fruit-Pigeons (Ptilopus zonurus), 1 Wallace’s Fruit-

Pigeon (Ptilopus wallacer), and 1 Yellow-bellied Ground-Pigeon

(Phlogenas helviventris), from the Aru Islands, all new to the

Collection, purchased on October 15th.

4 White-throated Tree-Partridges (Arboricola crudigularis),

from Formosa, new to the Collection, received on deposit on

October 23rd.

3 Leith’s Snakes (Psammophis leithii), from 8.W. Asia, new to

the Collection, purchased on October 15th.

The Secretary exhibited a photograph of the Hainan Gibbon

(Hylobates hainanus) in natural colours, taken by Messrs. Elliott

& Fry, by the Lumiere process.

Mr. G. A. Boutenesr, F.R.S., F.Z.8., exhibited a specimen,

155 mm. long, of the African Cichlid Perch (Hemichromis bimacu-

latus Gill), which had recently died in Capt. Vipan’s aquarium.

The specimen was remarkable for its large size. Although the

species is common over a considerable part of Africa, and hundreds

of specimens have been collected in various localities, no wild

example is known.to exceed a length of 100 mm.

This Hemichromis is tinged with bright red about the head

and on the fins, and, as was noticed in the ‘ Fishes of the Nile,’

p- 462, this red is soluble in spirit, which, a few hours after the

immersion of the fish, acquires a bright orange-red colour.

Mr. Epwarp Gerrrarp exhibited the skull of an Indian

Rhinoceros (Rhinoceros unicornis) which had recently died in

the Society's Gardens. He pointed out that abscesses had been

formed at the base of the lower incisors, which had been much

rubbed down so that the nerves had become exposed. The

animal must have suffered great pain, and this no doubt was the

cause of his dashing his head against the walls and bars of his

den in the violent manner he at times did.

My. E. G. Boutencer, F.Z.S., Curator of Reptiles, exhibited a

living specimen of the Salamander, Amblystoma tigrinum, which

Ie, 4. &; ISIS. Pl, Wx,

oes

¥.

=a/

Photo. D. Seth-Smith.

NEST MADE BY ORANG-UTAN.

MR. OLDFIELD THOMAS ON AN AMAZONIAN MONKEY. 3

he had obtained from an Axolotl placed in special conditions

with the object of inducing the metamorphosis.

Mr. D. Sers-Siru, F.Z.8., Curator of Birds, gave a lantern

demonstration of photographs of the young of Hemprich’s Gull

(Larus hemprichi), of the Black-necked Swan (Cygnus melano-

coryphus), and of the White Stork (Ciconia alba) which had been

hatched in the Society’s Gardens during the year.

Three specimens of Larus hemprichi had lived in the Gardens

since 1906, but had shown no inclination to breed until the

summer of 1912. In July a pair kept in the Great Aviary

constructed a nest, composed of a few bits of grass and leaves:

under a bush, laying two eggs of a pale greenish-grey colour,

spotted and blotched with bluish grey and olive-brown. As there:

were birds in the aviary which would have devoured the young,,

the eggs were taken when just about to hatch and placed under:

a bantam. Both hatched, but one died on leaving the shell.

The other was successfully raised to maturity. The colour of the:

down was pale sandy-whitish, without any definite markings,

though the back was of a darker shade than the underparts. In

the first plumage it was of a buffish-brown colour, the feathers:

having paler edges.

The Black-necked Swan had not bred in the Gardens for over

thirty years until the past summer, when two cygnets were

hatched, one of which was successfully reared to maturity. The

eygnets, which are pure white in colour when hatched, have a

second down plumage of a dirty buftish-brown colour.

The White Storks, kept in the large flying aviary with the

Seagulls, had hatched five young birds, which unfortunately died

when from two to three weeks old.

Mr. Seth-Smith also showed slides of the nest, in a tree near

the Apes’ House, made by the large Sumatran Orang-utan (Simia

satyrus) which escaped from its cage on November 3rd, 1912

(Plate IX.). Mr. R. I. Pocock gave an account of the escape and.

subsequent capture of the Orang-utan.

November 26, 1912.

Dr. A. Smirn Woopwarb, F.R.8., Vice-President,

in the Chair.

Mr. C. Tare Recan, M.A., F.Z.S., exhibited male and female

examples of Cynolebias bellottii, a Cyprinodont Fish from the

La Plata, to illustrate the remarkable sexual characters.

Mr. Ouprietp Tuomas, F.R.S8., F.Z.5., exhibited a specimen of

an Amazonian monkey referable to a species he had deseribed in

4 DR. F. E. BEDDARD ON

1904, from an example without a skull, as Midas goeldii, but

which had recently been redeseribed by Dr. Ribeiro, from the

living animal, as Callimico snethlageri, a new genus and species

A intermediate between Callicebus and Mico.”

Mr. Thomas's examination of Dr. Ribeiro’s type specimen, sent

over by the authorities of the Para Museum, showed that Calli-

mico really was intermediate between the Cebide and Callitrichide,

having the external characters of a Marmoset, notably the elon-

gated ‘claws, combined with the shape of skull and molar formula

wf the Cebide, The molars themselves possessed no hypocone, as

in the Marmosets.

The animal being, therefore, intermediate in character be-

tween the two families Cebidee and Callitrichide, there was great

difficulty in deciding as to the effect its discovery should have on

the systematic arrangement of the American Monkeys, and as to

whether these two families ought still to be kept separate.

On the whole, as causing least disturbance, Mr. Thomas thought

‘that the best plan would be to form a special subfamily, the Calli-

miconine, for Callimico, and to include this as a second subfamily

with the Cebine in the fainily Cebide. But that Callimico was

areal genetic link between the two families there seemed to be

no doubt whatever.

PAPERS,

1. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea. By Frank HE. Bepparp, D.Sc.,

M.A., F.R.S., F.Z.8., Prosector to the Society.

[Received September 27, 1912: Read October 29, 1912. ]

(Text-figures 1-10.)

VII. On Srx Spectres oF TAPEWORMS FROM REPTILES,

BELONGING 10 THE GENUS [CHTHYOTZNIA (s. l.).

INDEX.

Systematic : Page

Species of Ichthyotenia (Acanthotenia) ............ 5

TIchthyotenia (Acanthot@nid) Sp ...... 0.660 see vee cee eee 6

T. (Acanthotenia) nilotica, 8p. NM. ..........0. 2-0... 11

T. (Acanthotenia) sp... Pier Meee neti: See at aL G,

I. (Acanthotenia) var ia, sp. "TS Sead astant nanan 17

Ih, (kee OEE DUG) GTROCHNIS, S18 Wo seeeostopeeedeseee 55 23

Ophidotenia, SOM MOVE were ec eae erie sees setae seen ie IMO

O. nai@, sp.n. Fob DES eae eA aR Eee pital 7)

Genera of Iehthyoteeniidae. COW ae atavs been ee Ne ay MMOS

a RY Sybil pla amn 5 HoH EE a0 *oeiche ian GACH SU BRSRE OPA GRA URED EER Scab acl Keo)

The genus Lchthyotenia (whose possible subdivisions I discuss

later) is chiefly found among freshwater fishes—that is, most of

REPTILIAN TAPEWORMS. 5

the species infest those vertebrates. ‘Two species, however, viz.

I. schultzei* and J. linnbergit, have been met with in amphi-

bians—the former in Rana adspersa, the latter in Necturus. The

other members of the genus are known to inhabit reptiles, both

snakes and lizards, and, if we allow Z'etrabothrium trionychiwm <.

to be of this genus, tortoises.

$ Some Species of Ichthyoteenia (Acanthotenia) from

Monitor Lizards.

Our knowledge of the parasites of Monitor ‘Lizards is obviously

not very extensive at present, for very few species of these lizards:

have been examined or at any rate have yielded parasites. These

have been referred to five genera, viz. Duthiersia, Scyphocephalus

(both of which are Bothriocephalids), Pancerina, Palaia, and

L[chithyotena (including <Acanthotenia). With the two first-

named genera I do not concern myself in the present communi-

cation. The name dAcanthotenia, whether necessary or not, was

apparently given by v. Linstow under a misconception of the

structure of the worm of which he gave an account under the

name of “ Acanthotenia shipleyi, nov. gen. et sp.”?§. In my

opinion it was probably by reason of having not seen the peri-

pherally situate vitelline glands, and of having wrongly termed

the ovaries vitelline glands, that v. Linstow created the genus,

rather than by virtue of the densely-set spinelets upon the scolex,

though the latter clearly suggested the name. Two worms from

another species of Varanws had been previously (in 1900) described

by v. Ratz || as members of the genus Jchthyctenia; and this

name is retained by Schwarz 4], who, however, was apparently

unacquainted with v. Linstow’s paper, which is not quoted in his.

list of literature. Finally, at about the same time as Schwarz,

T. H. Johnston ** described under the name of Acanthotenia

a fourth species from Varanus varius. So far as I am aware,

this exhausts the species of Jehthyotenia which have been

recorded from lizards of the genus Varanus.

It appears to me to be probable that Palaia varani, a new

species and genus described by Dr. Shipley tt from Varanus indicus,

is also to be referred to this genus Jchthyotenia. It is true that

Johnston,in his memoir upon Lchthyotenia (Acanthotenia) tidswelli,

is not of that opinion ; there are, however, certain reasons which

seem to be favourable to its inclusion within the present genus.

* Hungerbthler, Jen. Denkschr. 1910.

+ Fuhrmann, “* Die Tznien der Amphibien,” Zool. Jahrb. ix. Anat. 1896.

{ Lonnberg, “ Ueber eine neue Tetrabothriumspecies, &c.,” Centralbl. Bakt. u.

Paras. xv. 1894.

§ Centralbl. Bakt. u. Paras. xxxiti. 1903, p. 534.

|| Centralbl. Bakt. u. Paras. 1900; C. R. Soc. Biol. 1900; Arch. de Paras. iv. 1900.

§ “Die Ichthyotzenien der Reptilien, &c.,” Inaug.-Diss., Univ. Basel, 1908. Fora

copy of this I am greatly indebted to the author.

** “On a new Reptilian Cestode,’ Proc. Roy. Soc. N.S. W. vol. xliii. 1909, p. 103.

yy Description of Entozoa é&e. in Willey’s © Zoological Results,’ Cambridge, 1900.

6 DR. F, E. BEDDARD ON

It must be remembered that when Dr. Shipley described Palaia

varani the genus Lchthyotenia had not actually been described

from Varanus, since Ratz’s memoirs appeared contemporaneously

with that of Dr. Shipley. Dr. Shipley himself thought that

Palaa came nearest to Oochoristica, a genus which also occurs in

reptiles. He did not consider Panceria, to which I shall refer

immediately in this connection.

The reasons which lead me to regard Palaia varani as a species

of Ichthyotenia are largely negative, but there are also one or

two positive facts. Firstly, the only Teenioid parasites of Varanus

are Ichthyotenia and Pancerina, and as to the former it is now

known that four species of Varanus harbour different species of

Ichthyotenia. This is not obviously conclusive against the

occurrence of another genus; but it may be borne in mind

as something of an argument. The general structure of Palaia

is not in the least opposed to its inclusion in the genus Jchthyo-

tenia. It is of course true that the same characters would fit

other genera, and unfortunately the reproductive system could

not be described in any detail. The author gives a figure of a

transverse section of the worm, whence it would appear that the

transverse muscular layer lies in the medullary parenchyma, as

has been described by Schwarz in his memoir referred to on this

genus Ichthyotenia. The most important positive reason for the

inclusion of Palaia varani in the genus Ichthyotenia is in the

character of the eggs, which are particularly described and figured

as lying in clumps, the individual eggs being adherent. This is

highly characteristic of the reptilian members of the genus, and

I have something in the present paper to say upon the matter.

The genus Pancerina (originally described under the pre-

occupied name of Panceria) occurs in Varanus arenarius, whence

it has been described by Sonsino* and Stossicht, more fully

by the former of these two authorities. Pancerina is placed by

systematists = im the neighbourhood of Oochoristica, to which

Dr. Shipley thought that Palaia was allied. The double repro-

ductive organs in each proglottid prevent any confusion between

this genus and Jchthyotenia. Nevertheless there do not appear

to me to be any positive facts which forbid the near relationship

of Pancerina and Ichthyotenia, there.being no definite statement

by Sonsino as to the vitelline gland.

In two specimens of Varanus niloticus a series of small tape-

worms was found which clearly belong to this genus /chthyotenia.

I am inclined to think that the worms from one specimen of this

Varanus are different from those found in another specimen of the

lizard, But inasmuch as the worms of one series were not mature,

J am not able to speak with certainty as to some necessary specific

characters. Moreover, I had the opportunity of examining one

series in the living condition, and am not therefore able to report

* Monit. Zool. Ital. vi. 1895, pp. 121 & 189.

T Boll. Soc. Ady. Trieste, xvi. 1895, p. 42. Described as Tenia varani.

i #. g., Ransom, Bull. U.S. Nat. Mus. no. 69, 1909.

REPTILIAN TAPEWORMS. 7

certain characters as obvious in the living tapeworm which

could not be detected in the alcohol-preserved examples from the

other lizard. In the meantime I furnish a separate description

of the two series of tapeworms.

The immature worms are distinctly smaller than the others,

which is again in favour of regarding them as different species.

It is interesting to note that in Varanus niloticus, as in Varenus

varius (as will be pointed out later), a small and a comparatively

large species of Ichthyotenia occur together. But it must be

borne in mind that in the case of Varanus niloticus they did not

infect the same individual.

These worms are indeed quite small, measuring not more than

from 10 to15 mm. They present quite the appearance of a fine

white thread, as in the case of Ichthyotenia tidswelli and of one

of the species which I describe in the present paper from Varanus

varius. I havestudied them entire and in transverse and sagittal

sections; but owing to their immaturity my description has many

lacunze, in consequence of which I do not propose to give a name

to the species. The most striking external character is one that

I have not met with in any other tapeworm. The base of the

scolex and a certain amount of the ensuing strobila was tinged of

a bright carmine, which I thought at first was merely a blood-

stain. Itis not, however, blood-colour by reflected or transmitted

light, and is distinctly bright pink. Moreover it resisted for a

certain period the action of alcohol, but was ultimately washed out.

This pigment does not occur in the form of granules, like the

black. pigment which is not rare in the heads of tapeworms. It

is diffused through the tissues, though I am not able to delimit

the regions where it occurs. It is highly characteristic and quite

unmistakable. Nothing of the kind appears to be mentioned by

Johnston in Jchthyotenia (Acanthotenia) tidswelli.

The scolex of this worm is very mobile, and the anterior end

(the rostellar region) can be protruded far or flattened down or

actually retracted, thus producing an apical and conical cavity

which might be mistaken for a sucker-like structure in an indi-

vidual which had been fixed in death in this position. As the

majority of tapeworms are always described from preserved

examples, it is worth while to dwell upon this great flexibility of

movement; and, moreover, Riggenbach has used the presence of

an apical dimple as an aid to the systematic arrangement of the

genus Ichthyotenia*. Among the preserved examples which I

also studied during life I find some with fully expanded and

others with retracted rostellum., This species is at least to be

contrasted in this respect with that described by Johnston, in

which the author particularly states that the rostellum is repre-

sented by a muscular plug which is not retractile. In the same

way the four suckers are very mobile and can be protruded in the

shape of finger-like processes. There are, of course, no hooks

* Rey. Zool. Suisse, iv. 1896, p. 165.

8 DR. F, E. BEDDARD ON

upon the -scolex; but there is a fine investment of minute

spinelets, which are most densely set but are absent from the apex

of the scolex. The suckers seem to look upward when in a state

of rest.

Text-fig. 1.

AWG

COM

“

Fm th

ee oo

pI se oy ine

Lpiipgy yyy’

Ga ina:

aun hin Wd Gio,

ss s May ZA

< é

Ree

e Hee ea

oy: OS IONE y=

S = J « Pap RIC

ae SS

ee &

FE ® e S

Two sections through the scolex of Ichthyotenia (Acanthotenia) sp.

The upper figure displays the partial protrusion of the rudimentary rostellum (R)

permitted (?) by the absence of external spinelets in this region as shown in

thefigure. The lower figure passes through a sucker (S), and here the covering

of spinelets is seen to extend into the interior of the sucker.

Transverse sections through the scolex (see text-fig. 1) show an

anteriorly situated muscular organ comparable with a rostelluim.

It is rudimentary, and not like that which I describe below as cha-

racteristic of the second species of tapeworm from this Varanus.

The suckers are borne at the base of the apical cone, and the

actual suckers are on processes of the body, and not sessile upon

REPTILIAN TAPEWORMS. 9

it, as they have been figured in some other species. Between the

layer of muscles which forms the actual sucker and*the tissue of

the body is a considerable plexus or convoluted section of the

water-vascular tube not shown in the figure. I should say that

I confirm Johnston’s statement for /chthyotenia tidswelli as to

the spinelets covering the interior of the sucker for the present

species. The strong muscles which retract the suckers and the

seolex generally are massed in the neck-region of the strobila into

a thick layer of longitudinally running fibres, which reduce the

medullary region of the body very considerably. The appearance

reminds one of Dasyurotenia robusta, where the neck-region has

also particularly stout muscles *,

Text-fig. 2

e aoe i ae

tae 3 ®

weed. :

b 5

Two transverse sections through the strobila of the same species of

Iehthyotenia (Acanthotenia).

The upper figure represents the “ueck”-region, and shows the strong longitudinal

muscle-bundles (M). ‘The lower figure is from the posterior region of the

body, and shows the absence of these muscles. TT, testes; on the right are

shown other parts of the reproductive system, also in an immature condition.

The water-vascular tubes are shown white in both figures.

The fibres (see text-fig. 2) are collected into bundles of various

sizes, which are close together and completely encircle the body.

* See Beddard, P. Z.S. 1912, p. 684, text-fig. 97.

10 DR. F. E. BEDDARD ON

Between this row of bundles and the subcuticular layer are other

longitudinal fibres which are not massed so obviously into a series

of separated bundles. Farther back in the body the bundles get

smaller and further apart, and ultimately disappear altogether.

The strobila of this tapeworm shows posteriorly a very marked

delimitation between the individual proglottids, which are con-

stricted at their junctions and thus produce a necklace-like appear-

ance, as is often seen in other tapeworms, e. g., Davainea. I describe

the same appearance in an equally small or rather smaller species

from Varanus varius. In transverse sections through the ripest

segments that I have been able to examine, but which were by

no means fully mature, the structure of the body is much lke

that of other Lchthyotenia. The general parenchyma is very

delicate ; the subcuticular layer of glandular cells is very marked,

and the cells are deeply stained as compared with the neighbouring

parenchyma.

I could find no layer of longitudinal fibres within the sub-

cuticular layer. They seem to ne to be as plainly absent from

this species as from the others with which I deal in the present

communication. Nor could I find any calcareous corpuscles with

an obvious appearance like those of the second species of Jchthyo-

tenia from this Varanus, which I shall describe later.

In the anterior seoments of the body there are two water-

vascular tubes on each side of the body, the dorsal tube nearly

accurately overlying the ventral; they are about equisized. In the

segments which I am now considering the ventral vessel is very

large, comparatively speaking, and often lies right up against the

subeuticular layer, thus emphasizing the difficulty—to which I

also direct attention in other species—of distinguishing the

cortical from the medullary layer. The much smaller dorsal

vessel is present, but it is not always very easy to see.

With reference to the generative organs I am not able to say

much, as they were immature. The ducts, however, pass between

the two water-vascular tubes on their way to the exterior.

The cirrus is represented by an elongated wide tube into which

the sperm-duct opens at the posterior end. This is surrounded

posteriorly by a muscular sac, which is the cirrus-sac. The cirrus

hardly narrows before its opening on to the exterior. The sperm-

duct is only coiled once or twice upon itself, and this part lies

within the cirrus-sac. The generative openings, it should be

mentioned, occur either on one side of the segment or the other.

The middle area of the proglottid is free from testes, of which

three or four lie on each side of it; so that ina given proglottid

the transverse rows of testes contain six or eight of these gonads.

The ovary is median. Iam aware that the above facts are not

in any way distinctive of the species. They only prove that it

belongs to the genus Jchthyotenia. From the characters other

than those afforded by the generative organs it is hardly safe to

attempt the definition of a new species of this genus. There is,

indeed, nothing which positively prevents the inclusion of this

REPTILIAN TAPEWORMS. lid

parasite of Varanus niloticus within the species [chthyotenia

tidswelli, excepting, indeed, the presence in the latter of a layer of

longitudinal muscular fibres in proglottids with fully developed

testes. These muscles I can only find anteriorly in my species.

It appears to me, however, to be imprudent to give a name to the

Ichthyotenia which I have just described until a good deal more is

known about its anatomy than is contained in the present paper.

In addition to the small species which has just been described,

Varanus niloticus 1s infested by a larger Jchthyotenia which

I term /. nilotica, regarding it as an undescribed species.

Tam uncertain about the length, inasmuch as I had to deal

with fragments only of various sizes. ‘The largest fragment with

a scolex attached measured about 20 mm. But as the posterior

region of this piece had not nearly acquired the dimensions of

obviously mature fragments, at least 10 mm. can be, I think, safely

added on to the length. The species is fairly slender, but not in

the least thread-like, and the stoutest proglottids do not reach

much more than ‘5 mm. in diameter, The most salient external

feature of this species differentiates it at once from /chthyotenia

varia and J. gracilis; there is no external appearance of seg-

mentation without a careful examination witha lens. Only the

slightest constriction marks the boundary line of adjacent pro-

glottids. In this feature the present species agrees with J. biroi

and J. tidswelli. ‘The ripe segments get to be longer than broad,

but they are apparently never very long.

The scolex has the usual Ichthyoteniid characters. I only

found two scolices, and in both the conical rostellum was long

and is possibly therefore not retractile. The four suckers are

clustered round its base and look forwards. I observed the usual

dense investment of spinelets which led v. Linstow to bestow the

name of Acanthotenia upon the genus. One of the two scolices

was cut into a series of transverse sections for the purposes of

further investigation. At the apex of the rostellum there was a

distinct sucker-like depression carried on a narrow prolongation

of the rostellum, which, however, showed no such great spe-

cialization in its tissues as the four suckers proper, though it is,

I am sure, a permanent structure and not a mere temporary

involution of the rostellar region. Such a structure has been

discerned in other species of reptilian Jchthyotenia, and has been

regarded as the homologue of the rostellum of other tapeworms.

In the smaller /ehthyotenia from Varanus niloticus that has just

been described the apical structure was obviously of the nature

of a retractile rostellum.

The four suckers are borne upon prolongations of the scolex in

which lie water-vascular tubes, which latter were not quite so

conspicuous as in the last mentioned species of Jchthyotenia.

It is these outgrowths of the scolex that are the motile parts of

the scolex when the suckers are protruded or withdrawn. Toa

naked-eye examination the suckers appear to be sessile upon

12 DR. F. E. BEDDARD ON

the scolex. It is not impossible that we have here an intermediate

state of affairs, where a true sucker (acetabulum), originally only

an accessory structure (as has been held), has nearly completely

replaced the phyllidium, of which comparatively small traces

remain. In connection with this matter, it must be borne in

mind that the systematic position of the. genus Ichthyotenia 1s

not yet settled. Some place it in a family, Ichthyoteniide of the

group Tetraphyllidea ; while Lonnberg * dwells upon its likeness

to Tetrabothriwm, and Lihe fT places the certainly closely allied if

not congeneric Cr apiclobalinion gerrardi also in the genus Tetra-

bothrium.

The muscles which move the scolex are well désolopad in this

species, but by no means so prominent as in the last species.

This is remarkable when we consider that Ichthyotenia nilotica

is the larger of the two species. On the other hand, I have no

information as to the mobility of the scolex in the present worm.

Immediately behind the scolex these muscles are massed into

regular and, of course, longitudinally running bundles of fibres,

which are not so large as are those of the last, and therefore do

not contain so many individual fibres. These bundles lie below

the subcuticular layer, and occupy exactly the position in which

the longitudinal fibres of other species of /chthyotenia are repre-

sented by Schwarz. But in the present species, as in the last,

they only exist, and for a very short length, in the part of the

body immediately following the scolex. I counted ten of these

bundles in a section in the neck-region, only just behind the

scolex; they form a complete ring, and thus lie without as well

as within the water-vascular tubes. Further back the bundles

decrease in number, and in mature segments are not recognisable.

The greatest number of fibres in a bundle is 7 or 8

The structures seen in a transverse section of a proglottid (text-

fig. 3) are not different from those of other species of this genus

which I describe in the present paper. ‘The delicate layer of longi-

tudinal fibres below the cuticle is plain and also the very strongly

marked subcuticular layer of darkly staining cells. I could find no

longitudinal muscular layer beneath this, nor is there any boundary

line that I could discover between the cortical and medullary

parenchyma. Calcareous bodies were very evident here and there.

The water-vascular tubes ave very plain in the anterior region as

two tubes on each side. These are not very different in calibre,

and the dorsal and ventral of each side were more or less accu-

rately superposed. In the mature segments I can find only one

tube on each side, which I am inclined, mainly on account of its

small size, to regard as the dorsal water-vascular tube. In longi-

tudinal horizontal sections I could detect no evidence of more

than a single water-vascular vessel on each side. JI have also

been unable to find any branching of this tube to join the

corresponding vessel on the opposite side of the body. A single

* Centralbl. Bakt. u. Paras. xv. 1894, p. 801.

y+ In a footnote to a paper, “ Zur Kone einiger Distomen,” Zool. Anz. 1899,

REPTILIAN TAPEWORMS, 133

water-vascular vessel on each side of the body is also described

for Ichthyotenia biroi. I find, however, that in completely ripe

Text-fig. 3.

Transverse section through the strobila of Ichthyotenia (Acanthotenia) nilotica.

se.l. Subcuticular layer. ¢. Testis. wt. Uterus appearing as an irregular series of

cavities with numerous contained embryos. v.g. Vitelline gland.

segments with embryos a much larger ventral vessel is present

in addition to the dorsal.

14 DR. F. E. BEDDARD ON

There are various points in the structure of the reproductive

system of organs which help to fix the distinctness of this species

from others, and especially from Jchthyotenia biroi, to which it is

most nearly akin. The éestes occupy the greater part of each

ripe proglottid, and there is no very definite median area free

from them. ‘There is nothing so definite, for instance, as the

conditions figured by Schwarz in Ichthyotenia wattereri*. In

horizontal sections of a series of mature segments the difficulty

of distinguishing the boundary lines between successive segments,

plainly felt on examination of the entire worm with a lens, is

increased by the lack of a definite break between the reproductive

systems of each segment. There is an appearance of complete

continuity not seen in corresponding sections of the other species

which I have studied.

The vas deferens behind the cirrus-sac is closely coiled, this

coiled region reaching quite or nearly halfway across the pro-

glottid. The calibre of this region of the vas deferens does not

vary in any appreciable way from point to point. That is to say,

there is not, so far as I have been able to ascertain, any vesicula

seminalis like that of Jchthyoteenia gracilis described in the

present paper. It 1s important to note this fact, since in J. birot

the existence of a vesicula behind the cirrus-sac has been stated,

but Schwarz was not able to see it. The tangled coil of the vas

deferens lies across the segment with a slight inclination towards

the anterior margin. Within the cirrus-sac the vas deferens is

also coiled.

The present species is the only one of those known to me at

first hand, and which are described in the present paper, which

possesses an armed civrus. This organ, when protruded, is covered

externally with very numerous and closely-set hooks. In sections

through the cirrus-sac, where it is not protruded, the spines can

be seen lining the canal of the cirrus. Schwarz figures in the

case of Ichthyotenia marenzelleri ~ an actual protrusion of the

cirrus-sac itself in an apparently unaltered condition. There is

no doubt that in the species with which I am here concerned

the outpushing of the cirrus is an eversion of the cavity of the

cirrus, since the spines lining the latter become external in the

everted organ.

The cirrus-sac and vagina open very slightiy in front of the

middle of the lateral margin of the segment. The generative

apertures, of course, alternate from the side of the body in

different segments, and there is also, as in many but not in all

species of the genus, an alternation in the relative positions of

the two orifices, the cirrus being sometimes anterior and some-

times posterior. The generative ducts pass between the dorsal

and ventral water-vascular tubes, and there is in the present

species no cloaca genitalis. This is rendered particularly evident

in one specimen which I mounted entire in glycerine, and of

* Loc. cit. Taf. i. fig. 2, h.

+ Loe. cit. Vat. iv. fig. 12, ¢, b.

REPTILIAN TAPEWORMS. 15

which the cirri in many segments were extruded in various

degrees of completeness. By the side of the cirrus in several

proglottids was an oval projection, which is clearly the slightly

everted or protruded vagina. Von Ratz has described the ovary

of Ichthyotenia biroi as possessing digitiform processes. The

ovary of Ichthyoteenia nilotica is solid, and the two wings are not

frayed out into processes ; its appearance is, in fact, that of most

species of the genus. The vagina when not fully mature is apt

to be arranged, in that part of it which runs parallel with the

cirrus-sac, im a sinuous line. Jn quite fully mature segments the

vagina runs in quite a straight course in this region, and is con-

siderably dilated, the calibre increasing as it passes back. It is

not easy, however, to distinguish any part of it as a receptaculum

seminis. But, on the other hand, it probably corresponds to the

“‘spindelfoérmige Anschwellung” seen by von Ratz but not by

Schwarz in the species /ehthyotenia biror. After the pear-shaped

swelling, which is quite plain in J. nilotica, the vagina dwindles

greatly in calibre and becomes quite a narrow tube. That this

dwindling occurs rather suddenly gives an additional appearance

of a receptaculum seminis to the tract lying in front of it. As in

other species, the vagina has an adventitious sheath formed by an

ingrowth of the subcuticular layer. The vagina, when it reaches

the middle of the body, passes back in a straight line, and is a

little coiled in the neighbourhood of the ovary and before it

becomes connected with the shell-gland. The vagina lies dorsally

of the ovary and the uterus lies ventrally of the ovary in this

region. The shell-gland is large, and extends across about half

of the dorso-ventral diameter of the proglottid. It lies behind

the median junction of the two wings of the ovary. Close to it,

but upon the ventral side of the median, is the very obvious and

strongly muscular ‘‘Schluckapparat.” The vitelline glands form

the usual lateral strips.

Before the eggs are ripe the wterus is visible as an empty cavity

running antero-posteriorly through the whole of the middle of

the segment. It is straight, with numerous closely adpressed and

irregular outpocketings. The appearance may also be described

as of a wide tube partially divided by numerous ingrowths of the

wall. When fully formed and containing ripe embryos the

uterus extends over the greater part of the segment from side to

side, though it does not touch the lateral vitelline glands. It has

an exaggeration of the same structure, the lateral diverticula

being prolonged. In transverse sections through ripe proglottids

(see text-fig. 3) it often appears as a series of more or less isolated

and circular cavities occupying the middle of the segment. The

ripe eggs lie loosely in the cavities of the uterus, generally

aggregated into masses. The shell of the egg is very thin and

unrecognisable, which distinguishes this species at once from

Ichthyotenia varia, described later in the present paper. The

eges of Ichthyotenia nilotica are also-—and, perhaps, therefore—

distinctly smaller than those of the other species just mentioned.

16 DR. F. E. BEDDARD ON

While there is no need to insist upon the difference between

Ichthyotenia nilotica and any of the other species described in the

present communication, it is equally clear that there are consider-

able resemblances between that species and J. biroi. They are

however, as I think, distinct. In the first place, our knowledge

of these parasites of Varanus shows so far that, while a particular

species of Varanws may harbour more than one species of [chthyo-

tenia, the same species of /chthyotenia does not infest more than

one species of Varanus. But there are also definite structural

distinguishing marks. Of these the most important is the form

of the ovary, which is quite normal in J. nilotica and peculiar in

T. biroi. 'The spiny lining of the cirrus appears to be a further

distinguishing mark, as Schwarz in his general account * of the

anatomy of the genus says nothing about it.

An example of Varanus varius contained specimens of tape-

worms which were clearly referable to two species—one very

much larger than the other. The smaller species was represented

by only one individual, which was a delicate thread-lke worm,

quite agreeing with Acanthotenia tidswelli of Johnston 7 in that

“to the naked eye it appears like a piece of fine white thread.”

Furthermore, the head is covered with minute densely-set

spinelets. In other external characters, however, this small

worm does not agree with Acanthotenia tidswelli. 1t is very

much smaller, not reaching beyond 6 mm. in length as compared

with 30 mm. for Acanthotenia tidswelli; there are abundant

calcareous bodies, and the segments are rather clearly defined

after the neck. The neck is quite short, and the segments soon

get to be as long as they are broad. I counted altogether forty |

distinct segments after the neck.

Of these the last few had an oval outline and were deeply con-

stricted at their junctions. Whether these were fully mature

segments or not I am unable to state, as I could not see the

organs of reproduction in the single specimen, which was mounted

entire in glycerine; nor, indeed, could I see the generative pores.

It is thus only by inference that the worm is referred to the

genus Ichthyotenia (from which it is, as I think, very difficult to

distinguish Acanthotenia), with which, however, all available

characters agree in uniting it. I should say that, as might be

expected, the head is entirely unarmed with hooks; it is con-

siderably wider than the ensuing neck and the body for some

distance. Mr. Johnston found the minute spinelets of the scolex

in A. tidswelli to occur also upon the following proglottids. I

have not been able to ascertain that this is so in my species.

On account of the very scanty amount of notes which I am able

to set down concerning this species, I do not for the present give

it a name.

The second very much larger species, which I propose to name

* Loc. cit. p. 13.

+ Proc. Roy. Soc. New South Wales, vol. xlii. 1909, p. 103.

REPTILIAN TAPEWORMS. iW

Ichthyotenia varia, was represented by a great number of seg-

ments, but there was not a complete worm; so that I cannot judge

of the length of the species. I can say something, however, of its

dimensions as compared with the other species of [chthyotenia

which inhabit the genus Varanus. The longest piece measured

28 mm., and as this consisted of long and mature proglottids

only, it is certain that a complete individual would measure

more than 30 mm. A single proglottid, indeed, in one or two

cases was 4 mm. long, and thus the worm is obviously relatively

a large one as compared with other species that inhabit Varanus.

The breadth of mature proglottids was fully a millimetre, so

that this species of Jchthyotenia is not at all thread-like but

flattened and ribbon-like: indeed, in all external characters a

typical member of the genus. Among the numerous fragments

of varying sizes and maturity was one piece which included a

scolex, which is, [ think, probably not to be referred to the same

species. I shall deal with the characters of this individual, which

measured 34 mm., later. The pieces of the large worm were

investigated by sections, and also pieces were mounted entire,

stained and unstained, in glycerine.

In transverse sections of mature proglottids (see text-fig. 4,

p. 18) the flattened character of the worm is very obvious. The

medullary parenchyma is much more coarsely spongy than I

have observed it to be in many tapeworms, and I have been

unable to detect a transverse layer of muscle separating the

medullary from the cortical layer. Moreover, the cortical layer

must be thin, for the coils of the vas deferens extend dorsally

and ventrally to just beneath the cellular subcuticular layer.

The latter layer is very strongly marked, as appears always to

be the case in this genus. The large cells are very deeply stained

by hematoxylin. Between it and the cuticle is a delicate layer

of longitudinal fibres; but I can find no other layer of longi-

tudinal muscles such as are figured by Schwarz * and Johnston 7,

or have been described by myself in the present paper in an

Ichthyoteenia from Varanus niloticus.

It would, I think, be difficult to miss such a layer in my prepa-

rations were it present. But while there is no definite layer of

muscles, there are certainly here and there large longitudinally

running fibres which I have observed both in transverse and

longitudinal sections. But they are scarce. As for the transverse

layer which Schwarz figures { as occupying the greater part of

the medulla, I have only been able to notice a rather stretched

condition of the parenchymatous meshwork, particularly at the

extremes of the proglottids where there are no gonads developed.

There are very probably muscles in this meshwork running in a

transverse direction; there certainly are such fibres running in

a dorso-ventral direction. I cannot, however, find thick layers

such as I understand Dr. Schwarz to insist upon in Jchthyotenia

calmetter. Owing to the coarseness of the medullar meshwork

* Loc. cit. Tat. i. fig. 4. + Loe. cit. £ Loe. cit. pl. iii. fig. 10.

Proc, Zoou. Soc.—1913, No. II. 2

18 DR. F, E, BEDDARD ON

and the fact that the spaces in the meshwork are often approxi-

mately circular in transverse section, it is not always easy to

detect the water-vascular tubes. They are, however, often quite

easy to see, and in such cases I] have seen two on each side—a

Text-fig. 4.

Transverse section through the strobila of Ichthyotenia (Acanthotenia) varia.

d.v. Dorsal water-vascular tube. lm. Layer of delicate longitudinal muscles lying

between cuticular and cellular subcorticular layer. 7. Testis. @ U. Cavity of

uterus. vé.g. Vitelline gland. ov. Ventral water-vascular tube.

REPTILIAN TAPEWORMS. 19

dorsal and a ventral, fairly evenly superposed. The calibre of

these vessels is not large, and they do not differ greatly in size.

I have observed branches going towards the exterior, but have

not traced them as far as an opening. The longitudinal trunks

lie sometimes to the inside of, and sometimes above and below, the

vitelline mass.

The testes, as can be best seen in transverse sections, form a

layer, only one deep, of 80-100, or even more, in a single pro-

glottid. While there is not a median space entirely free from

testes in this species such as occurs in some other species of the

genus, the testes are often not quite so densely arranged in the

median area; they occur there, however, even when the uterus.

is fully formed, in the thickness of the strands which form its

diverticula. The testes do not extend back to the sides of the

ovary, but anteriorly they may (or may not) reach quite to the

end of the segment.

The vas deferens, after issuing from the cirrus-sac, forms a

dense coil, which extends over about half of the breadth of the

proglottis. It les at right angles to the long axis of the pro-

glottis, and is directed neither forwards nor backwards. The

tube forming this coil is thin-walled and of wide calibre. Within

the cirrus-sac the sperm-duct is also coiled, but it is here of much

less calibre. J was not able to see any hooks in connection with

the copulatory apparatus. The aperture of the cirrus-sac was

always in front of the orifice of the vagina. The cirrus-sac is

flask-shaped as in other Ichthyoteniids and in many tapeworms,

but there is not so sharp a demarcation between the two regions

as is socommonly the case. The walls are thin and obviously

muscular; they are not thicker in the neck of the cirrus. It.

is necessary to insist upon the muscularity of the walls of the.

cirrus-sac since Schwarz* has written of that of Ichthyotenia.

natiereri: ‘Die Blase (i. e. of the cirrus-sac) ist diinn, nicht.

muskulés.” It should be mentioned that the alternating gene-

rative pores lie some way behind the middle of each proglottid .

The ovary is, as usual, in two portions, with a slender bridge im

the middle line uniting the two. The contour of the ovary,

though irregular, is not pulled out into digitiform processes as in

Ichthyotenia biroi. The vagina runs parallel with the cirrus-sac

and posterior to it, as already mentioned. This region of the tube

is wider than that lying behind; but the vagina is here not

coiled, and, indeed, scarcely wavy in outline. Posteriorly between

the wings of the ovary the vagina is coiled for a short distance.

* Loe. cit. p. 20.

+ In one proglottid, which was fully ripe and contained numerous embryos, I

observed a remarkable variation which I have not seen before in a tapeworm. The.

cirrus-sac with the succeeding large and complex coil of the vas deferens was particu-.

larly obvious ; but the most careful examination failed to show a vagina lying either

in front of or behind the cirrus-sac. But by following up the vagina from the ovary

forwards it was found to bend in the reverse direction and to open on to the opposite

side of the proglottid, nearly on a level with, but slightly behind, the line of opening

of the male pore. I regard this state of affairs as merely a variation, but I have not

read of any parallel mstance.

Das

2.0 Dk. F. E. BEDDARD ON

The dilated region of the vagina (the dilatation is very slight)

has apparently not got a ciliated lining. It has a covering of

glandular cells externally which is obviously continuous with, and

presumably derived from, the glandular subcuticular layer of the

body generally. The shell-gland is large and conspicuous, and

the “Schluckapparat” is particularly muscular and strong. I

have observed a duct arising from each peripheral vitelline series,

and there is in short nothing of particular note in this part of

the generative system.

View of a ripe proglottid of Ichthyotenia (Acanthotenia) varia, to illustrate

the extent of the mature uterus.

BE. Egg-clumps scattered through the uterus, which occupies the greater part of

the proglottid. ZT. Testes.

The wterus of this species when fully formed and containing

embryos occupies a good deal of the middle region of ripe

proglottids (see text-fig. 5). Its aspect in transverse sections is

quite like that of Ichthyotenia nilotica described above, and no

definite specific differences appear to me to be obtainable from

a comparison of these organs. The eggs, however, are highly

characteristic when the present species is compared with [chthyo-

tenia nilotica. They are disposed for the most part within the

cavity of the uterus in clumps of varying numbers, perhaps 12-20

in each batch of eggs. Three or four of these, or often more,

may be visible in a single transverse section. The mode of

ageregation of the individual eggs (which, of course, contain

embryos) in Jchthyotenia varia is thus different from that which

is found in Jchthyotenia nilotica. They also appear very con-

siderably larger, the difference in size being principally a question

REPTILIAN TAPEWORMS. Al

Text-fig. 6.

Transverse sections through two compartments of the uterus of

Ichthyotenia (Acanthotenia) varia.

m. Outer sheath enclosing several embryos, each of which has also its own separate

sheath. TJ. Testes. U. Cavity of uterus. W. Wall of uterus.

In the upper figure a clump of embryos is shown in which the outer membrane is

not present or just represented by a rudiment on the left side. In the lower

figure is shown a clump of embryos in which there is an outer sheath

involving the whole mass,

YE DR. F. E. BEDDARD ON

of the increased shell. Text-figure 6 (p. 21) shows a clump of

eggs in transverse section within a compartment of the uterus,

pressed to whose boundary walls on either side is a testis.

This figure may be compared with text-figure 3 (p. 13), which

illustrates a corresponding view of an egg-clump in Jchthyotenia

niloticn.

The masses of eggs of Ichthyotenia varia are, however, some-

times rather different from that represented in the drawing to

which attention has just been called. Occasionally (as shown in

text-figs. 6, 7) each mass of eggs is surrounded by a common

Text-fig. 7.

Two isolated masses of embryos from the uterus of Ichthyotenia (Acanthotenia)

varia, viewed entire as transparent objects, to show the sheath which encloses

a variable number of embryos.

sheath or shell, within which lie the eggs, or rather embryos, each

with its own shell. Thesame state of affairs is quite plainly visible

in preparations of a proglottid mounted entire (text-fig. 5) in

glycerine, where the eggs appear to be contained within a glass

ball. In the text-figure to which I have referred the outer sheath

is represented as thin; it is often considerably thicker, and I

have seen it with a few thin obliquely-set spiny processes. An

aggregation of the eggs within the uterus appears also to occur

in Ichthyotenia saccifera of v. Ratz, but there are not sufficient

details given to compare with the conditions observed by me in

I. varia. As to the egg-masses enclosed within a common sheath,

they suggest spores within a sporangium, and give rise to the

idea (which I admit to be otherwise unsupported) of the division

REPTILIAN TAPEWORMS. 23

of an originally single ovum to form a number—thus lying within

one sheath.

In describing the last mentioned species I have referred to a

scolex which probably belongs to a third species, and a further

examination of this and of some other fragments has convinced me

that this is so, and that I am right in describing these worms

under a different name.

The present species is intermediate in size between the two

last mentioned. As already suggested, it is probably at least

40 mm. in length, for the largest fragment, that including the

scolex, is quite 34 mm. long. ‘The head is wider than the body

which follows, and the greatest breadth is not attained for some

time; the elongated ripe segments are about °5 mm. in width.

This worm is therefore much more slender than /chthyotenia varia.

The rostellar region was depressed and the four suckers look dis-

tinctly upwards. I ascertained that the scolex was densely beset

with fine spinelets as in other species. There are, of course, no

hooks. The neck is very short indeed, and the proglottids very

soon get to be as long as they are broad ; later on they become

about three times as long as they are broad.

The general structure of the body when examined in transverse

and longitudinal sections is very like that of Jchthyotenia varia,

which has just been described. The strands forming the meshwork

of the parenchyma are, however, much finer, and the interstitial

homogeneous ground substance is deeply stained, whereas a

corresponding immersion in the staining-fluid of pieces of

Ichthyotenia varia produced no result in the way of tinging the

ground substance.

The segments, however, from which I made these sections,

although elongated, are rather younger than those which I

examined (and which alone I was able to examine) in Jchthyotenia

varia. The outer longitudinal muscular layer lying outside of

the subcuticular layer’ was obvious; but, as in J. varia, I could

find no other definite longitudinal layer. The subcuticular cells

were strongly developed and very darkly stained. The water-

vascular tubes are two on each side and superposed, the dorsal

lying often fairly accurately above the ventral.

The disproportion in calibre between the two tubes and their

actual size is greater than in the last species. I can find no trace

of transverse vessels. I could find no caleareous bodies, which are

present, though not abundant, in /.varia. The principal difference,

however, between the present species, which I propose to call

Ichthyotenia gracilis, and J. varia lies in the reproductive organs.

In the first place, the opening of the external pore is rather

nearer to the middle of the proglottids—sometimes, in the case of

shorter proglottids, actually median on the Jateral margin. It is

never so posteriorly situated as in J. varia. I counted 80 testes

in one proglottid, which are more definitely arranged in this

species in two lateral bands, the median area being largely and

24 DR. F. E. BEDDARD ON

sometimes entirely free from testes. The sperm-duct shows an

important difference from that of Jchthyotenia varia. Almost

immediately after leaving the cirrus-sac it becomes dilated into a

vesicula seminalis in a fashion not met with in the last described

species. Moreover, the coil of sperm-duct which ensues is con-

siderably less in extent than that of J. varia, and instead of

running out towards the middle of the segment is directed rather

obliquely and forwards. There is thus a greater angle formed

with the vagina. The structure of the cirrus-sac appears to me

to show no obvious differences from that of the Jchthyotenia

which has just been described. The vas deferens is coiled within

it in a precisely similar fashion. As a rule the vagina, which lies

parallel with the cirrus and is here somewhat dilated, lies

posteriorly to that sac; but I found it in one instance to open in

front of the cirrus-sac. Ichthyotenia gracilis thus differs in this

character also from Jchthyotenia varia. In the neighbourhood

of the ovary the vagina is a good deal coiled ; but my account of

the female organs of this species is very incomplete, since they

were not fully mature. The shell-gland was large and well

forward in development ; but I could not definitely recognise the

lateral vitelline strips. The uterus was in the condition of a

delicate thread commencing anteriorly and extending far back in

the middle line of the segment.

I am therefore able to distinguish plainly three species of

Ichthyotemia occurring in the gut of Varanus varius, which are

apparently graduated in size, and of which the smallest (to which

I do not venture to give a name for the present) is the only one

that bears any external likeness to the only species of Jchthyo-

tenia (viz. I. tidswelli) as yet recorded from this particular

species of Varanus. I have, however, shown ample reasons for

regarding it as different. There can also be no question as to the

marked distinctness of J. varia and J. tidswelli quite apart from

size, and here there appears to be too great a discrepancy to allow

of specific identity. J. tidswelli differs from my species by its

total lack of calcareous corpuscles, by the alternating relative

position of cirrus-sac and vagina, by the disappearance of the

uterus and the lodgment of the ripe ova in the parenchyma*, by

the freedom from testes of the middle region of the proglottid,

and by the strong muscular layers of longitudinal fibres lying

within the subcuticular layers of cells.

With my Ichthyotenia gracilis Johnston’s LF. tidswelli agrees

more closely; yet they are, as I think, different. In the first

place, /. gracilis cannot possibly be described as ‘like a piece of

fine white thread,” a description applied by Johnston to

I. tidswelli. I can find neither in J. gracilis nor in J. varia the

strong muscular layer figured by Johnston, and also in other

species by Schwarz. Finally, the existence of what may be

* The author of the paper upon TZ. tidswelli is not, however, able to insist

absolutely upon this.

REPTILIAN TAPEWORMS. 25

termed a vesicula seminalis appears to be distinctive of J. gracilis,

and thus contributes largely to the establishment of its distinct-

Ness as a species.

This latter character also precludes the possible identification of

I. gracilis with either of the two species described by v. Ratz,

from which both J. gracilis and /. varia differ also in other

particulars. In J. birot the genital pore lies in front of the

middle of the segment instead of behind it, the neck is long, and

the proglottids, even posteriorly, are not plainly separable to the

eye; the water-vascular tube is single on each side. It is therefore

not identical with either of my species. In J. saccifera the

positions of male and female pores alternate as in /. gracilis; but

the cirrus-sac is described as round, as opposed, I presume, to oval,

which is its shape in J. gracilis, in which species, moreover, the

proglottids are longer; but the two species are obviously very

near to each other.

Finally, I do not think that these species can be confused with

v. Linstow’s Acanthotenia shipleyt from Varanus salvator, for it

is very minute, only 13°8 mm. in length by :1—-4 mm. in breadth ;

the segmentation is not marked; the cirrus-sac lies behind the

vagina and is crescentic in shape. The author, however, seems

to have missed the marginally situated vitelline glands, since he

identifies as such what is, I think, certainly the ovary. The

description setting forth the characters of a new genus and a new

species is, however, a very brief one.

§ An Ichthyoteniid from the Indian Cobra (Naia tripudians).

OPHIDOTENIA NAIM, gen. et sp.n. (See p. 35.)

The genus is already known to exist in a good number of

snakes, of which a list was given by v. Linstow in 1907 * and by

Schwarz in the following year+. There are at most seven species

known from snakes, if we except Crepidobothrium gerrardi< from

Boa, and regard it, mainly by reason of its horseshoe-shaped

suckers, as a distinct genus. I have now to describe a form

which occurs in a species of snake which has not hitherto yielded

examples of the genus. I obtained at least three specimens

(there were three scolices) of an Ichthyoteniid from an Indian

Cobra which died in the Society’s Gardens on April 18th, 1912

The longest individual measured, after preservation in aleohol,

110 mm., and had a greatest breadth posteriorly of 1-5 mm.

The anterior end of the body gradually narrows up to the small

scolex, which is, however, wider than the neck which immediately

ensues. In none of the specimens was the rostellum very con-

spicuous for its size and extension beyond the suckers. Indeed,

T found that in the specimen which I examined by means of

transverse sections there was no prolongation of the head at all

* “ Welminthen von Java,’ Notes Leyd. Mus. xxxix. 1907-8, p. 85.

+ “Die Ichthyotenien der Reptilien, &c.,”’ Inaug.-Diss. Univ. Basel, 1908.

t Baird, Proc. Zool. Soc. 1860, p. 446; Monticelli, Atti Soc. Nat. e Mat. Modena,

(4) 1. 1899, p. 9.

26 DR. F. E. BEDDARD ON

beyond the suckers. The very first section of the series, which

was nearly transverse, showed simply two suckers back to back

with very little tissue between them. The scolex, indeed, appears

to be very like that of Jchthyotenia nattereri as figured by Schwarz.

Naturally, therefore, as in that species, the present is provided

with neither rostellum nor a fifth sucker. Nor is there the least

trace, that I can discover, of the presence of minute spines upon

the scolex, such as characterise the species of Jchthyotenia from

Varanus. I gather from Schwarz that in other Ichthyoteniids

from Ophidia there is the same lack of minute spinelets upon

the rostellum. I may add that I cannot find these spines

lining the cavity of the suckers, a matter upon which I lay some

weight, since Johnston has particularly asserted their presence in

this situation in J. tidswelli.

The actual neck of this worm—i. ¢., that portion of the strobila

immediately following the scolex—is short, the segmentation of the

body commencing early. The proportion between depth of the

segments and transverse diameter is more equal here than later,

where the body appears, on account of its great breadth, to be very

thin. Also the dorso-ventral diameter is not greater in the neck-

region than posteriorly. In transverse sections through this

neck-region, one important difference is apparent between this

species and those which I have been able to examine among

the species peculiar to the genus Varanus. The parenchyma of

the body is uniform throughout, and appears to contain no

muscular fibres at all. It may be that there are a few which I

have missed on account of their tenuity; but it is quite clear

that there is in the present species nothing like the thick

encircling layer of longitudinal fibres which I have described

above in Ichthyotenia nilotica and Ichthyotenia sp. from the

Nilotic Monitor. This peripheral layer of muscles, were it

present, could hardly, I should imagine, have been missed, so

conspicuous is it in the two species from Varanus.

In the wider posterior part of the body the segmentation is

quite clear, though the segments, as is the case with the other

species of the genus, do not overlap. They get to be about twice

the length of their width. In transverse sections, through not

fully mature segments, the width in comparison to the depth is

greater than that which is figured by Schwarz for some species

of Ichthyotenia from Serpents. In some quite mature segments

which I examined the difference from J. nattereri, etc., in this

respect appears to be less; and in those segments the two ends

of the section were wider than the middle region and formed

an almost circular-projecting area, the whole section having thus

a dumbbell-form with a long cross-bar. The minute structure

of the body differs in some respects from that of the Jchthyo-

tenia considered in the present communication, but seems to

agree on the whole with that of species of Ichthyotenia from

Ophidia. The cuticle is thick and homogeneous; the layer

of longitudinal fibres beneath it is very distinct. The sub-

REPTILIAN TAPEWORMS. nT

cuticular layer of large pear-shaped cells is deeper than in the

Ichthyotenia from Varanus, and, like the layer in them, is very

conspicuous and deeply staining. Some way below this glandular

layer is a layer of stout longitudinal fibres, such as are figured

by Schwarz in species of /chthyotenia from Serpents, a layer

which J have been quite unable to find in such species from

Varanus as I have examined. This layer in the present species

is very conspicuous, and the differential staining of logwood

emphasises it very well. The layer is usually more than a single

fibre deep, but in places only a single fibre deep. The greatest

number of fibres which I have found in a dorgo-ventral direction

is not more than four. The layer thus appears to be thicker

than in Jchthyotenia nattereri and J. calmettei figured* by Schwarz.

The layer seems, as Schwarz has pointed out, to form the boundary

line between the cortical and medullary regions of the proglottids.

The latter author has directed attention to a peculiarity in the

disposition of the transverse fibres which differs from that of all

other tapeworms. He described and figured? these fibres as

lying within—~. e., not on—the boundary of the medullary paren-

chyma. I have myself observed in the present species many such

fibres in proglottids not fully mature. In fully mature proglottids

IT have not detected them. They are irregular and certainly

occupy a good deal of the medullary region. They are not, how-

ever, by any means so plain as the longitudinal fibres just

described.

The water-vascular tubes in the neck-region are plainly two on

each side. They show a character here which is at least not.

very common among Tapeworms. Both the tubes, which are

fairly accurately superposed, have distinctly thick walls and are

rather larger than usual. Moreover, the lumen is slit-like, the

tubes presenting the appearance of being compressed dorso-

ventrally ; there is no doubt that in the fully mature proglottids

only one of these tubes is present on each side of the body.

The generative organs conform with the Ichthyoteniid plan, but

show numerous differences in details from those of the other

species already considered in the present paper. The testes

present the usual arrangement, but are, perhaps, more strictly

confined to the lateral regions of the proglottid than in some

other species described in the present paper. When they are

fully ripe the testis seems to become entirely converted into a

mass of spermatozoa, which does not wholly fill the cavity in

which it lies. In transverse sections a series of these cavities are

seen to lie side by side, only separated by a thin strand of

medullary parenchyma.

The vas deferens forms the usual Ichthyoteniid coil, but seems

to be of less calibre than in the species of this genus from

Varanus. ‘This coiled region extends to quite the middle of the

* Toe. cit. Taf. i. fig. 4 and Taf. ii. fig. 10, Um.

+ Loc. cit. Taf. iii. fig. 10, tum.

DR. F. E. BEDDARD ON

A series of transverse sections through generative ducts of Ophidotenia naie.

B. The extremity of the cirrus (3) and the va

rt of the cirrus lying without the cirrus

A. The cloaca genitalis.

gina (2).

a Te. |

-sac (¢') and the

sac ( $) containing coiled

he cirrus are shown black.

vagina (2) already narrowed in calibre. D. Cirrus-

C. The terminal pa

cirrus; () vagina.

The glandular cells besetting t

Re a ae

REPTILIAN TAPEWORMS. 29

proglottid. The cirus-sac is longer than broad, but not of a very

regular oval; its walls are thin but muscular in structure, the

number of fibres in a longitudinal section not appearing to be

more than one or two. Within the cirrus-sac lies the cirrus,

which widens out considerably in its last section—7. ¢., that lying

immediately after the extefnal aperture; the cirrus lies in many

coils and is throughout of considerable calibre. In structure it

differs from the cirrus of many other tapeworms (see text-fig. 8)

by the very thick layer of circular muscular fibres which sur-

rounds it. The muscles are increased at the broad terminal

ortion of the cirrus. Outside the muscles is a layer of pear-

shaped glandular cells which deck the whole of the cirrus. The

same arrangement, or something very lke it, apparently exists

in Ichthyotenia nattereri according to the figure and description

by Schwarz*. In that species, however, the glandular layer is

neither so thick nor so widely distributed as in the present.

I have observed in several proglottids the complete protrusion

of the cirrus, which is everted in the usual way and thus is broader

at the base than at the tip.

J have found no protrusion of the cirrus-sac itself such as has

been referred to by Schwarz. The cirrus when fully everted is

not half the width of the proglottid. There is thus a difference

from J. nattereri, in which species (very like the present in many

respects) Schwarz asserts that the protruded organ equals in

length the whole diameter of the segment. Moreover, he figures

the cirrus as very much more closely coiled within the cirrus-sac

and therefore longer than I have found to be the case with my

new species. I have not found any spines upon the cirrus. As

in many species of Jchthyotenia, the cirrus opens sometimes

in front of and sometimes behind the vagina. The cloaca

genitalis into which both open is of some depth.

At its outer end (see text-fig. 8 B) the vagina is quite as wide

as the somewhat dilated end of the cirrus; but it very soon

narrows into a tube of less calibre. The wide distal region is

-fully as muscular as the cirrus and it is furnished in addition

with a strong circular muscular sphincter. Outside of this is a

layer of glandular cells, which forms with those belonging to

the cirrus, and which have been already described, a common

Jayer surrounding the two tubes in one circle. In this region the

vagina is not ciliated. A little further back the vagina has its

own separate coat of glandular cells as has the cirrus. Later

on the vagina becomes narrower and is ciliated internally. It

bends back in the usual position and is coiled posteriorly in

the region of the ovary. The straight part of the vagina which

passes backwards lies dorsally to the uterus, but not quite in the

median line.

The ovary extends on either side up to the laterally placed

vitelline strips. I could not find any signs of a shell-gland which

* Toe. cit. Taf. ii. fig. 5

30 DR. F. E. BEDDARD ON

is so conspicuous in certain species from Varanus described above.

On the other hand, the “Schluckapparat” was large and

Text-fig. 9.

Ophidotenia naie.

A. Section through a portion of a ripe proglottid.

d. A diverticulum of the uterus. P. External pore of the uterus. V. Vagina.

a. A mass of secreted substance lying within the main stem of the uterus.

B. A section through the uterus cut transversely and showing the diverticula (a)

of the median stem. V. Vagina.

C. Part of a transverse section of a ripe proglottid.

In the centre is seen the uterus, the median stem of which sends down an outgrowth

towards the exterior of the body, of which the external orifice is seen in A.

D. A mass of secreted substance lying in one diverticulum. d. Eggsin process

of development lying in the other diverticulum. V. Vagina.

REPTILIAN TAPEWORMS. 31

muscular. This and adjacent parts of the female ducts extended

back beyond the ovary to the hinder border of the proglottid.

The uterus of this worm shows a very remarkable feature

which in detail is different from anything that has been hitherto

described in any tapeworm. I believe that I am right in saying

that an independent uterine opening is not found in any of the

Tetraphyllidea or Cyclophylliidea—that it is, in fact, confined to

the Pseudophyllidea. The present species, allied to a genus

which perhaps stands intermediate between the Tetraphyllidea

and Cyclophyllidea, is thus so far unique in possessing inde-

pendent uterine openings which lie, as in Bothr tocephalus, eter,

upon the ventral surface of the body. While there is this general

resemblance to the lower group of tapeworms, there are plenty

of differences in detail between the conditions which obtain in

this species and those which characterise the Bothriocephalide

and their immediate allies. When the ripe proglottid is viewed

as a transparent object in its entirety, 7. e., without undue

pressure, the uterus is seen to oceupy the middle region of the

body, and to be a quite narrow tube with regular paired diver-

ticula, which extend but a little way laterally.

In addition to this there were in the median line of the pro-

elottid a series of medianly situate orifices, one behind the other,

extending through the greater part of the proglottid and rendered

especially conspicuous by virtue of the fact that granules of black

pigment had accumulated in every instance near to these orifices.

f could not in such preparations actually see the tube which put

this series of’ orifices into communication with the underlying

uterus. But the orifices, on focussing down, were quite plain and

could not be missed. In transverse sections (text-fig. 9) all of the

structural characters were easily confirmed and extended. The

uterus is seen to occupy but a comparatively small space in the

middle of the proglottid, by reason of which the present species

contrasts with Ichthyotenia varia or LI. wilotica. The uterus in

a given section consists of a median stem which is a small tube ;

from this arise continually a series of paired diverticula, which

are at times somewhat ear-shaped or crescentic in outlme. The

curvature is downwards and they expand at their blind ends

(text-fig. 9B). The diverticula furthermore show a difference in

structure from the median stem of which they are outgrowths.

Their exterior is more or less covered with a layer (text- fig. 9 A)

of pear-shaped glandular-looking cells, which resemble those

found decking the cirrus and which are also of the same nature

as the glandular cells of the subcuticular layer.

These uterine pores are quite as visible in sagittal sections

(text-fig. 10) and are then seen to exist along the entire length of

the proglottid, being situated at intervals from each other. Such

sections also show very much better than the transverse sections

the fact that the median stem of the uterus is continuous

throughout the proglottid; they furthermore enable me to state

that the number of paired diverticula of the median stem in a

32 DR. F. E. BEDDARD ON

given proglottid was over twenty; they doubtless vary, but it is

important to note that they are numerous. In this particular

Text-fig. 10.

Sagittal section through part of a proglottid of Ophidotenia naie.

Zm. Longitudinal muscular layer. sc.t. Subcuticular layer.

ut. Diverticula of uterus seen in transverse section.

feature the present species does not differ in any way from other

Ichthyoteniids. I do not think that the number of external

REPTILIAN TAPEWORMS, 33

pores is as great as that of the diverticula. But I cannot reckon

these pores up exactly. The actual pore itself is as a rule small,

but its position is rendered obvious by the gradual thinning to a

point of the cuticular layer on either side. Where the pore is

small the downwardly directed tube from the median stein of

the uterus is also slender and narrow.

Not infrequently, however, the pores are greatly enlarged, and

in such cases I have observed the orifice plugged with granular

matter exuding from the orifice. Very often too, and in association

with the stretching of the uterine pores, the ventrally directed

outgrowths of the uterus are much dilated, It might be suspected

perhaps that these large “pores” are simply the expression of a

rupture caused by the turgescence of the uterus, That this is

not the case is shown by the thinning of the chitinous cuticle

round the pore; it is clearly not torn across, as it would be were

it a question of a rupture,

If there were any doubt as to this tube with its paired

diverticula being the uterus, the existence of eggs within the tube

dissipates that doubt. These possess a tolerably thick shell and

they appear to resemble the eggs of Ichthyotenia varia. But in

addition to these eggs (which had segmented and were of course

really embryos) there were others with no apparent shell (text-

fig. 9, Cd). Attention has been called to the diverticula of the

uterus, which are small and whose walls are beset with numerous

long-stalked pear-shaped glandular cells. If it were not for the

comparatively wide lumen of these diverticula they might easily

be mistaken for shell-glands—so similar is their glandular invest-

ment to that of a shell-gland. I am disposed, indeed, to believe

that this is the actual function of the diverticula. For there are

to be noted here and there in the uterus round homogeneous

masses of a substance (text-fig. 9 A a, C p) which stains precisely

like the egg-shell, and which may very well be a seeretion of the

glandular cells referred to. It will be furthermore recollected

that this worm apparently does not possess a proper shell-

gland *.

The function of the uterus is thus enlarged, and it is in this

species not merely an egg-holder. ;

* It seems to me to be possible that an analogous state of affairs exists in

Mesocestoides. I have lately spent some time in studying a species of that genus

from a black Serval, which I do not feel able to place accurately but have regarded

as M. litterata. In the uterus masses of amorphous matter lie here and there among

the embryos. This substance is represented by Hamann (Zeitschr. f, wiss. Zool.

1885) as a nucleated tissue, with which I cannot agree. I think that we have to do

here, as in the species described above, with a secretion of uterine glands. The

hinder part of the uterus is regarded by Hamann as the equivalent for shell-gland

which he has stated to be absent in Mesocestoides. Zschokke, however (Mém. Inst.

Genéve, 1889), has asserted the presence of a real shell-gland in the normal position,

If a shell-gland is present in the species which I studied it is certainly very incon-

spicuous. I may point out, moreover, that in the genus with which the present

paper deals the shell-gland may be present or absent. In any case, the commence-

ment of the uterus in the Mesocestoides examined by myself had a glandular wall,

which quite possibly corresponds with the glandular diverticula of the uterus in the

species from the Indian Cobra.

Proc. Zoo, Soc.—1913, No. II. 3

34 DR. F. E. BEDDARD ON

It should be mentioned that the eggs were not in any way

aggregated into balls such as I have described in the species

Ichthyotenia varia.

I have finally to point out that although I have not come across

any description of external uterine pores in any Cyclophyllidean,

Spitlich* has described and figured in Tetrabothrium a cord of

cells passing from the uterus which may well be, as he suggests,

the remains of an external duct and pore. This comparison gains

some additional force by other points of likeness between Jchthyo-

tenia and Tetrabothrium insisted upon by Lonnberg T

§ On the Genera of the Family Ichthyoteniide.

Having surveyed the anatomy of a number of worms un-

doubtedly referable to this family of Tapeworms, it remains to be

seen how they should be placed in genera, whether in more than

one genus.

Relying mainly upon variations in the scolex and its suckers,

Braun {, in 1899, ailowed three genera—viz., Ichthyotenia,

Corallobothrium, and Crepidobothrium. Braun does not accept

Lonnberg’s § view that Zetrabothrium should be included in the

same family; neither does Ransom in his review of the Cyclo-

phyllidea ||, nor Fithrmann @ in his well-known essay upon the

Cestodes of Birds. It appears to me, however, that Lonnberg’s

Tetrabothrium trionychium is not congeneric with other species

referred to that genus. It has, in fact, the peripherally situated

Vitelline glands ‘of Ichthg pili with which it also agrees in its

alternating generative pores. In Tetrabothrium** the genera-

tive pores are unilateral and the vitelline gland Tr les in front

of the ovary.

While, therefore, there are certain points of agreement between

Tetrabothrium and Ichthyotenia, 1 do not think that they can be

placed together so closely as would be implied by. union in so small

a subdivision as a Cestode family. Linstow{t gave the name

of Acanthotenia to a small Ichthyotzniid from Varanus salvator,

which name was also accepted by Johnston §$§ in his account of a

worm of the same genus from Varanus varius. This generic name

is perhaps hardly accepted by Schwarz |||) in his recent monograph

upon reptile Ichthyoteniids. But the last-named author has

* Z ol. Jahrb. xxvit., 1909.

+ Centralbl. Bakt. u. Paras. xv. 1894, p. 801.

+ In Bronn’s ‘ Klassen und Ordnungen des Thierreichs, Bd. iv. p. 1706.

§ Tioe. cit.

|| “Lhe Tenioid Cestodes of North American Birds,’ Bull. U.S. Nat. Mus.

no. 69, 1909.

~ © “Die Cestoden der Végel,” Zool. Jahrb. Suppl.-Bd. x., 1908.

** Spatlich, “Studien tiber Tetrabothreen,” Zool. Jahrb. xxviii. 1909 ; and generic

diagnosis of Ransom.

++ Spatlich, Joc. cit. Taf. 28. fig. 25, dst.

+* Centralbl. Bakt. u. Paras. Xxxiil. 1903, p. 534.

§§ Proce. Roy. Soc. N.S. W. 1909.

i\|| “* Die TaNehonenien &e., > Inaug.-Diss., Basel, 1908.

REPTILIAN TAPEWORMS. 35

pointed out that in his opinion the reptile Ichthyoteniids do

agree together and differ from the fish Ichthyoteniids ina number

of points which he enumerates.

Schwarz comes to the conclusion that the reptilian Ichthyo-

teniids form a group which is characterised by the peculiar spiny

outgrowths of the egg-shell, by the fine spines upon the rostellum,

by the complicated coil of vas deferens and the protrusible cirrus-

‘sac, and finally by the arrangements of the transverse muscles.

It does not appear to me that all of these features are distinctive

of the reptilian Ichthyoteniids; for the coiled vas deferens is

found in other species of Jchthyotenia not from reptiles; the

fine spines upon the rostellum and early segments of the body are

at least not to be found in the species from the Cobra which I

describe in the present paper. The peculiar and delicate spines

which deck the egg-shell are figured and described by Schwarz

only in J. nattereri, The cirrus-sac does not seem to be protru-

sible in the way which he figures it, in the species of IJchthyo-

tenia which I have described in the present paper from various

species of Varanus. These characters are, in fact, not sufficiently

general or not sufficiently distinctive to forma group. It must

be admitted, however, that Schwarz suggests no name for his

group, and thus does not unduly emphasize it.

It appears to me, on the other hand, that we can separate off as

a distinct group of generic rank the species of /chthyotenia which

inhabit, not reptiles generally, but the lizard genus Varanus

only. Andif so, we must obviously term this genus Acanthotenia,

adopting v, Linstow’s name. The genus will be chiefly distin-

guished by the fine spiny covering of the scolex’ and anterior

segments, and, as I think, also by the character of the longitudinal

muscular layer. J am disposed to think that, as I have described

in several species in the present paper, the internal layer of

longitudinal fibres is found only in the anterior region of the

body. A third character is perhaps afforded by the com-

paratively large size of the rostellar region, which is apt to be

insignificant in the snake Ichthyotniids. Further than this [

do not think we can go at present, I propose the name of

Ophidctenia for a new genus to include at any rate the species

which I describe in the present paper from the Indian Cobra,

Its most salient anatomical character is of course afforded by the

structure of the uterus and by the ventrally and medianly placed

external orifices of the same.

This genus Ophidotenia also contrasts with Acanthotenia in

the lack of internal longitudinal muscles anteriorly and in their

presence posteriorly. ‘lhe inconspicuous. rostellar region. affords

another distinguishing mark. ‘In all of these characters, with the

possible exception of those offered by the uterus, this genus agrees

with other snake Ichthyoteniids. We cannot, therefore, as yet

attempt to fix the limits of this genus Ophidotenia, in fact not

until the structure of the uterus has been reinvestigated in other

Ichthyotznids in the light of my own discovery recorded here.

36 ON REPTILIAN TAPEWORMS.

While it is not apparently possible to distinguish as a separate

genus the two species which inhabit Amphibians, I am of opinion

that Lonnberg’s species from Zrionyx ferox will prove to be not a

Tetrabothrium or an Ichthyoteinia, but another genus closely allied

to the latter. At present the chief anatomical difference appears

to be the limitation of the testes to the middle of the proglottid.

Finally, I think that we must retain Crepidobothrium for those

Ichthyoteniids, inhabitants of Ophidia, which possess horseshoe-

shaped suckers.

§ Résumé of the principal new Facts.

It may be useful to abstract from the foregoing pages a short

résumé of the more striking new facts which I have been able

to ascertain :—

(1) Worms of a group belonging (in agreement with its usual

definition) to the genus /chthyotenia are more conveniently to be

removed and placed in the genus Acanthotenia of von Linstow.

These worms are found in several species of Varanus and are

limited to that genus. The same species does not occur in more |

than one species of Varanus, but a single species of Varanus may

contain more than one species of Acanthotenia parasitic in it.

(2) In some at any rate of the species of Acanthotenia there is

a layer of longitudinal muscles limited to the head and neck and

separating the medulla from the cortical region ; these fibres are

arranged in bundles. This layer disappears and does not occur

in the body behind the neck.

(3) It follows from the above statement that in the body

generally there is no demarcation between the medulla and

cortex.

(4) The existence of numerous ventrally situated openings of

the uterus in a tapeworm which is assigned to a new genus

Ophidotenia is a new fact so far as concerns the Cyclophyllidea

(= Tetracotylea).

(5) The diverticula of the uterus in Ophidotenia, instead of

being of a character similar to the median stem, are closely

beset with gland-cells and probably secrete the shell of the ova,

there being no shell-gland in this worm. There is here a possible

likeness to Mesocestoides, but the details are different.

(6) The mature eggs of Acanthotenia varia not only tend to

cling together in balls, as has been described in other species

of the genus, but a series of distinct eggs are enclosed also,

in many cases, within a common sheath, which suggests a division

of one original egg.

(7) A vemarkable abnormality is recorded in the same

species, where in one proglottid the male and female ducts open

on to opposite sides of the body.

IPAS. Aas dey, I.

West, Newman lith.

_H.R.Hogg del. -

FALKLAND ISLAND SPIDERS.

ZS. 1913) Pies

Wiley e

nore (ee

H.R Hogg del. West,Newman lith.

FALKLAND ISLAND SPIDERS.

ON FALKLAND ISLAND SPIDERS. 37

2. Some Falkland Island Spiders.

By H. R. Hoge, M.A., F.Z.8.

[Received October 8, 1912: Read November 12, 1912. |

(Plates I. & II.*)

INDEX.

Page

Systematic :—

LNRECHS QHMGDEHI Ig SVs We non coccegapdens sonecaccabncadsoasadoecnese BY

ZURGTOCIS GUODOGGIPs Fo Ds cop can aovene accoos2esase poo coosouaaneosonoann OL)

Tetragnatha inswlata, SP. MW. ...... 0. .ce cee coe veceeccseeessevseces 41

IER HISCE, COULMUCEE Fa Mo Gonceosc0 cd enseoosaedasaeo sconcvondeonvacees 22

Hinmenomma falklandica, sp. i. ... 2.2.00... .c. eee veeseeeee cence eee 45

LEQUPICUS SUGMAEMS, SDS Ws” roaccesen dvb odd caehes cso cooasacnsesanesescecn CAB

SOCUB COU EAGT SDs Bs 20 000000 son cee soo ede 07666009 nbnw7a ban senOAE 48

The small collection of spiders from the Falkland Islands

described below was brought to this country by Mr. Rupert

Vallentin, who has been good enough to afford me the oppor-

tunity of examining them.

Connected generically with Patagonia and the Magellanic

Region, they are all apparently new species. Gathered as they

have been over a period of two years, they comprise only six

species of spiders and one Opilio, of Sorensen’s genus Sadocus,

with a fair number of specimens of some of the species.

Three of these are Argiopide which are easily conveyed by the

wind, and supply no argument for a former land connection

between these islands and the mainland of America.

Family ARGIOPID &,

Group ARGIOPES.

Genus ARANEUS Clerck.

ARANEUS VALLENTINI, sp.n. (PI. I. figs. 3-3 ¢.)

The colour as seen in specimens taken from spirit is as

follows :—

Cephalothorax pale yellow-brown with longitudinal streaks of

long white hair, rather bare on the sides but a pale marginal

streak all round. Mandibles pale yellow-brown at the base, dark

brown at the apex. The fangs are dark brown at the base,

dingy yellow-brown at the points.

* For explanation of the Plates see p. 50.

38 MR. H. R. HOGG ON

co}

outer and lower portions being black-brown, fringes dark grey.

The sternum is black-brown, with upstanding grey hair.

The coxe of the lees are br ieht yellow-brown, the femora reddish

brown with pale grey hair, the patelle black- brown, tibiee yellow-

brown with dark band at fore end. Metatarsi and tarsi darker

yellow, with brown rings and dark brown spines and bristles.

The abdomen is pale yellow-brown on the back and sides, with five

pairs of dark muscle-spots on the former, each side of a scalloped

pale yellow longitudinal streak r eaching from the base to the

posterior end. ‘On the under side in front of the genital fold is a

wedge-shaped dark streak, narrowing posteriorly, with a narrow

dark streak on each side and smaller dark streaks nearer to the

rear outside the latter again. ‘The spinnerets and epigyne are

black-brown.

Mr. Vallentin has furnished me with a coloured drawing of

the specimens when first taken. This shows a greyish-green

cephalothorax and bright emerald-green abdomen with white

longitudinal streak. The femora ave bright crimson, the other

joints yellow with brown rings.

The cephalothorax is longer than broad, rounded at the sides,

convex, with the cephalic part well raised and distinctly separated

by lateral depressions from the thoracic part. The hairs are long,

pointing forward, and laid in streaks on the cephalic part ; on the

thoracic they point downwards on the sides and upwards on the

marginal streak. From the rear margin as far up as the end of

the pars cephalica isa rhomboidal area bare of hair ; this, however,

is hidden by the overhanging abdomen.

The rear row of eyes is slightly recurved seen from above, but

straight from in front. The front row strongly recurved from

each position.

The four median eyes are raised ona distinct prominence which

overhangs the clypeus. ‘They ave equal in size, the rear pair a

diameter apart; the front pair, 2 diameters apart, are 14 diameters

fromtherear. The side eyes are similarly situated on rather large

prominences, and are about half a diameter from each other. The

clypeus is about 14 diameters of the front median eyes, but curves

under the eye-space to the insertion of the mandibles, and vertically

is not so much.

The mandibles are slightly convex at the base but thence rather

straight to the front edge. A few straggling hairs at the base and

along the inner edge. The fangs are long and powerful. There

are four teeth on the outer margin of the falx-sheath, the third

one being stout and twice as long as the others; on the inner

margin are three equal teeth of moderate length.

The lip is broader than long, oblong at the base to about half

of its height, thence sloping toa point anteriorly. This part is

all pale. The mamille are roughly triangular, broadest in front,

with very broad pale margins.

The sternum is a broad shield-shape, slightly convex, hollowed

The lip and maxille have broad margins of pale yellow, the

FALKLAND ISLAND SPIDERS, 39

in front, with rounded projections at each fore corner. It narrows

to a point posteriorly, where there are two little round bosses, one

each side; in front of each coxa it is incuryed, and there isa

raised hump.

The legs are moderately thick in the femoral, patellar, and tibial

joints, the latter being thickened at the anterior end; the meta-

tarsi and tarsi are cylindrical and much finer. On tibia 1 and 2

are thick upstanding bristles, and a single row of six upstanding

spines on the under side. There are about five pectinations on

the superior tarsal claws, thin at the base but thickened anteriorly.

The femoral joint of the palpi is short; the tibial joint is longer

than the patellar, and there are upstanding bristles and spines on

the distal joint. There are several pectinations on the palp-claw.

The abdomen, which overhangs the cephalothorax as far as the

lower end of the cephalic part, is ovate, convex, rounded in front

but slightly pointed. It is widest at about one-third of its length

from the base, where there are small corner humps which disappear

In some specimens, thence narrowing to a blunt rounded point at

the rear end. There are two pairs of large muscle-spots followed

by three pairs smaller and less distinct.

The epigyne has a scape furnished with two prominent lobes

on the lower margin, and a specially long appendix of the

A. productus type, 2mm. long, which reaches more than halfway

to the membranous base of the spinnerets. The latter are large

and well developed, and of the normal type.

The hair covering is short, fine, and downlying, but there is a

ring of longer bristly hairs round the base of the spinnerets.

The measurements (in millimetres) are as follows :—

Long. Broad.

3y *

Cephalothorax... 44 ee a

Abdomen......... i 10°

Mandibles ...... 2

Trochanter Patella Metatarsus

Coxa. &femur. &tibia. & tarsus.

JUSS Re Re, Ie 3 D 54 See

2. 4 5 5 ew IG

Sue at ula 2 = 108

4, lt 5) 5 ee lot

Ballot ict eas yse nee 2 2 2 Z 0 = 62

There are numerous specimens of females but no males. The

species is said to be common about the Camp. There was a

green-coloured egg-sac with the specimens but it was torn out

of shape.

ARANEUS GLOBIGER, sp.n. (PI. I. figs. 4-4.)

Male. Cephalothorax dingy yellow-brown, lightest on the

cephalic part, darker on the thoracic, with a marginal stripe

again paler. The hairs are long, forward pointing, yellowish

AOL > MR. H. R. HOGG ON

grey. The mandibles are brown at the base and pale yellow

brown anteriorly. Fangs pale yellow-grey at the point, browner

at their base.

The lip and maxille have a margin, equal to half their respective

breadths, of pale yellow-grey, the remainder black-brown. The

sternum is black-brown with yellow-grey hairs. The legs and

palpi are bright yellow, with darker bands, yellowish-grey spines

and hair and brown bristles.

The abdomen, both on the upper and under sides, is dark brown

irregularly mottled with paler yellow spots not forming any

special pattern, and grey hair.

The cephalothorax is longer than broad, only slightly convex,

straight in front, and is rounded at the sides and rear; it has a

short longitudinal fovea behind the cephalic part leading into

a broader transverse depression. The cephalic part is bounded by

long shallow depressions, and is rather thickly covered with bristly

downlying hair all pointing forward ; the thoracic part is sparsely

covered with shorter hair, with the exception of a marginal stripe

which is again more closely covered.

The rear row of eyes is straight or, seen from in front, slightly

procurved. Both the median four and each side pair of eyes are

on one raised prominence overhanging the clypeus in front; the

median are equal in size, a diameter apart behind and two

diameters in front; the laterals are adjacent, the rear rather larger.

The clypeus is narrow.

The mandibles are short and flat, with short curved fangs.

The lip is broader than long, curved at the sides, pointed at

the apex, and less than half the length of the maxille. The latter

are broadest at the top, curving rather away from where, on the

inner side, they meet the lip, and have pale margins half their

own width.

The sternum is shield-shaped, convex, hollowed from the two

front corners, broadest in the middle, pointed at the rear where

the rear coxe are contiguous. The hairs are long, rough, and

rather upstanding.

The legs are thickly covered with downlying hair, upstanding

bristles, and numerous short thick spines. The tibie of the first

pair are dilated at the anterior end, being there twice the

thickness of the metatarsi, and are furnished with short spines.

The tibie have a single row of short spines on the outer side,

besides numerous spines on the inner and under side. The palpi

are short and slight, with a remarkably large development of bulb

stylus and other organs, covering the whole front of the man-

dibles and standing out at each side; a iong elbowed stylus has

a projection at the bend. The tibial joint is short and globular,

with two Jong bristles protruding therefrom.

The abdomen is roughly quadrangular, longer than broad,

narrowed but rounded at the front and rear ends, with angles at

the broadest part not quite one-half of the distance from the

base to the rear; at these angles are slight rounded prominences

FALKLAND ISLAND SPIDERS. Al

and two smaller ones at the rear end. It overhangs the cephalo-

thorax almost to the end of the cephalic part.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax ... 4 { he in front.

INOGIOWNETN gs osc0ns00ne 4} 3

Mandibles ......... 13

Pat. & Metat.

Coxa. Tr. & fem. tib. & tars.

Tesst 2, (ons A A fe aigyh

a eee eo Mie 2a) ae

a 2 24 2 Qf = 74

gt Saath Ry 9 Rye ane

IE] Ov dimea eae eens 4 1 z 2 — 4%

There is one male specimen.

An undeveloped female, similarly coloured, has the rear and

side prominences much more prominent, and a pale median

longitudinal streak down the abdomen on the upper side.

Subfamily TETRAGNATHINA.

Genus TerracnaTHA Latreille.

TETRAGNATHA INSULATA, sp. n. (PI. II. figs. 6 a—6 e.)

Female. The cephalothorax, mandibles, legs, and palpi pale

yellow, the hairing on the latter dark grey; the eyes black, the

laterals on biack tubercles and black patches behind the rear

median. The sternum nearly black at the edges, but yellow in the

central streak, with dark grey hair. The abdomen is uniformly

dark yellow-grey all over, with dark grey hair.

The cephalothorax is longer than broad, the cephalic part being

raised and divided off from the thoracic by distinct depressions.

The eyes are all equal in size. Both the front and rear row

are recurved, the laterals being as far apart from each other as

the front and rear medians. The rear median are 14 diameters

apart and 23 from the laterals. The front median are one

diameter apart, 13 diameters from those of the rear row, and

3 from their laterals. The front median eyes are one diameter

distant from the margin of the clypeus.

The mandible in the female has on the inner margin of the

falx-sheath one tooth at the upper end and two teeth at the

jiower end, about the level of the top of the maxille; on the

outer margin there is one tooth at the upper end, and two at

the lower end below those on the inner margin.

In the male there are eight teeth on the inner margin of the

falx-sheath. Near the upper end of the outer margin are two

much larger teeth, of which the second is the longest, and then

492 "MR. H. R. HOGG ON

after an interval, five about the same size as those on the inner

margin but reaching farther down the edge. On the outer side

near the fore end is a projection bifurcated at its anterior end.

The sternum, lip, and maxille are of the normal type.

The abdomen slightly. overhangs the cephalothorax, it is

broadest at about one-fourth of the distance from the base, thence

tapering to the posterior rounded end. On the under side in the

female is a raised fold of the skin reaching from the centre of

the epigyne to the spinnerets.

The measurements (in millimetres) are as follows :—

Male.

Long. Broad.

Cephalothorax ... 2m \ ' 1 in front.

Abdomen............ 5 1 2

Mandibles ......... 2

Pat. & Metat.

Coxa, Tr. & fem. tib. & tars.

Megs wee 3 64 Se eas

2 3 4A 4A 5 = 143

Seige 94 2 a 4

4 a 43 4 1 = 134

Aallyoizn hese ae. eile 3 13 2 2 = 32

Patella shorter than tibia.

Female,

Long Broad

Cephalothorax Pal { a in front

Abdomentesseee eee 7 24

Mandibles ......... le

Pat. & Metat.

Coxa. Tr. & fem. tib. & tars.

Thesisie. cin ee 6 63 ih = 202

2 3 4 Ald 4, = Igz

aera: 4 2 2

Suen ss 23 2 = i

Ales 98 5 4 4 = 133

Palle eh. rst 2 if 13 les 5

Patella much shorter than tibia.

Family CLUBIONIDA.

Group CLUBIONEA.

Genus Puitisca E. Simon.

PHILISCA COLULATA, sp.n. (PI. IT. figs. 5-5 d.)

Female. The cephalothorax is dark yellow-brown in the centre

of the cephalic part, dark brown at the sides. The thoracic part

FALKLAND ISLAND SPIDERS. 43

is dark yellow-brown, with a narrow darker brown median longi-

tudinal stripe in the anterior portion, and a dark brown marginal

stripe; the downlying hairs are yellowish grey, with a few

longer upstanding brown hairs over the posterior part.

The mandibles are black-brown, with upstanding grey hairs

along the inner side. The fangs black-brown at the base, become

bright red towards the anterior end. The lip and maxille are

yellow-brown with paler edges, brown hairs on these and the

sternum, which is much darker, almost black-brown.

At the base of the abdomen, on the upper side, are two pale

yellow areas divided by a longitudinal brown median streak, and

bounded by brown at the sides and right in front. The median

streak reaches to about halfway, where it spreads out into three

chevrons followed posteriorly by a uniformly mottled brown area.

It is thinly covered all over with yellow-grey downlying hair

and brown upstanding bristles.

On the under side a darkly mottled wedge-shaped area broadest

in front extends the whole distance from the base to the

spinnerets ; this is bounded by pale yellow-brown which extends

to the dark brown sides.

Legs and palpi dingy yellow, with dark grey hairs and spines.

The cephalothorax is longer than broad, convex, straight in

front, slightly rounded at the sides, with no side depressions but

a quite short median longitudinal fovea at the upper part of the

rear slope.

Hyes. The rear row of eyes is slightly procurved ; they are

about equal in size. ‘The median are two diameters apart and one

diameter from their respective laterals. The front row is shorter

and straight. The laterals are as large as the rear eyes and

slightly more than their diameter therefrom; the median, two-

thirds the diameter of the laterals, are their own diameter apart

and one-half of the same from the side eyes. The area covered

by the four median eyes is broader than long.

The clypeus is as broad as the front side eyes.

The mandibles, rather stout and broad, are kneed at the base,

thence descending perpendicularly. Fangsrather long and strong.

Two teeth on the inner margin of the falx-sheath, and one longer

between two small on the outer margin.

The maxille are upright, straight on the inner side, broadest

at the upper margin, thence curving inwards to the insertion of

the palpi. There is a long tuft of bristles on the upper part

of the truncature followed by shorter bristles to the lower end

of same.

The lip is rather longer than broad and more than half the

Jength of the maxille. It is narrowed at the base, just above

which it is widest, narrowing to a slightly hollowed truncate

anterior end.

The sternwm is convex, 13 times longer than broad, straight

in front, whence it widens out to the middle and then again

narrows to a point between the not quite contiguous rear coxe.

44 MR. H. R. HOGG ON

It is hollowed for each coxa, between which and itself is a

marginal space. The base of the lip joins it on the upper edge

but not the maxille. The rough upstanding hair thereon is

thicker near the edge than in the centre.

The abdomen is longer than broad, truncate in front, broadest

about the middle and pointed at the rear end, where the

spinnerets are terminal. It is moderately thickly covered with

short upstanding and finer downlying hair. The ventral tracheal

fold is rather close behind the spinnerets.

The inferior spinnerets are one-jointed and conical, close

together and rising from a membranous base; the superior are

longer, flatter and broader, with a short pointed second joint.

There is a large well-formed colulus in front of the inferior pair.

The legs are moderately stout, the tarsi and metatarsi cylin-

drical. On the two latter joints of all legs isa scopula with claw-

tufts of flat pointed bristles. The two claws have about five pectin-

ations. There are numerous long spines on the upper side of the

femora and on the tibial and metatarsal joints.

The palpi are rather short and fine, the femoral joints incurved,

and spines on the tibial and distal joints. The tibia is longer

than the patella.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax... 4A { me a leon,

Abdomen......... 6 33

Mandibles ...... 12

Pat. Metat.

Coxa. Tr. & fem. & tib. & tars.

Be Sic cas Alle ] 33 oF 33 = 103

Me 1 3} 3f 3 = 107

eles 4 DO Mis cas

Palipias isso ues z ls Va I 43

This would appear to belong to M. Simon’s genus Philisca

from Tierra del Fuego etc. It differs from P. navarinensis Tull.,

to which it approximates, in having the side eyes quite 12 dia-

meters apart instead of 1 diameter; in having the rear median

twice as far from one another as from the side eyes instead of

equidistant, and the row procurved instead of straight; the front

median decidedly less than three-fourths of the laterals; the

second pair of legs as long as the first, instead of shorter ; at the

rear end of the dark median line on the back three black arrow-

headed cross markings instead of testaceous spots; and the fourth

pair of legs longer in proportion to the others ; the epigyne is also

apparently different.

One female found under a stone in Camp.

FALKLAND ISLAND SPIDERS. 45

Family AGALENID#.

Subfamily CyBa#1na.

Group CyBHEs.

Genus EMMENomMaA KH. Simon.

EMMENOMMA FALKLANDICA, sp. n. (PI. I. figs. 2-2 6.)

Female. The cephalothorax is yellow-brown, darker in the

lateral depressions, black-brown over the eye-space, rather thickly

covered with yellowish-grey hair. Mandibles, lip, and maxille

bright yellow-brown with dark grey hair ; fangs dark red-brown ;

sternum somewhat darker, with a brown median patch and

upstanding brown bristles towards the rear.

The abdomen is a greyish yellow, with a pair of black rectan-

gular longitudinal patches on each side of the base and a median

black quadrilateral, broadest in the middle and pointed at each

end, reaching to rather more than halfway ; on each side of this

again are longitudinal black patches and two more similar at

the rear end. The under side is dingy yellow all over, with pale

yellow-grey hairs and brown upstanding bristles.

The legs are rather bright yellow, with dark grey hairs and

spines and brown upstanding bristles on the coxz and under side

of the femora, and pale grey rings on the femoral, tibial, and

metatarsal joints.

The cephalothorax is longer than broad, with about four well-

marked depressions on each side, and a long and deep longitudinal

fovea reaching from the rear slope to the cephalic part.

The rear row of eyes is recurved; the tops of the median pair,

which are three-fourths of their diameter apart, reach to the

line joining the lower side of the laterals, which are rather more

than their diameter away. The front lateral eyes are 14 times

the diameter of the rear median, 11 times their diameter from

one another, and about 14 times that distance from the rear

laterals. The front median are one-fourth the diameter of, and

in a line with, the upper edge of the laterals.

The mandibles are stout and strong, kneed at the base, where

there are long upstanding bristles; at the apex there is a fringe

of downlying hair.

The lip is about as broad as long, straight in front, and

broadest one-third of its height therefrom. The mawille, slightly

bending over the lip, are rounded anteriorly, where they are

broadest and about twice the height of the lip.

The séernwm is shield-shaped, 14 times as long as broad,

hollowed in front and pointed at rear end between the not quite

contiguous last pair of coxee.

The abdomen is oval, truncate in front, the hair covering

scanty, rather short and fine.

46 MR. H. R. HOGG ON

The legs are only moderately stout, tarsi and metatarsi cylin-

drical; long stout spines on the metatarsi and tibie of all legs.

A brush of short bristles, not amounting to a scopula, on the

under side of the tarsi.

The superior claws are long and rather straight, with a few

pectinations at the basal end only.

The tibial joint of the palp is twice as long as the patella, and

the distal joint is furnished with spines and bristles.

The measurements (in millimetres) are as follows :—

Loug. Broad.

Cephalothorax... 4 | 31 ao

‘Abdomen. ssee: 6 32

Mandibles ...... 2

Pat. Metat.

Coxa. Tr. & fem. & tib. & tars.

ees syne i el 33 AD 8d =e lie

2 1 37 4 a. 113

3) i 34 ore 4 = 11?

4, 1 4. 4} 6 = lod

LEP ay ee SRG eeeadonee 3 }3 ee els ae 43

This agrees with M. Simon’s genus Hmmenomma from Cape

Horn, except as to its median eye area. It differs from H#. ocu-

latum in this being only 14 times as long as broad instead of

3 times the width.

One female only.

Family THOMISID.

Subfamily PHiLopRoMIN#.

Genus Pretrricus EB. Simon.

PErRICUS SIGNATUS, sp. n. (PI. I. figs. 1-1.)

Female. Cephalothorax black-brown mottled with dark yellow

brown, and scantily covered with downlying flat white hairs ;

a thick band of same across the front of the clypeus.

Mandibles black-brown, smooth at the base, with a few brown

bristly hairs towards the anterior end. Fangs dark yellow-brown.

Sternum, lip, and maxille patchy dingy yellow-brown, with rough

white hair. Coxe of legs dingy darkish yellow, with dark brown

on the under side of the femur, patella, and tibia, and downlying

dark hairs and brown upstanding bristles and spines. Metatarsus

and tarsus paler yellow, with thick whitish scopula on the latter

and anterior part of the former.

The upper side of the abdomen is b!ack-lLrown with down-

lying flat white hairs and upstanding brown bristles; a dark

longitudinal median stripe extends from the base to about two-

FALKLAND ISLAND SPIDERS. AT

thirds the length, and there are four dark stripes perpendicularly

down each side at the rear end. Underneath, dingy dark yellow-

brown with thinner whitish downlying hair.

The cephalic part of the cephalothorax, which is only slightly

longer than broad, is convex, thick, and square anteriorly,

separated by a broad shallow depression from the thoracic part,

which is also convex and slopes evenly to the margin.

The rear row of eyes is straight, or viewed from above slightly

recurved, but, owing to the curve of the headline, slightly pro-

curved from in front; it lies on the front margin of the upper

part. Krom this point the forehead falls perpendicularly, so that

the recurved front row, two-thirds the length of the rear row,

looks straight forward. The laterals of each row are equal in size,

each on separate tubercles. The eyes of the rear row are

equidistant, the median slightly smaller than the lateral. The

front median pair are half the diameter of the side, three of their

diameters apart, and two of the same from their laterals. A line

drawn across their upper margin lies near but well below that

across the lower margin of the laterals. The clypeus is as broad

as the length of the quadrangle of median eyes, which is twice as

long as its breadth.

The mandibles are conical, perpendicular, smooth at the base,

with patches of bristly hair across the lower end, and on the smooth

under side are a few scattered bristles. The fangs are short and

strongly curved.

The lip is broader than long, widest at the base, narrowing to,

and rounded at, the anterior margin. It reaches to more than

half the height of the mandibles, which slope over it. These are

rounded anteriorly, and the outer margin curves continuously

to the insertion of the trochanter of the palps.

The sternum is convex, of a broad shield-shape, almost as

wide as long, truncate in front, and narrowing to a point pos-

teriorly between the not quite contiguous rear coxe.

The abdomen is ovate, broadest two-thirds of the distance

from its base, where it is rounded and has a median longitudinal

depression. The downlying flat hairs are slightly plumose, all

pointing with their heads to the rear.

The legs are moderately fine and even in length, the second

pair being only about one-fifth longer than the shortest of the

other three pairs, the fourth is slightly longer than the first and

third, but they do not vary much. The tarsal and rather thinner

metatarsal joints are cylindrical. There are two pairs of long

spines and one shorter on the under side of the tibia and meta-

tarsus of each leg. ‘They are rather thickly covered with the

same downlying flat pointed hairs as the cephalothorax and

abdomen, and a few upstanding bristles, also long upstanding

spines.

The scopule and claw-tufts on all feet are of the same type of

flat pointed, not spatulate, bristles, shorter and more upstanding

48 MR. H. R. HOGG ON

than those on the other joints. They are not divided longi-

tudinally as in Wiswmena. The inner of the pair of tarsal claws

has about three rather long pectinations on the shaft curving with

the curved point of the claw.

The palpi are short and fine, with hair and bristles as on the

legs, and spines on the tibial joint,

The measurements (in millimetres) of the largest are as

follows :—

Long. Broad.

lin fr

Cephalothorax... 32 {33 meee

Abdomen......... 5 35

Mandible......... ile

Pat. & Metat

Coxa, Tr, & fem tib & tars

Legs ates A ; fe C a = i

Sain 31 31 oo ll

how il 32 33 33) eee SVs

Ballon seas 4 1? 1 egal

This species conforms to M. Simon’s genus Petricus from

South America, and is closer to his P, nivews than to any of

Mr. Tullgren’s species, It differs, however, from the former in

its front row of eyes being very clearly recurved instead of slightly ;

the median are 3 diameters apart instead of one, and nearer to

the lateral than to one another instead of equidistant; the

lateral eyes of the front and rear rows about 4 diameters apart

instead of 14; the area of the median quadrilateral equally broad

in front and rear, instead of much broader in front. The front

pair of legs are longer and the others shorter in proportion to tne

cephalothorax than in P, niveus.

There are nine females from various districts, apparently the

same but differing in size of adults and brightness of colour.

OPILIONES.

Family GONYLEPTOIDA,

Genus Sapocus Sor.

SADOCUS VALLENTINI, sp.n. (Pl. II. figs. 7-7c.)

Carapace yellow-brown mottled with black-brown anteriorly,

nearly all dark brown at posterior end; eye-tubercle yellow in

middle, black-brown at each side.

Trochanters yellow; other joints yellow mottled with plack-

brown ; anterior ends of tarsi of rear two pairs yellow with yellow

FALKLAND ISLAND SPIDERS. 49

claws. Hair generally pale yellow-grey. The under side of the

abdomen is paler than the upper side, having more yellow and less

dark mottling.

The eye-tubercle is oval, twice as broad as long, the eyes being

at the extreme side edges, It is situated in the front half of the

cephalic part, the portion in front of it being its own width and

twice that distance behind.

The thoracic part of the carapace is slightly convex with a

distinct marginal suleus all round. There are small globular

tubercles in the median area of the cephalic part, but not extend-

ing to the sides, reaching from the eye-tubercle to the rear margin.

There are smaller sparsely scattered granulations on the second,

third, and fourth divisions of the scutum, and a pair of tubercles

on the latter. Behind this are four segmental divisions thickly

covered with tubercles, succeeded by a posterior broader marginal

area, also tuberiferous, which ends, in the female, in a median

pointed process as long as the rear trochanters. In what appears

to be the male thisis wanting. On the under side of the carapace

the flat coxal areas of the second and third pairs of legs extend

to a narrow sternal depression, those of the first pair to round

the mouth-parts. There is a small lower lip above the sternum

and movable chitinous maxille unconnected with the coxe of the

palps. A much larger upper lip is divided into two parts on

the under side by a muscular double wall. On the under side of the

tibial joint of the palp are three spinous processes on the outer

margin, and four on the inner. On the tarsal joint of same are

four spines alternately iong and short on the outer side, and three

on the inner side of the falx-sheath. The long, slightly curved,

movable fang is about the length of the tarsal jot. The man-

dibles reach up to about the anterior end of the patellar joint of

the palp. There are three equal teeth on the inside of the fixed

claw, one large between two smaller on the movable claw. The

coxal area of each of the fourth pair of legs is twice as broad as

the median portion of the thorax lying between them. They

extend along each side twice the length of the three anterior coxal

areas. At the posterior end of this coxaa chitinous process projects

from the outer corner as long as the breadth of the trochanter

in the females, and twice that length in the males. Behind this

is around convex boss.

At the posterior end of the median area, lying between the

fourth coxe, are the pulmonary apertures with oval convex lids,

-and on the inner side between these are a pair of oval hollow

depressions.

The three anterior pairs of legs are of about the-same diameter,

rather slender, all without spines but covered with short hair.

On the tarsal joint of the third and fourth pairs are two curved

smooth claws. Tarsus 1 has five joints; tarsus 2 eight; tarsus

3 and 4 six each. The trochantal joint of the fourth pair is as

long as broad and thickly covered with spinous tubercles.

Proc. Zoon, Soc.—1913, No. LV. ob

50 ON FALKLAND ISLAND SPIDERS.

The femoral joint is thickly covered with long chitinous

spinous processes, which in the male are twice as long as in the

female. ‘The patellar joint is enlarged and globular, the other

joints are without spines but thickly covered with hair.

The measurements (in millimetres) are as follows :—

Long. Broad.

: ‘

nee es } part of scutum... 2 3

Abdominal do. rikC Nepean 25) 5

Mianmdiblesiniine!spacaaceceleok ges y

Coxal Pat. & Metat.

part. Tr. & fem. tib. & tars.

Legs.... iL 3 3 2 a =e

2 ] 34 en —

oe 1 34 3 4 == te

4 3 4 33 5 = libs

Pali en: Beet Me ies fel 12 i 1 = 4+

This differs from S. vitellinosulcatus Sor. (believed to have

come from 8. America and described from a single female) in

having a single terminal process above the anal tubercle at the

posterior end of the abdomen, instead of a pair of quite small

ones; in having the anterior edge of the scutum straight instead

of two pointed processes in the middle; in having no spine on

the femoral joint of the palpi and no pattern on the back of the

abdomen, as well as in its smaller size. There is no process on the

eye-tubercle.

One male, 6 females and 2 non-adult, found among decaying

roots of ferns.

EXPLANATION OF THE PLATES.

Puate I.

Petricus signatus, sp. n., X 3. a.Eyes. 6. Lip and maxilla. c. Epigyne.

. Emmenomma falklandica, sp.n., X 2. a. Eyes. 6. Epigyne.

. Araneus vallentini, sp.n.,nat.size. a. Underside of abdomen. 4. Profile.

ec. Front of cephalothorax and eyes. d. Lip and maxille. e. Epigyne.

. Araneus globiger, sp.n., nat. size., g. a. Hyes. 6. Male palp from side.

c. Male palp from underneath. d. Tibia II, e. Coxa I.

= coor

Puate II.

Fig. 5. Philisea colulata, sp. u., X 2. a. Kyes. 6b. Mandible. c. Epigyne.

d. Spinnerets.

. a. Kyes of Tetragnatha insulata, sp. n. 6. Underside of mandible, 3.

c. Fore-end of mandible from above. d. Male palp. e. Epigyne.

- Sadocus vallentini, sp.n., X 2. a. Underside of carapace. 6. Mouth parts.

c. Mandible and palp.

“I co

WHIAUO) ais) Ol ish) WAAL SSO eIWASt GNI ESE SLiNshsy WSOUSHAI/ El” YL

‘cut 38 YI] [Sp Useap p

WE Neal SS UGIL Sal

—"

MR. G. A. BOULENGER ON NEW AFRICAN FISHES.

3. Descriptions of Three new Fishes discovered in the Gold

Coast by Wr: HiaGe ih, Spurcelli MEAS ZS: > By, Gi.

A. BouLrencer, F.R.S., F.Z.8.*

| Received October 28, 1912: Read November 12, 1912. ]

(Plate IIT.+)

INDEX.

Systematic : Page

IHRE SOUR Hy BOs Wo: coscesoossonesonsenaddecasocnecesso+o1 Ob

JE OUOUS TOLLCROSIOMI Gy BDo Ds edccccscsneosenannconossoneesncas Opll

T3ORTITES Sf OOFRAM G5 06 Wo cocondabaccodscnovandcccose coocuvss OH

Dr. Spurrell, who for some years has enriched our collection of

living animals with many examples of Reptiles from the Gold

Coast, has lately turned his attention to Fishes, and has presented

two small collections to the British Museum. Among them I

have found examples of what appear to me three new species.

These fishes are from the vicinity of Bibianaha, near Dunkwa,

between the watersheds of the Tano and Ankobra Rivers. Notes

on the fresh coloration have been supphed by Dr. Spurrell.

BARBUS SPURRELLI, sp. n. (PI. III. fig. 1.)

Depth of body 3 to 33 times in total length, length of head 33 to

32 times. Snout rounded, as longas eye, 33 to 34 times in length

of head, interorbital width 2? to 3 times; mouth subinferior ;

lips moderately developed ; two barbels on each side, posterior as

long as eye and twice as long as anterior. Dorsal IIT 8, equally

distant from centre or anterior border of eye and from root of

caudal, border feebly concave; last simple ray not enlarged, as

long ashead. Anal III 5, not reaching caudal. Pectoral + length

of head, reaching ventral; base of latter below anterior rays of

dorsal. Caudal peduncle 1 to 13 times as long as deep. Scales

radiately striated, 24-26 = 2 between lateral line and ventral,

10 or 12 round caudal peduncle. Greenish silvery, darker on the

back, the scales dark-edged or with a dark base; this dark base

often more marked on the scales of the lateral line, which may be

further obscured by black dots, forming an ill-defined dark

lateral band; fins greyish.

Total length 75 mm.

Several specimens.

This species comes very close to 4. ablabes Blkr.

BARILIUS MACROSTOMA, sp. n. (PI. III. figs. 2, 2 a.)

Depth of body equal to length of head, 3 times in total length.

Head 24 times as long as broad; snout pointed, projecting

beyond mouth, 12 times as long as eye, which is 44 times in

* By permission of the Trustees of the British Museum.

+ For explanation of the Plate see p. 43.

2 MR. G. A. BOULENGER ON NEW AFRICAN FISHES.

length of head, 14 times in interorbital width ; mouth extending

nearly to below posterior border of eye; no barbels; second

suborbital deep, extending posteriorly to vertical of posterior

border of eye; naked space between preoperculum and suborbital

about 7 diameter of eye. Gill-rakers few and very rudimentary.

Dorsal III 8, originating at equal distance from occiput and from

root of caudal; posterior third of its base above anal; anterior rays

longest, a little less than 3 length of head. Anal III 14, notched,

anterior lobe rounded. Pectoral pointed, § length of head, not

reaching ventral, which is much shorter and reaches vent.

Caudal crescentie when fully spread out. Caudal peduncle

1? times as long as deep. Scales with radiating strie, 52 ie 3

between lateral line and ventral, 16 round caudal peduncle.

Silvery, with 13 or 14 dark bars on the side of the body above

the iateral line; dorsal fin greyish, anal reddish, caudal red with

a black edge.

Total length 155 mm.

A single specimen.

This species is closely allied to B. senegalensis Stdr., and

B. loati Blgr., both of which occur in West Africa. It agrees

with the former in the extension of the dorsal over the anterior

third of the anal and in the larger eye, but differs in the longer

mouth and in the lower number of scales in the lateral line

(52 instead of 59-63); with the latter it agrees in the number of

scales in the lateral line, but the larger eye, the longer mouth and

the position of the origin of the anal with regard to the dorsal

suffice to separate it; and, finally, the number of scales in a

; ms 92-102\ 4-4: . -

transverse series (Ce instead of foc distinguishes it from both

2 309

its nearest allies.

FunpDuLus SPURRELLI, Sp. n,

Depth of body 4 to 43 times in total length, length of head

34 to 5? times. Head flat above; snout short and broad, shorter

than eye; mouth directed upwards; lower jaw projecting; eye

34 times in length of head, 13 times in interorbital width ; space

between eye and lip about + diameter of former. Dorsal 13-14,

originating at equal distance from head and from base of caudal,

longest (posterior) rays = to ?# length of head. Anal 15-16,

originating slightly in advance of dorsal. Pectoral nearly $ length

of head, not quite reaching base of ventral, which is small and mid-

way between end of snout and base of caudal. Caudal rounded in.

the female, subtruncate in the male. Caudal peduncle longer

than deep. 29-31 scales in longitudinal series, 24-26 round

body in front of ventrals; lateral line indicated by a more or less

complete series of pits. Male pale yellowish green, with numerous

narrow, often paired, vertical bars of dark carmine; sides of head

metallic green, variegated with carmiine; gular (branchiostegal)

region of a dark, rich blue; pectoral fin whitish, with an oblique

crimson streak, ventral with red tip; vertical fins grey, dotted

ON TREMATODE PARASITES.

with carmine and broadly edged with yellow or orange, the

yellow bands occupying the upper and lower fourths of the

caudal. Female paler, more translucent, at times pinkish ; fins

white, dorsal and anal dotted with carmine.

Total length 42 mm.

Several specimens.

Allied to /#. gardneri Bley.

This species will be figured in the forthcoming third volume of

the British Museum Catalogue of African Freshwater Fishes.

EXPLANATION OF PLATE III.

Fig.l. Barbus spurrelli.

2. Barilius macrostema.

2a. 8 Sy Head from above. Natural size.

4. On some Parasites of the Seoter Duck (G’dema nigra),

and their Relation to the Pearl-inducing Trematode in

the Edible Mussel (Mfytilus edulis). By H. LystEr

Jameson, M.A., D.Sc., Ph.D., and Wittisw Nicoxy,

Loe Dasress IMLID)s

{ Received October 18, 1912: Read November 12, 1912.]

(Text-figures 11 & 12.)

INDEX.

Systematic : Page

Gymnophallus edemie, nom. nov. ..................... BT

(GSO FUENIS SP uM orn he cya cen panic chet eat hed, sa Os ete Gant NOS

Ch TIGKCTPROMOPUS, Se Wass asocsucedsbbosbccasengecedaededaasun l)

GOUOPLEUs Spleen ees eeer eee eee etna ee roe 62

Of the many questions connected with the formation of pearls

in the common Edible Mussel (J/ytilus edulis) the identity and

life-history of the pearl-inducing organism is one of the most

important. It was shown ten years ago by Jameson (1902) that

the agent in this particular case is the larva of a parasitic

Trematode, which, instead of secreting a cyst of its own, as is

usual with such larvee, stimulates the mussel to form around it a

sac of epidermal cells. These cells possess the same physiological

properties as the outer shell-secreting epidermis, and eventually,

on the death of the Trematode larva, secrete conchyolin and

calcareous salts, which, deposited in concentric layers around the

remains of the worm, become the pearl. Attempts were made by

Jameson to trace the life-history of this parasite, but the diffi-

culties in the way of working out the complete life-cycle of

digenetic Trematodes are considerable, and the results obtained

by him in 1902 have not been accepted as entirely conclusive.

__ With regard to the parasite in Mytilus, the two main questions

to be solved were : (1) Whence does it come? and (2) Whither does

o4 DR. H. LYSTER JAMESON AND DR. W. NICOLL ON

it go? From a consideration of Trematode development in

general, it was obvious that these larve in Mytilus must have

passed a previous stage, as a sporocyst or redia, in some other

molluscan host, and it was equally obvious that they were destined

to become adult in some final vertebrate host, to be sought among

the animals that eat the mussel.

With regard to the earlier part of the life-history, Jameson

was struck by the occurrence in Zapes decussatus, which lives

associated with the pearl-bearing mussels in the harbour of

Billiers (Morbihan), Brittany, of sporocysts containing Cercarice

closely resembling those in JMJ/ytilus and differing from them

chiefly in size. Jameson subsequently found similar sporo-

cysts in a different situation in the common Cockle (Cardiwm

edule) on the pearl-bearing mussel-beds at Piel, Lancashire, and

these have since been rediscovered by Lebour (1906) and

Nicoll (1906). The occurrence of these sporocysts in Cardiwm is of

particular interest in view of the fact that the cockle, as a rule,

lives in close association with the mussel and is on that account

a not unlikely intermediate host for the mussel parasite.

Herdman (1903-6) failed to find this parasite in Piel cockles,

and seems to have doubted Jameson’s assertion that it occurred

there; but we have found it over and over again in cockles both

from the original station at Piel, and from the cockle-bed at

Foulney in the same neighbourhood.

Infection-experiments were undertaken by Jameson in 1901,

in order to prove the transference of the parasite from Tapes to

Mytilus, but although he claimed success in these experiments,

they are still open to the objection that some at least of the

mussels used were already infected before the experiment was

undertaken *.

Pending the results of further experiments, there are only

the structural characters of the two larval forms to go upon.

Morphologically there is a very close agreement between the

cercaria in Cardiwm and that in Mytilus. They both agree in

having the oral sucker not more than one-fifth of the body-

length, and in its being not more than one-third as large again

as the ventral sucker. In both the pharynx is comparatively

large. The differences which exist are small. For instance,

the oral sucker in the J/ytilus cerearia is relatively larger than

that in the Oardium cercaria; the ventral sucker is larger in

the former in proportion to the oral and it is slightly nearer the

posterior end of the body instead of, as might have been expected,

further forward. Again, the csophagus is shorter and the

diverticula longer. These discrepancies, although minute, are

sufficient to give pause in too hastily concluding that the two

forms belong to one and the same species.

With regard to the final host of this parasite, there can be little

doubt but that Jameson was right in supposing it to be one of the

* [Iam at present repeating these experiments, with the aid of a Government

Grant from the Royal Society.—H. lL. J.]

TREMATODE PARASITES. 55

mussel-eating ducks, such as the Scoter and the Hider; but the

form he figured as the adult in 1902 from the Common Scoter,

which he referred to Gymnophallus (Lecithodendrium) somateric,

and which we are now describing below under the name G’. edemie,

is certainly not the adult of the larva which occurs in Mytilus,

but is a much smaller species *.

In order to reconcile the small size of the larvee whicn he found

in the Scoter at Billiers in 1902 with the larger dimensions of

the larvee in the mussel, Jameson was obliged to have recourse to

an unlikely hypothesis. This was commented on by Odhner,

who suggested that Jameson had probably encountered a mixture

of two species, one probably being G. bursicola and the other a

new species of the same genus. As the present investigation will

show, the latter part of Odhner’s supposition was correct ; while,

with regard to the former, until further specimens from Billiers can

be examined, it is impossible to say whether the larger examples

observed by Jameson at that station were G. bursicola or G'. dapsilis.

Since the publication of Jameson’s paper, two species, namely

Gymnophallus bursicola Odhner and G. dapsilis Nicoll, have been

suggested as the possible adult form. Before the discovery of the

second of these species Odhner (1904) gave it as his opinion that

the probable adult was G. bursicola, a parasite of the Bursa

Fabricii of the Hider Duck (Somateria mollissima). At the same

time he stated that a larva identical with that in d/ytilus occurs

in Saxicava rugosa in the Arctic regions. At that time there

could be little question as to the probable correctness of his view,

for no other known species was so likely to be the adult of the

parasite in Wytilus. The discovery by Nicoll (1907) in the Scoters

of a second bursicolous Gymnophallus closely resembling, yet

distinct from, G. bwrsicola, showed that Odhner’s conclusions

could not be accepted without further study, for G. dapsiti.

appears to have even stronger claims than G. bursicola to be

considered the adult of the larva in the Piel pearl-bearing M/ytiliy.

The two chief facts in favour of such a view are that the sizes

of the suckers and the position of the ventral sucker, in the

Piel pearl-inducing Trematode, correspond more closely with

those in G. dapsilis, while the firm brittle consistency of the

body reminds one more of G. dapsilis than of the softer G. bur-

sicola. The position of the testis in relation to the ventral

sucker is also a feature of some moment. It is, of course,

quite possible that two closely allied Trematodes are concerned

with pearl-formation in Mytilus—the one derived from Tapes,

the other from Cardiwm,—and it may be that the adults of these

two species are respectively G'. bursiccla and G. dapsilis.

Jameson foresaw that it would be necessary to prove in some

* [I distinctly remember the occurrence of asmall number of larger Gymnophalli

in the Billiers @demie, though it did not occur to me at the time that more than

one species might be present. I referred them all to the only species then known to

me, G. somaterie.—H. L. J.

+ We have not so far had an opportunity of re-examining the larvw found in the

Billiers mussels.

56 DR. H. LYSTER JAMESON AND DR. W. NICOLL ON

conclusive fashion the correctness of his views in regard to the

life-history of the pearl-forming Trematode in Mytilus, and

towards that end an endeavour was made in 1901 to perform

feeding-experiments with a Pochard (fuligula ferina), the only

likely diving duck which was available at the time. These

experiments were unsuccessful.

It was in continuance of this work that the present investigation

was undertaken. Arrangements were made* to obtain some live

Scoters (Hdemia nigra) and have them fed upon mussels infected

with the pearl-inducing cercariez. Three Scoters were pur-

chased on our behalf by the Zoological Society from D. G. Schuijl,

of Rotterdam, who stated that they were all caught on the

Zwarte Water, in the province of Overijsel. In addition, four

dead Scoters, stated to come from the same locality, were kindly

sent to us for examination by the Superintendent of the Brighton

Aquarium. With one exception, these four birds were found

to be heavily infected with intestinal parasites, including a

considerable number of Gymnophalli. These will be referred

to later. Of the three live Scoters, which were housed in the

Society’s Gardens, one died shortly after arrival, and was found

to harbour numerous specimens of T'ocotrema concavum and

immature Cestodes in its intestine. The second Scoter was

removed to the Lister Institute with the object of making a

feeding-experiment, but it died within 24 hours. Its intestine

contained only afew immature Cestodes. The third was kept from

8th to 24th December, 1911, when the experiment was started.

It was fed at intervals with mussels from the beds at Foulney,

near Piel, which were infected with the pearl-inducing Gymno-

phallus. Altogether about 1000 mussels were given toit. Check-

examinations of samples of these mussels showed that out of

78 mussels 32 were infected, the number of live Gymnophalli

being 64. On February 27th, 1912, the bird was killed and

thoroughly examined. The only parasitic worms present were a

few immature tapeworms in the intestine and two specimens of

Metorchis xanthosomus in the gall-bladder.

It is difficult to draw any satisfactory conclusion from the

result of this experiment. At first sight the most obvious in-

ference would appear to be that the larva in Mytilus does not

become adult in @demia, but in some other host. This inference,

however, is not without objection, for, quite apart from the

evidence furnished by the close resemblance between the parasite

in the mussel and in the duck, other factors may require to be taken

into consideration—for instance, the somewhat unnatural mode

of feeding, the unsuitability of the season, the effect of cap-

tivity, and so forth. In this connection, the complete absence

of other Trematodes from the intestine is significant, as suggest-

ing that the intestine may possibly have been cleared of parasites

as a result of digestive derangements following on change of

* [Thanks to a grant from the Government Grant Committee of the Royal

Soeiety.—H. L. J.|

TREMATODE PARASITES, 57

food. It is, however, useless to speculate on these matters, and

for the present we are content to record the negative result of

this particular experiment.

Our main object in submitting the present communication is to

give an account of the new species of G'ymnophallus which were

encountered in the course of examining the Scoters, and also

to note the occurrence of a few other Trematode parasites which

have hitherto not been recorded from this lost. Nicoll (1907)

gives the following list of Trematode parasites from Cdemia

nigra examined at St. Andrews :—

Gymnophallus dapsilis Nicoll. Bursa Fabricii.

Psilostomum brevicolle Creplin. Intestine.

Spelotrema pygmeum Levinsen. Intestine.

The following have been recorded by other authors :—

Psilochasmus oxyurus Creplin. Intestine.

Echinostomum revolutum Froelich. Intestine.

Monostomum sp. Respiratory tract.

Strigea tarda Steenstrup. Intestine.

In the course of our examination we found the following :—

Psilostomum brevicolle Creplin. Intestine.

Tocotrema concavum Creplin. Intestine.

Levinseniella brachysoma Creplin. Intestine.

Metorchis xcanthosomus Creplin. Gall-bladder.

Gymnophallus dapsilis Nicoll. Intestine.

Gynnophallus bursicola Odhner Bursa Fabrici.

Gymnophallus edemice, sp. n. Intestine.

Gymnophallus affinis, sp. 1. Intestine.

Gymnophallus macroporus, nom. noy. Intestine.

Gymnophallus ovoplenus, sp. n. Intestine.

Catatropis verrucosa Froelich. Bursa Fabricii.

This makes a total of 15 distinct species from this single

host, of which 9 are here recorded for the first time. Such a

wealth of varieties of Trematode parasites in a single host is

rather remarkable *.

The new species which we are describing were all obtained from

the Scoters supplied by the Brighton Aquarium. The birds

originally came from the same quarter as those purchased through

the Zoological Society, and it is rather striking that they should

have been so heavily infected, while the others were almost free

from Trematodes.

GYMNOPHALLUS @DEMI&, nom. nov.

= Lecithodendrium somaterie Jameson, 1902 {ex parte).

This was the least common of the four species, and occurred in

* [To the list must be added Paramonostomum alveatwn Mehlis, which was

obtained from a Scoter ((Hdemia nigra) which died in the Society’s Gardens on

Noy. 19th, 1911.—W. N. |

58 Dit. H. LYSTER JAMESON AND DR. W. NICOLL ON

only two of the birds examined. It is the adult form depicted by

Jameson (1902, pl. xvi. fig. 11) and to which the greater part of

his description (pp. 159-160) applies.

The body is somewhat flat and oval, with pointed extremities.

Tts length is -19—-25 mm. and its maximum breadth is usually a

little more than half the length. The cuticle is entirely covered

with minute spines. The oral sucker measures *030-"037 mm. in

diameter and the ventral sucker ‘024-032 mm., the ratio being

approximately 6:5. The ventral sucker is situated at a distance

of ‘11-15 mm. from the anterior end, the distance being about

three-fifths of the body-length.

The pharynx is contiguous with the oral sucker and is about

°017 mm.long. It is followed by a narrow cesophagus which may

be nearly twice as long as the pharynx. The intestinal diver-

ticula diverge fairly widely and barely reach the middle of the

body. The excretory vesicle consists of a very wide main stem

and two long branches, which nearly reach the oral sucker.

The testes lie some distance behind the ventral sucker, but

nearer it than the end of the body. They are small and irregu-

larly globular. The vesicula seminalis is situated over the left

side of the ventral sucker, usually a little in front, but sometimes

extending to the posterior border of the sucker. From it issues

a short pars prostatica, of the usual G'ymnophallus type, opening

on the anterior lip of the sucker.

The ovary lies on the right side, on the same level as the

ventral sucker, and it is nearly as large as the sucker. The yolk-

glands are situated over the anterior half of the ventral sucker.

They are somewhat irregular. The uterus lies almost entirely in

front of the ventral sucker, and extends forwards almost to the

oral sucker, filling up most of the anterior part of the body. The

eggs, which usually number 30-100, measure -018—-020 mm. in

length and :013-—-014 mm. in breadth.

GYMNOPHALLUS AFFINIS, sp. n. (Text-fig. 11.)

This species, although about the same size as G. wdemie, is at

once distinguished from it by the large size of the oral sucker and

by the disposition of the uterus. It measures *2—25 mm. in

length and -11—-13 mm. in breadth, the body being flat and oval.

The ends are not usually so pointed as in the previous species.

The oral sucker has a transverse diameter of -065--085 mm.

The ventral sucker is very much smaller, measuring only -030—

-045 mm., and the ratio is usually about 2:1. The ventral sucker

is situated -16—18 mm. from the anterior end, 7. ¢., a little more

than two-thirds of the body-length.

The pharynx, which is contiguous with the oral sucker,

measures ‘015—018 mm. in length by :012-017 mm. in breadth.

It is followed by an esophagus of about the same length. The

intestinal diverticula diverge very widely and. are usually much

dilated. As a rule, they do not reach the middle of the body

TREMATODE PARASITES. 59

The excretory vesicle is V-shaped, the main stem being very

short.

The testes are two elongated oval bodies, measuring about

025x-014 mm. That on the left lies on the same level as the

ventral sucker, the one on the right being somewhat behind. The

vesicula seminalis lies adjacent to the anterior border of the

ventral sucker, and from its anterior end issues the pars prostatica.

Text-fig. 11.

Gymnophallus affinis, sp.n. Ventral view, x 400.

Ov. Ovary. T. Testes. V.S. Vesicula seminalis.

The ovary is considerably larger than the testes, measuring

-027 x:023 mm., and is more nearly globular. It is situated

almost immediately in front of the right testis and on the level

of the anterior border of the ventral sucker. The yolk-glands lie

over the anterior half of the ventral sucker. Each gland consists

of a somewhat crescentic mass of very irregular contour, the con-

cavities of the crescent being directed outwards.

The uterus lies mostly behind and to the left side of the

60: DR. H. LYSTER JAMESON AND DR. W. NICOLL ON

ventral sucker. It seems to form a single loop, starting from the

ovary, passing behind the ventral sucker and up along the left.

side of the body to some distance in front of the sucker, then

returning in the same way. The eggs are of conspicuously large

size, measuring °021—-028 x 015-018 mm.

There were a considerable number of much smaller specimens.

(13-19 mm. in length) bearing a close resemblance to the fore-

going. Even the smallest of them, measuring only °135 mm.

in length, was fully mature and contained about a dozen large

eggs. All intermediate sizes between this and ‘2 mm. were

observed. It is difficult to decide whether these small specimens

are the same as Gymnophallus affinis or whether they represent

a distinct species. The differences, apart from the difference

in size, are not very easy to detect. The body is altogether more

plump and the various organs more packed together. The

suckers are relatively larger. The intestinal diverticula extend

past the middle of the body and are in contact with the ovary

and the vesicula seminalis. The yolk-glands are slightly further

back and appear to be partly fused or at any rate very close

together, while the uterus extends forwards to the oral suckers

on the left side. ‘These differences, although noticeable, do not

appear sufficiently constant to warrant the creation of a further

new species, so that for the present we shall content ourselves

with noting the existence of these smaller forms.

GYMNOPHALLUS MACROPORUS, sp. n. (Text-fig. 12.)

This is the largest of the four species, and measures ‘4—"5 mm.

in length by *14--23 mm. in maximum breadth. The greatest

breadth is across the middle of the body, but is not very much

greater than that of the oral sucker, and this gives the animal

a very characteristic shape. The lips of the sucker are usually

everted and project well beyond the sides of the body. From

the anterior end the body tapers gradually towards the pointed

tail, though there is usually a slight inflation a little in front

of the ventral sucker. The oral sucker has a transverse diameter

of -14—-17 mm., the ventral sucker :065—075 mm. ‘The ratio

is generally about 9:4. The ventral sucker is situated at a

distance of about °3--4 mm. from the anterior end of the body,

i. e. about three-fourths of body-length.

Almost contiguous with the oral sucker is a comparatively

large pharynx measuring -040x°:035 mm. ‘The cesophagus is

slightly longer than the pharynx, measuring ‘05mm. ‘The

intestinal diverticula are usually moderately distended and form

an acute angle with each other. Their ends reach well past the

middle of the body. 1

The excretory vesicle is V-shaped and the limbs are frequently

enormously distended, compressing the intestine and giving the,

anterior part of the body a hollow appearance.

Owing to the close packing together of the eggs the ovary and

testes were extremely difficult to discern. They appear, however,

-'PREMATODE PARASITES, 61

to have much the same disposition with regard to the ventral

sucker as in Gymnophallus affinis, the ovary lying on the right

at the level of the anterior border of the sucker, the right testis

being at the posterior border and the left testis occupying an

intermediate level. The yolk-glands differ from those in

G. affinis in being much more compact, their outlines, in fact,

being quite oval. They usually lie over the anterior half of the

ventral sucker or a little in front of it.

Text-fig. 12.

Sa wos

eR RA os

eT 3

Gymnophallus macroporus, sp. n. Ventral view, X 250,

Ov. Ovary. T. Testes. V.S. Vesicula seminalis.

The vesicula seminalis lies immediately in front of the yolk-

glands, sometimes median, at other times displaced laterally.

From its anterior end issues a short wide pars prostatica, ruaning

down towards the ventral sucker. The uterus is disposed around

the ventral sucker and does not extend in front of the intestinal

62 ON TREMATODE PARASITES.

diverticula. The eggs do not show the same tendency to be

massed towards the left side as is seen in G. affinis. They are

of relatively enormous size, measuring ‘029-034 x -015—020 mm.,

the average being (032 x-0175 mm. There are usually about

30-100 eggs.

GYMNOPHALLUS OVOPLENUS, Sp. Nl.

This fourth species, which is undoubtedly distinct from all the

others, was characterised by its extremely minute size and by the

enormous overgrowth of the uterus, which completely filled the

body. The numerous eggs seen in the smallest specimen entirely

obscured the other organs, rendering a description of the internal

anatomy impossible. On that account, it is to some extent

a matter of conjecture as to whether the species actually belongs

to the genus Gymnophallus or not, but from its general appearance

there seems little doubt that it does.

It is a very plump, subcylindrical species with somewhat

pointed ends. The length is *11--13 mm. and the maximum

breadth about ‘07 mm. The oral sucker does not measure more

than 024mm. indiameter,and it isnot much larger than theventral

sucker (‘019 mm.), which is situated about two-thirds of the body-

length from the anterior end. The uterus fills the body, except

for a small space at the tip of the tail which usually remains

free. Anteriorly the eggs are packed tightly around the oral

sucker, rendering it almost invisible. They measure -016—019 x

011-012 mm.

It was at first thought that these specimens were simply young

forms of either G. affinis or G. edemie, but it seems impossible

that such can be thecase. Even when the animal is only -12 mm.

long the uterus attains a stage of hypertrophied development

which is never seen in either of the other species even when

twice as large. It might be conjectured that either or both of

these species may become prematurely ripe and that egg-pro-

duction falls off later. This, however, is a hypothesis which has

nothing to support it. Moreover, as has already been mentioned,

specimens measuring -13 mm. and closely resembling G. affinis

have been found, and in them the uterus occupied only a relatively

small proportion of the body.

Judging by the relative sizes of the suckers, this species is

more closely allied to G. wdemie than to G. affinis.

References.

(1) Herpman, W. A. 1903-6.—Report on the Pearl Fisheries of

the Gulf of Manaar.

(2) Jameson, H. L. 1902.—‘‘ On the Origin of Pearls.” P.Z.S.

1902, pp. 140-166.

(3) Lesour, M. V. 1906.—A Preliminary Report ona Trematode

Parasite in Cardiwm edule. Northumberland Sea Fisheries

Report for 1905 (1906).

PS 191s Pave

West,Newmean lith.

STRUCTURE OF BORNEAN DRAGONFLIES.

ON BORNEAN DRAGONFLIES. 63

(4) Nicort, W. 1906.—‘‘ Notes on Trematode Parasites of the

Cockle and Mussel.” Ann. Mag. Nat. Hist. (7) xvii.

pp. 148-155.

(5) Nicoti, W. 1907.—“‘ Observations on the Trematode Parasites

of British Birds.” bid. xx. pp. 245-271.

(6) Opuner, T. 1904.—‘ Die Trematoden des arktischen Gebietes.”

Fauna Arctica, iv. pp. 291-372.

5. Contributions to a Study of the Dragonfly Fauna of

Borneo.—Part I. The Corduliine: The Genus Amphi-

enemis: The Legion Protoneura. By F. F. Larpiaw,

MnAns Ee ZnS.

[ Received October 17, 1912: Read November 12, 1912. |

(Plate 1V.*)

INDEX.

Morphology : Page

Structure of terminal abdominal segments of Oe

EAC ICMOGEO! \ sharon anocedaccuebese cae sed dedonaece bon apmctondancg hee

Geographical Zoology :

Corduliinz of Borneo ....... doitepanenanrieatibar w alore|

Distribution of the genus Amphicnemis. Siete peace enter 70

Distribution of “certain species of the genus

DiS POROM CUR OA sah x dee siaiah satus Sashes Gata alee Aa NN!

Systematic :

The classification of the Corduliine ..................00.-... 63

Idionyx dohrni Kruger, subsp. borneensis move, Jee Ga

Amphicnemis louise, S| Oop Una ne ride santncr eco pecaearer ease doa tear LO

A. madelene, sp. un. ....... hue: oa Flt

List of the known species of the genus ‘Amphicnemis . Soache 74

The verticalis section of the genus Disparoneura ......... 75

I DEG DAREOCA LOA) Wh dleaen coeecnbon cen qoondarcheneoccacarecanccaceane: tLe

MDS WOSCUsASD AN steers Seas ee eae TCE RES EG NOTES 78

I RODS HCE DERSUGOUOR, SVs Wo scoconcovenaccccca coaenvoowsocecons {hss

ANISOPTERA.

CORDULIINE.

The most recent classification of this subfamily is that

suggested by Tillyardy. He proposes to arrange the various

genera composing it into four groups, as below :—

i. EKucordulina ;

ul. Idocordulina ;

ili. Macromina ;

iv. Synthemina.

Of these groups 1., iil., and iv. are on the whole well charac-

terised, the larvee are moderately well known, and the geographical

* For explanation of the Plate see p. 79.

+ Tillyard, Proc, Linn. Soc. N.S. W. 1912, xxxvi. 2, pp. 381-386.

64 MR. F. F. LAIDLAW ON

distribution of the genera fits in well with the proposed

arrangement. The second group, Idocordulina, however, is not

in so satisfactory a condition. The species referred to it are

mostly rare, largely tropical in distribution and hence their

larval forms are but little known, and the characters of

their venation are very diverse. In consequence there is reason

to suspect that the group will prove not to be a natural one,

although as no criterion is yet available for a better grouping of

the genera referred to it, needs must for the present that it be

retained in its existing form.

The group Synthemina, being purely an Australian one, does

not concern us here.

Of each of the other three Borneo has representatives.

Writing in 1899, Kriiger * was able to record only one

Corduline for Borneo, and but fourteen for the whole Malay

Archipelago.

Martin in his Monograph of the Corduliine, in the “ Collections

Zoologiques du Baron Edm. de Sélys Longchamps, Fase. xvi.”

published in 1906, gives a total for the island of seven.

In the present paper, thanks to the amount of material sent

to me from the Sarawak Museum by Mr. Moulton, the Curator,

Iam able to give the following list of species referable to this

subfamily :—

Eucordulina :

Hemicordulia assimilis Seélys.

Idocordulina :

Metaphya micans Laidlaw.

Idionyx dohrni borneensis, subsp. n.

Macromina :

Macromia cincta Ramb.

ts borneensis Kruger.

35 gerstaeckert Kruger.

5 cingulata Ramb.

as westwoodi Sélys.

5 sp. ?

Epophthalmia ausiralis Hagen.

i vittigera Ramb.

One may, with tolerable confidence, predict additions to this

list in the future.

Group 1. EUCORDULINA.

1. HemicorDULIA AssIMILis Sélys.

M. Martin has very kindly examined a male of this species for

me and determined its identification. It is new to the Bornean

* Kriiger, Stettin. Ent. Zeit. 1899, pp. 321-338.

BORNEAN DRAGONFLIES. 65

fauna, having hitherto been met with in the Celebes, New

Guinea, and the Solomon Islands.

The bulk of the genera referred to the Eucordulina group are

massed in the Holarctic and in the Australian regions ; with a few

species in Extra-tropical 8S. America, and outliers in the Oriental

region, the Seychelles, and Madagascar. On the whole the

distribution agrees fairly closely with that of the Conifer, and

suggests that the two groups must be of approximately equal

geological age.

Tillyard * has pointed out that the Australian genera are not

to be regarded as more primitive than those of the northern

hemisphere but show specialisation along lines of their own.

Somatochlora, perhaps the most primitive, is bi-polar.

| Vote.—In defining the EKucordulina, I believe that stress should

be laid on the convergence of M, and Cu, in the front wing as a

character especially marking the group. Accordingly I refer to the

group all the genera included by Williamsony in his group i.,

adding to them Cordulephya and perhaps Hesperocordulia, the

latter in deference to Tillyard’s views, but I would exclude from it

Oxygastra and the genera referred by Williamson to his groups

ii. and iii. |

Group 11. [bocorRDULINA.

As I have already remarked, I do not look on this group as

satisfactorily defined at present. Here I use it to hold those

genera which do not, in my opinion, fit into the Eucordulina on

the one hand nor yet into the Macromina on the other. These

are genera which fall into groups li., 1i., iv., of Williamson’s

classification. Whether such genera will not ultimately be

found to fall into one or more groups of equal value with the

HKucordulina and Macromina I cannot now conjecture.

2. Merapuya micans Laidlaw t. (PI. IV. figs. 1-3.)

12. Matang Rd. §, 3.10.10.

Length of hind wing 23 mm., of abdomen 20 mm.

The female is more brilliantly coloured than the male, and is

amongst the few Cordulines which have really brightly coloured

wings. It agrees closely in proportions and in details of vena-

tion with the male.

The membranule is large and uniformly grey. This is also

the case with the male. By an error I described it as having

the upper third of the membranule white, the lower part dark

brown. ‘This description should apply to the single cell forming

the anal triangle of the male.

* Tillyard, loc. cié.

+ Williamson, Ent. News, Nov. 1908, pp. 428-434, pl. xviii.

{ Laidlaw, Sarawak Mus. Journ. No. 2, 1912, pp. 65-67, pl. i.

§ All localities mentioned in this paper are in Sarawak, North-west Borneo.

Proc. Zoou. Soc.—1913, No. V. 5

66 _MR F, F. LAIDLAW ON

In the female both pairs of wings have a smoky tinge all over

them, most marked towards the apices. Further, the base of the

fore wings has a rich red-brown tint, best marked in the sub-

median space and in the sub-costal space, extending as far as the

level of the second antenodal. The base of the hind wing has,

too, a darker colour, except along the anal margin beyond the

level of the anal angle, but on this wing the colour is for

the most part of a dark brownish black (very similar to the

colour on the wings of a Rhyothemis) with a metallic glaze,

except that the median and cubital spaces have only the

transparent red-brown tint of the base of the fore wing. The

dark colour extends as far as the fourth antenodal nerve, and

posteriorly has a regularly curved margin. Body-colour similar

to that of the male. Upper surfaces of head and thorax metallic

blue-green, abdomen shiny black, under surface of thorax and

base of legs dark brown, the rest of the legs black.

The abdomen has been flattened and, especially at its distal

extremity, distorted. It does not show quite so distinct an

expansion of segments 7,8, 9 as does the male; none the less

there is a distinct enlargement.

The structure of these terminal segments is worth remark.

Segment 8 is about 3 the length of 9. Its lateral plates are pro-

duced ventrally and posteriorly into a pair of pointed spur-like

projections. The lateral plates of 9 have a similar arrangement

on a smaller scale.

The ventral plate of 8 is long, produced backwards to the

level of the end of 9 at least, its posterior margin running to a

median acute angle. It appears to fuse with the ventral plate

of 9, which is produced backwards beyond the level of the tip of

the abdomen and has exactly the shape of a spoon, with the

concavity lying upwards. The ventral plates of both 8 and 9

have a median longitudinal keel. Segment 10 is so much

crushed that it is impossible to discuss it. The appendages

are small, and reach about to the level of the end of the

spoon.

Neither in general organisation nor yet in the structure of the

genital appendages does Metaphya appear to show particular

kinship with Jdionyx. I figure (Pl. IV. fig. 4) for comparison

with the terminal segments of Metaphya, an outline drawing of

the same part of a female of Jdionyx dohrni Kriger from the

Peninsula of Malacca (Skeat Expedition). The anal appendages

of the male Jdionya distinctly approximate to the type found

in Macromia, whilst those of Jetaphya, which I also figure

here, are very different.

I have not been able to examine specimens of the Tropical

American Gomphomacromia paradoxa. In appearance, judging

from Martin’s figure *, there are grounds for considering relation-

ship between it and Metaphyw fairly close. From de Sélys’s

* Martin, op. cit. pl. ii. fig. 9.

BORNEAN DRAGONFLIES. 67

description of the anal appendages, in neither sex do these bear

any very marked resemblance to those of Wetaphya.

The genus /dionyx, hitherto unrecorded from Borneo, is

represented by a form of J. dohrni Kriiger, of Sumatra. Other

species in all probability await discovery.

3. IDIONYX DOHENI Kriiger, subsp. BORNEENSIS nov.

266. Matang, 1905-1907.

Length of hind wing............ 27°5 mm.

abdomen sens -caeaen: 7S) os

anal appendages ... 3 ,,

Fore wing: antenodals 13, postnodals 6, supratriangulars 1,

cross-nerves in median space 1.

Hind wing: antenodals 8, postnodals 9, one supratriangular

and two cross-nerves in basal space.

Head: under lip brownish-yellow, upper lip yellow with black

margin thickest in the middle line, rest of the front of head

black with green and violet metallic shades. Occiput black.

Prothorax biack above, dark brownish yellow at the sides.

Thorax metallic green above, with three yellow marks on

either side. The first is continued up from the coxe of the

second pair of legs, lying immediately in front of the humeral

suture, ending halfway up the suture. The second begins at

the coxee of the hindermost legs and runs up between the wings

asa narrow band The third is a rounded mark lying below the

base of the hinder wing.

The abdomen is thin and eylindrical, slightly widened at

segments 7—9, almost entirely black ; the four anterior segments

of a shiny texture, the rest dull. Under surface of 2 and 3 yellow.

Wings smoky, with faint yellow tinge at base; membranule

small, grey.

Legs: cox and base of femurs of first two pairs yellow-brown.

Lower third of first pair of tibias, and nearly the whole of second

and third pairs, red-brown. ‘The rest very dark brown or black.

Anal appendages a little longer than the last two segments of

the abdomen. Upper pair black, lower appendage very dark

brown, black at the tip. Seen in profile the upper pair are

cylindrical, slightly bowed upwards at their middle, tapering very

gradually to the extremity, which ends ina downwardly directed

point. The lower appendage slightly overlaps them, and is curved

upwards towards its extremity, which carries a minute backwardly

directed point.

Seen from above, the upper pair are thick for the first two-fifths

of their length, then rather thinner, approximated a little at

their middle, then diverging slightly, lastly turning inward again

towards their extr emities, Swhich have a rounded outline and

carry a fine tuft of hairs on their outer sides.

The lower appendage for the first two-fifths of its lent is

68 _ MR. F. F. LAIDLAW ON

rather ovoid in shape; at its middle it carries on either side

a blunt tooth or spur; the last two-fifths of its course it has

nearly parallel sides fringed regularly with fine hairs and its apex

blunt in outline.

The 10th abdominal segment carries on its dorsal surface a

small, laterally compressed truncate projection.

On the underside of segment 7, at the junction of its middle

and distal thirds, is a fine bunch of yellow-brown hairs pro-

jecting downwards. This character is perhaps specific, but may

occur in the males of other species; I have not seen it noted,

and have not been able to examine other specimens *.

I. dohrni borneensis differs from the typical J. dohrni from

Sumatra chiefly as follows :—

I, dohrni Kriger. I. dohrni borneensis.

i, All the cox yellow. First two pairs of coxe yellow.

ii. Lower anal appendage not Lower anal appendage slightly

overlapping upper pair. overlapping upper pair.

ili. Yellow markings on ab-

domen more extensive.

Group ii. Macromina.

Genus MAcRoMIA.

According to Martin, the following species have been recorded

from Borneo :—

Macromia cincta Ramb.

ak borneensis Kruger.

" gerstaechert Kruger.

is cingulata Ramb.

rr westwoodi Sélys.

I give a brief description of a very large female specimen,

collected by Mr. Moulton, which must probably be referred to an

-unnamed species.

4, MACROMIA sp.

A single 9 taken in October 1911 at Sadong.

Length of hind WTI teeter . 58°5 mm.

te pterostigma ...... 3 ss

The specimen is unfortunately badly damaged. It is remarkable

on account of its great size, and is most probably unnamed. I

prefer to leave it so for the present, in the hope that more

material may be forthcoming shortly.

Fore wing: antenodals 17-19, postnodals 10, supratriangulars

4, median cross-nerves 9.

Hind wing: antenodals 11, postnodals 12, supratriangulars 2,

median cross-nerves 4.

* See note at the end of this paper.

BORNEAN DRAGONFLIES. 69

- Wings of a smoky tint, without any basal colour.

Front of head entirely russet-brown, with metallic-green reflex

above ;.occiput black. Prothorax dark brown. Thorax brown

with a faint metallic-green reflex, antealar sinus bright brown.

‘The lateral stripe is of a pale brown colour, and is bordered on

either side with more richly metallic colouring.

Abdomen entirely bronze-black, except for a square yellow

mark on the base of segment 7 occupying not quite a third of the

length of the segment. The first four segments have a strong

metallic lustre, the rest are duller.

Legs black, slender.

In the present specimen the span of the fully extended fore

wings is 125 mm.

5. Macromia crncta Ramb.

1S. Baram.

Length of abdomen 49 mm., of hind wing 45 mm.

This specimen agrees exactly with de Sélys’s account of

M. cincta, save that segments 6 and 7 of the abdomen are

entirely black.

[Mote—The large female Macromia described above has

certainly a very close resemblance to J/. westwood, especially in

the colouring and in the absence of a basal mark to the wings.

De Sélys described M/. westwoodi 2 as having ‘“‘deux large bandes

antéhumeérales fauves,” a description which would scarcely apply

to the specimen I have described. It is further very considerably

larger, but the range of size in species of this genus is not well

known so far as Oriental species go, so that whilst I incline to

believe that Mr. Moulton’s specimen represents an undescribed

species, I feel bound to await further material before describing

it.

by gerstaeckeri is readily distinguished from other Bornean

species by its relatively small size (span about 70 mm. in the

male, 80 mm. in the female), by the possession of a narrow yellow

antemedial line, incomplete above, and by the position of the

external tooth on the upper anal appendages of the male, nearly

at the extremities of the appendages instead of being at their

middles, as in the other Bornean species so far as is known.

Further, it has a yellow band across the nasus. I confess that

the position of JJ. borneensis Kriiger seems to me a little

doubtful, it certainly comes very near J. cincta. It is 1m-

possible, however, to study these species satisfactorily without

the advantages of having a good series before one. J/. cingulata

Ramb., with much yellow on the face and abdomen, is very

distinct.

Genus HpopHTHALMIA.

6. EroPHTHALMIA AUSTRALIS Hagen.

IT have examined a young male of this species, collected by

70 MR. F. F. LAIDLAW ON

Mr. Moulton. The species has been recently discussed by

Dr. Ris.

7. EPOPHTHALMIA VITTIGERA Ramb.

ZYGOPTERA.

AGRIONIN&.

Genus AMPHICNEMIS Sélys.

Certainly one of the most characteristic genera of the Malay

province. It has been recorded only from Borneo, Sumatra, and

the Philippine Islands. It will ultimately, no doubt, be found

to occur in the Malay Peninsula. In its recorded area it is

probably represented by very many species.

The genus is notable for several reasons. One of these is

the remarkable sculpturing of the hinder lobe of the prothorax

found in several species, either in both sexes or in the males

alone.

A second, perhaps more remarkable character, is, that whereas

males of most of the species at any rate present a very uniform

system of colouring of the body, the females, on the other hand,

are often more brilliantly coloured than the males, and show, so

far as I can judge,a far greater diversity between the species in

this respect.

The extreme delicacy of these creatures, and the somewhat

bizarre form of the anal appendages of the males, together with

the curious prothoracic armature (closely paralleled in the case of

Disparoneura and some other genera), suggest that the genus is

highly specialised and “‘ gerontic” : to be compared, perhaps, with

Opisthostoma amongst the land molluscs, and Calamaria amongst

the snakes of the same province. No observations are available

on the habits or life-history of the species.

The material I have studied consists of six male specimens

belonging to five species, and of five females belonging to four

species ; representing in all, probably, six distinct species.

1. AMPHICNEMIS WALLACEI Sélys*. (PI. IV. fig. 7.)

1g. Baram, 15.10.10 (adult).

Length of abdomen...... 32°5 mm. (without appendages). —

a hind wing ... 185 ,,

This specimen agrees closely with de Sélys’s description of the

type, whilst the anal appendages bear an exact resemblance to

those figured by Dr. Ris for a specimen from Sintang.

[The female is said by Dr. Rist to have the whole thorax, the

femora, and tibize blood-red, the tarsi yellow, and the spines of the

legs dark. The prothorax is without the median spine which

occurs in the male sex. |

* De Sélys, Synops. des Platyenem is, no. 2, Bull. Acad. Belg. 1863,

+ Ris, Aun. Soc. Ent. Belg. lv. 1911, pp. 236-287, figs. 4 & 6.

BORNEAN DRAGONFLIES. VOI

2. AMPHICNEMIS LOUIS&, sp.n. (PI. IV. figs. 5, 5a.)

3. Baram, Sarawak, 1910 (adult).

12. Limbang River, 3.4.10 (adult).

3. Very similar in appearance to A. wallacei.

Lower lip dark brown, upper lip dark bronze-green ; the rest of

the dorsal surface of the head black, with metallic reflex.

Prothorax : aboveand at the sides bronze-black ; below whitish.

A small spine rises from the middle of the posterior margin, it is

hooked vertically upwards, and is about one-half the size of

that found in A. wallacet, Seen from above, the lateral angles

of the posterior margin are produced as a very small pair of

outwardly directed spurs.

Thorax: above dark metallic green, below brownish white.

Abdomen: segments 1 and 2 metallic green above, brownish

white below. The succeeding segments are of a dull brown

colour, becoming progressively darker, their under surfaces paler.

Pterostigmata black, with a very fine grey margin, which is

much narrower than in 4. wallacei. 13 antenodals in the fore wing.

Legs white (first two pairs lost), with articulation between

femur and tibia black, a fine black line running along the whole

posterior surface of the femur. ‘Tarsus missing.

Anal appendages whitish, lower pair about four-fifths the length

of upper pair. The upper pair are curved a little downward, their

extremities flattened laterally and folded in on themselves; Each

has at its middle a rounded projection directed inwards. Lower

pair much as in A. wallacei, but ending in an upturned point.

@. Agrees with the male in the characters of the prothoracic

posterior margin. The whole of the prothorax and thorax.is of

a rich orange-yellow colour, the alar sinuses have dark metallic-

green spots, and the prothoracic spine is tipped with the same

colour. The legs are blood-red, with black articulations and

black spines, the tarsi yellow. Abdomen similar in colour to

that of the male, but duller.

The vivid colouring of the female gives it a very striking

appearance.

3. AMPHICNEMIS MADELEN#, sp. n. (PI. IV. figs. 6, 6 a.)

Dono wuchine st 09%

¢ adult. Length of abdomen ......... 32 mm.

As hind wing ~..-... 18) 5

Lower lip yellowish white, upper lip pale yellow, with its base

brown, and a median and two lateral minute black spots.

Epistome black; upper surface of head dark green ; basal joints

of antennz pale brown.

Prothorax : wpper surface and sides metallic green, under surface

pale yellow with slight orange tinge. Posterior margin with a

long, median cylindrical horn, nearly vertical, dark at its base,

pale towards the extremity, proportionately much longer than in

72 MR. F, F. LAIDLAW ON

A. wallacei. Seen from above the lateral angles of the posterior

margin are acute, but scarcely produced to form spurs.

Thorax: above metallic green, on the sides a pale pearly green,

yellowish white below.

Abdomen with segments 1 and 2 metallic green above,

yellowish white below ; the rest brownish black above, paler below,

progressively darker backwards; 9 and 10 uniform very dark

brown, almost black. Wings with the pterostigmata of the hinder

pair bright orange, darker in the centre. Those of front wings

dark grey with paler margin. Legs entirely yellowish brown,

darker at the articulation, spines dark brown.

Anal appendages white in the younger male, tipped with purple-

brown in the more mature specimen.

Upper pair slender, cylindrical, slightly bowed downwards,

dilated at the extremities, the dilated part being folded over on

itself so that there is a ventral groove or channel. There is alxo

a small blunt internal projection at about the middle of the length

of each. Lower pair rather stout, antler-like, a little compressed

laterally, each with a strong tine directed inwards and upwards

at its middle.

Female unknown.

4, AMPHICNEMIS REMIGER Laidlaw *.

1 ¢. Batu Lawi.

[I have described this species elsewhere. Here I give its cha-

racters very briefly. No prothoracic spine. A small lateral spine

to posterior prothoraciec margin. Pterostigmata of all four wings

grey. Legs primrose-yellow with a black ring at each articulation.

Anal appendages white, upper pair slender with a small dorsal

tooth at their middles, extremities flattened to form an oval

paddle-shaped expansion. ‘Lower pair shorter, slender, ending in

a fine upturned point. Female unknown. |

5, AMPHICNEMIS MARTINI Ris 7

1 $6. limbang. 1 9. Matang Rd., near Kuching.

The male has been compared with the type by Dr. Ris, who has

kindly informed me that he can find nothing to distinguish it

therefrom, save that whilst im the type the pterostigmata of the

hinder wings are of a yellowish white, in Mr. Moulton’s specimen

they are bright orange.

The posterior prothoracic margin is without projections. The

legs are pale orange-yellow with yellow spines and black tibio-

femoral articulations. Theanal appendages are figured by Dr. Ris.

They are missing, together with the last three abdominal segments,

in this specimen.

The single female included here has also been examined by

* Journ. Str. Br. Roy. Asiat. Soc. 1912 (paper not yet published).

+ Ris, loc. cit. pp. 2387-288, tig. 6.

BORNEAN DRAGONFLIES. 73

Dr. Ris, who believes it to be rightly referred to the present

species. He observes that the shape of the posterior prothoracic

Margin is similar to that of A. wallacet 2 , but not identical with

it. It is gently convex, with marked lateral angles. The colouring

is very different from that of the female A. wallacet, and approxi-

mates to the colouring of the male. The dorsal surface of the pro-

thorax and thorax is bronze-green, their sides a pale pearly green ;

under surfaces and legs whitish yellow. There is a complete longi-

tudinal black line on the posterior surfaces of the femurs, black

spines, and black articulations. ‘The abdomen has segments | and

2 bronze-green above, dull yellow below; the remaining segments

are of a dull brown colour, paler below, progressively darker from

before backwards.

Length of abdomen 32 mm., of hind wing 20 mm.

The head is too much crushed and shrivelled to permit of any

description.

6. AMPHICNEMIS sp.

2929. Baram, 14.10.10.

Length of abdomen 33°5 mm., of hind wing 20 mm.

Upper surface of head entirely dark metallic green. Prothorax

red-orange, rather paler below ; its posterior dorsal margin gently

convex, produced on either side into a fine backwardly directed

short spur. Thorax with a rather narrow bright metallic-green

band, succeeded laterally by blood-red colouring, which fades into

a dull orange-red on the under surface; alar sinuses metallic

green.

Abdomen: segments | and 2 lustrous brown above, each with a

terminal metallic-green ring. The rest of the abdomen brown

above, pale whitish brown below, darker posteriorly.

Pterostigmata grey-brown with pale margin.

Legs red, tarsi whitish yellow, articular markings black, spines

dark brown.

The colouring of these specimens resembles very closely that of

Teinobasis rajah recently described by me. ‘There can be no

doubt that these specimens should be referred to Amphicnemvis

and not to ZVeinobasis. They have been examined by Dr. Ris,

who has favoured me with the subjoined remarks on them :—

‘“*T am ata loss to give good characters for separating Teinobasis

and Amphienemis—this although I believe that the two are quite

distinct genera, as proven by the widely different type of 3

appendages. There is certainly a difference in stature also,

Amphicnemis being decidedly the more delicate, with especially a

very narrow and feebly built thorax. But a good character that

would do for both sexes and for all the species is still to be sought

for, my material is so very insufficient for such an investigation.

I believe the great similarity in colour of the two forms in

question (Amphicnemis sp. Q and 7’. rajah) is merely a case of

convergence.’

74 MR. F. F, LAIDLAW ON

The following brief characterisation of the known species of

Amphicnemis may be useful :—

A median posterior prothoracic spine present in both sexes.

Spine similar in both sexes; upper surface of thorax of female without

metallic colouring. A. louise, sp.n. Borneo.

Spine in female Jonger than in the male; thorax above with golden

. bronze marking. A. gracilis * Kriger. Sumatra.

Prothoracic spine in male only.

Large species (abdomen g 45 mm.); thorax of female with metallic

band above, and with marked lateral projections to posterior margin

of prothorax. A, lestoides Brauer}. Mindanao.

Smaller: species (abdomen ¢ 36 mm.); thorax of female entirely

blood-red. A, wallacei Sélys. Borneo.

No prothoracic spine in males.

Pterostigmata of hind wings orange-yellow in male ;. colouring of

female similar to thatof male, also without prothoracic spine; small

species (abdomen 9 32 mm.). A. martini Ris. Borneo.

Pterostigmata of hind wings of male grey or black; small species

(¢ abdomen 34 mm.), 2 unknown. A. remiger Laidlaw. Borneo.

Pterostigmata of hind wings of male grey ; large species (¢ abdomen

41 mm.) ; female coloured very much like the male. No prothoracic

projections. A, ecornuta Sélys{. Sumatra.

Lastly, 4. furcata Brauer § from Luzon has no median spine

but a lateral pair in the male, whilst the pterostigmata are all

black ; A. madelene, sp. n., from Borneo, has the pterostigmata of

the hind wings orange-yellow, and a long cylindrical median

prothoracie spine in the male; in both species the female is

unknown.

Legion PROTONEURA.

Genus DispAroneuRA Sélys, Ris emend. ||

Lower lip with short rounded lobes; pterostigma rhomboidal

or lozenge-shaped ; lower section of triangle present as a vestige

or absent. No supplementary basal postcostal nerve. Basal

postcostal lying between the level of the antenodal nerves.

The genus so defined ranges from the Cape of Good Hope

through Tropical Africa to India, Ceylon, Burmah, and the Malay

Peninsula and Great Malay Islands. It appears to show advancing

specialisation from west to east both as regards colour and venation

characters, reaching its maximum in Borneo, east of which island

its occurrence is doubtful. [Two species described by de Sélys

as belonging to the genus Allonewra, from the Philippine Islands,

were subsequently referred to Disparoneura by him, but as in his

description no special reference is made to the position of the

basal postcostal nerve, which is the character used here to

distinguish the two genera, and as I have not been able to

examine examples of these two species, their exact position I

* Kriiger, Stett. Ent. Zeit. 1898, pp. 121-123.

+ Brauer, Verh. zool.-botan. Gesell. Wien, 1868.

< De Sélys, Ann. del Mus. Civ. di Genova (2) vii. 1889. Krtiger, luc. cié.

§ Brauer, loc. cit. ;

\| See note at the end of this paper.

BORNEAN DRAGONFLIES. 5)

treat as doubtful. They are Disparoneura? integra Sélys, and

Disparoneura ® obsoleta Sélys. |

Further east the genus is replaced by the closely allied Caco-

neura (Allonewra), which in regard to venation is still further

specialised.

Like many other genera which are richly developed in the

Malayan region, it appears to be but poorly represented in Java.

The Malayan species appear to fall into three groups characterised

by the coloration of the males.

One of these groups, possibly the most primitive, has the males

black with yellow markings on the head and thorax. A second

is that in which the males show blue markings on a black

ground.

In the third group the males are black, with carmine, brick-

red, and orange-coloured markings.

The females of all these groups appear to be very similar as

regards colouring; they are black, with dull yellow or orange

markings. They are, however, remarkable in the possession of

various curious developments of the margin of the prothorax.

The grouping of the species according to the colouring of the

male is suggested by Forster*, who regards certain forms with

black and red males as races or subspecies of D. verticalis. I

have here extended the limits of his “ verticalis section ” to include

all the black and red Malayan species with no lower sector of the

quadrilateral, or with only a trace of it. I think it unnecessary

to treat these species as mere subspecies of verticals; 1 believe

them to be well characterised and readily recognisable species.

I propose to group them as follows :—

A. Head of the male with a red band passing from eye to eye, across the

ocelli.

1. g. Anterior surface of thorax orange-red, upper lip black, anal

appendages red-brown.

9. Head black with orange band passing from eye to eye; prothorax

black with lateral orange lines, the posterior margin deeply

lobed (échancré) at its middle. Thorax black, with three

yellow bands on either side. D. dorsalis Sélys. Borneo.

2. g. Upper lip red; broad red bands on front of thorax.

2. Head black with yellow band from eye to eye; prothorax black

with livid-red lateral lines, its posterior margin carries on

either side a strongly curved point. Thorax black, with three

yellow or livid-brown lines on either side.

D. verticalis Sélys. Borneo.

3. g. Small red antehumeral lines on the anterior surface of the

thorax ; prothorax with a large red spot on either side.

Q. Head with complete yellow band from eye to eye; prothorax

with large red spot on either side. Posterior margin? Small

red humeral band on prothorax, and two yellow lateral ones

on either side. D. delia Karsch. Sumatra.

. Jaya (Forster).

(None of the males of D. dorsalis that I have examined has any trace of

a lower sector of the quadrilateral. In D. verticalis this is at least usually

present.)

% Worster, Fascic. Malay., Zool. pt. iv. Odonata, pt. u. p. 14.

76 MR. F. F. LAIDLAW ON

B. Upper surface of the head of the male entirely black.

[1. gd. Upper lip pale yellow, prothorax entirely black ; no antehumeral

band on thorax, two yellow lateral stripes on either side.

Segments 38-7 of the abdomen carmine-red above, anal ap-

pendages yellow above.

¢- Unknown. D. hyperythra Sélys. Borneo.]

2. &. Gene of a bright yellow colour, head otherwise entirely black.

Prothorax with lateral carmine spots. Thorax with a fairly

broad antehumeral carmine band, a brick-red succeeded by an

orange line on either side of the thorax. Abdomen with carmine-

red on segments 1 and 2 above.

- Unknown. :

?. Unknow D. hosei, sp.n. Borneo.

3. g. Head entirely black, prothorax with small lateral carmine spots.

A fine carmine antehumeral stripe on thorax, with lateral

brick-red and orange lines; abdomen entirely black.

Q. Head with yellow band from eye to eye. Prothorax with a

small pair of lateral bosses on the median lobe, the posterior

margin produced backwards into a flat rounded median collar.

D. humeralis Sélys. Malacca ;

3a. g. Antehumeral line absent, prothorax all black. Java.

D. humeralis, var. nigra Forster.

Pahang.

4. &. Head entirely black, prothorax entirely carmine above. Abdomen

with segments 1, 2, 3 carmine above.

(2?) 2. Head with very narrow incomplete line passing from eye to eye ;

lateral ends of posterior margin of prothorax produced to form

two large forwardly directed horns (cf. D. verticalis).

D. peramend, sp. Nn. Borneo.

The presence of a lower sector of the quadrilateral appears

to be fairly constant in D. humeralis, and perhaps generally absent

in the other species.

Of the group which includes males with blue and black

markings, Borneo possesses the following species :—

D. interrupta Sélys,

D. collaris Sélys,

D. lansbergi Sélys (?) ;

whilst D. moultont Laidlaw and D. gracillima Sélys perhaps

represent a third group, with yellow markings on a black ground.

D. moultoni shows, I believe, some relationship to D. hyperythra,

and may be a melanotic species allied to it, and not primitive.

Lastly, D. awrantiaca Sélysis a handsome Bornean species with

the anterior surface of the thorax bright orange, and plentiful

orange markings on the abdomen in the male, which is very

distinct in its venation and eolouring from other members of the

genus found in the island. The posterior margin of the pro-

thorax of the female carries subtriangular, strongly recurved,

plate-like extensions.

Description of new species.

DIsPARONEURA PERAMGNA, sp.n. (PI. IV. figs. 8, 8a.)

23 6,2 29. Lawas and Limbang, Aug., Sept., 1909.

Length of abdomen: ¢ 29°5 mm.

Sy hind wing: ¢ 16 mm.

BORNEAN DRAGONFLIES. 77

No trace of lower sector of quadrilateral. Upper sector of

quadrilateral of fore wing extending to the first cross-nerve after

the quadrilateral in all the specimens. In the hind wing variable,

ranging from the nodal cross-vein to that immediately beyond it.

Postnodals of fore wing 14-15.

3. Head entirely black, including the lower surface.

Prothorax: above of a rich carmine, anterior and posterior

lobes delimited by a fine transverse black line. Sides and under

surface black.

Thorax black ; in front a pair of carmine bands, slightly cres-

centic, extending for about two-thirds of its length from its

anterior margin, succeeded at the top of the thorax by a minute

‘carmine spot. Antealar sinuses of the same colour. Anterior

lateral thoracic band of a brick-red colour, the posterior, on the

metepimeron, yellow.

Legs black, coxee and a ring round the base of the femora brown.

Abdomen black, segments | and 2 carmine above; 3 has at its

base above a very long narrow carmine triangle with its apex

directed backwards, extending for about one-fourth the length of

the segment.

Anal appendages black, dark brown at their bases.

The upper pair seen from the side have their posterior margin

rather crescentic, with an inwardly directed spur on the lower

side. Lower pair with the posterior margin @-shaped, the

terminal hook rather more slender than in most species of the

genus.

@. Head black, upper lip yellow; a very fine yellow mark runs

inward from each eye to the ocelli, but does not form a complete

band.

Prothorax black; median lobe with a pair of round lateral

yellow spots, anterior lobe with a very small lateral yellow mark.

The median lobe is almost hemispherical. The posterior margin

is produced on either side so as to form a curious horn-like

projection directed upwards and then forwards.

The thorax is black. There is a fine yellow antehumeral stripe

incomplete above, extending about one-third the length of the

upper surface. There is a well-marked yellow lateral band, and

behind this the yellowish grey of the under surface shows at the

side of the metepimeron.

Legs brownish black, the anterior surfaces of the femora and

tibias yellow.

Abdomen dull brown, paler beneath, with a small subterminal

pale ring on segments 3-6; the last three segments almost

entirely black.

The male is characterised especially by the colour of the head

and prothorax; the female appears to approach that of D. verticalis,

but differs in details of coloration. I believe I have correctly

‘referred the females described above to this species, but as was

the case with that described by de Sélys for D. verticalis, the

evidence is not conclusive.

78 MR. F. F, LAIDLAW ON

DIsPARONEURA HOSEI, Sp. N.

1 ¢. Baram, Borneo.

Length of abdomen ............... 32 mm.

a lawine) WAN?” boneoneceses Teh

Fourteen postnodal costals on the fore wing.

Head entirely black above and below, save for a fine yellow

mark on the gene. Prothorax black above, with a small carmine

spot on each side of the three lobes on either side; yellow

markings underneath about the articulations of the limbs.

Thorax black, with a rather broad complete pair of antehumeral

bands of a rich carmine colour, a median lateral pair with an

orange tone, and a sharply defined posterior pair yellow in colour.

Ventrally the thorax is black, save for yellow points at the

articulation of the limbs, as in the prothorax.

Abdomen black; segment 1 has a small transverse, terminal,

carmine band dorsally, 2 is entirely carmine above, 3 has a fine

carmine line above running nearly the whole length of the seg-

ment, tapering to a very fine point posteriorly.

Legs black, the femurs have a yellow ring at their bases, best

marked on the posterior pair.

The abdominal appendages are black, the upper pair seen in

profile are about equal to the lower pair; these latter in profile

appear blunt, but seen from above they end each in an incurved

hook, as do those of allied species.

This species is very near D. humeralis from Malacca. It is

characterised by the yellow gene, the broad carmineantehumeral

bands, and the red markings on the first three abdominal

segments.

The single specimen was received some years ago from Mr. Hose

along with other material from Baram.

Genus Prorosticta Sélys.

PROTOSsTICTA VERSICOLOR, sp. 1.

1 2. Lawas, 15.9.09 (27-6).

Length of abdomen 27 mm., of hind wing 19 mm.

Fifteen antenodal nerves in fore wing. Basal postcostal nerve

lies at a level widely proximal to that of the first antenodal costal

nerve. A second postcostal nerve, probably representing a trace

of the lower sector of the quadrilateral, lies at a level just proxi-

mal to the second antenodal costal. Pterostigma trapezoid,

covering more than one cell. Upper sector of triangle reaching

some distance beyond the nodus in fore wings.

Head bronze-black, upper lip greyish white, lower lip brown.

Prothorax and thorax chocolate-coloured above and on the sides,

with a metallic lustre. Under surfaces pitch-black.

The prothorax is provided with a remarkable spine resembling

very closely that found in certain species of the genus Amphi-

enemis. It rises from the middle of the posterior margin and is

BORNEAN DRAGONFLIES, 79

about 1 mm. in length. It projects upwards with a slight

forward curving.

The abdomen is very slender. Segment 9 longer than 8.

Segments 1-8 of a dull brown colour growing darker from before

backwards, each of the segments 2-8 with a pale narrow ring at

either end, 9-10 of a chocolate bronze-colour. ‘The legs are dull

yellow, with dark articulations and cilia.

I cannot at the moment recall any other Malayan Agrionid

which has the under surface of the thorax actually darker than

the upper side. This peculiar colouring of the thorax, together

with the pale yellow legs, gives this insect a very striking appear-

ance. It is further distinguished from other species of the genus,

which is new to Borneo, by the prothoracic spine.

Genus Puatysticra Sélys.

Puavysricra RuFostTIGMA Sélys. (Pl. IV. fig. 9.)

1 $. Lawas, August 1909. (No. 34.)

Length of abdomen ............ 34 mm.

ss onuaNG! WTO Goncoenoe vag No

The specimen agrees in every particular, excepting size, with

the type from Labuan. The latter is distinctly smaller, whilst

Mr. Moulton’s specimen approaches P. quadrata in size. The

anal appendages further resemble those of the type of P. rufosti PLE

and differ completely from that figured by me for P. guadrata*

Iam thus able to record 13 species referable to the legion

‘ Protoneura,’ arranged in three genera, for the island of Borneo.

EXPLANATION OF PLATE IV.

Fig. 1. Ventral view of terminal segments of abdomen of Metaphya micans

Laidlaw. @.

2. Lateral view of second abdominal segment of Metaphya micans Laidlaw.

8. Lateral view of anal appendages of the same.

4, Terminal segments ot Idionya dohrni Kriger 92, for comparison with

fig. 1.

+5. Profile view of prothorax of Amphicnemis louise, sp.n. 6.

ba. “4 » anal appendages of the same.

6. » prothorax of A. madeleneé, sp.n. 6.

6a. Lateral view of anal appendages of the same.

7. Profile view of prothorax of A. wallacei Sélys.

8. Disparoneura peramena, sp.n. § (anterior

end to the right). :

8a. Similar view of male anal appendages of the same species.

9. Lateral view of anal appendages of Platysticta rufostigma Sélys. ¢.

Norr.—Since the above notes were written I have received

from Dr. Ris copies of two accounts recently published by him

dealing with Dragonflies from the Orient. The first of these

(Supplementa Entomologica, Deutsch, Ent. Mus. No. 1, Aug. 1912,

* Laidlaw & Forster, in Fascic. Malay., Zool. pt. iv. Odonata, pt. ii. p. 9, fig. 1.

+ The profile figures are drawn on one plane, and, in consequence, the lateral

angles are not shown.

80 MR. E. S. GOODRICH ON THE

pp. 44-84, Taf. ili-v.), discusses new Dragonflies from Formosa,

South China, Tonkin and the Philippine Islands. Amongst others,

two species of the genus Jdionyx are described as new. In the

case of one of these, 7. claudia Ris, from Tsa-Yin-San, the male

carries on the under side of segment 7 of the abdomen a brush of

hairs exactly similar to that which I have described as occurring

in J. dohrni borneensis. Dr. Ris figures this feature in his paper

(loc. cit. p. 83, fig. 18). The second paper (Tijdschrift voor

Entomologie, Deel lv. 1912, pp. 158-182, pls. 6, 7, 8) contains an

account of Odonata from Java and Krakatau. The characters of

the genus Disparoneura and Caconewra are discussed, and a

specimen of D. humeratis from Mula (Java) is recorded.

6. On the Structure of Bone in Fishes: a Contribution

to Paleohistology. By Epwin 8. Goopricu, M.A.,

F.R.S., F.Z.8., Fellow of Merton College, Oxford.

[Received November 7, 1912: Read November 26, 1912.]

(Text-figures 13-16.)

INDEX. Page

WHOTROINOIE? “coo ueross sneacasaaeoe opanoucenmodesecounoscacseods ‘tl)

Phylogeny of Teleostei...............00cceceeeteeeeees renee 83

In a paper on the Scales of Fishes, published in the ‘ Pro-

ceedings’ of this Society five years ago (1), I showed that the

so-called ‘“Ganoid” scales are of two kinds, differing funda-

mentally in minute structure and mode of growth. Scales of the

first kind, to which the name Cosmoid was given, are typically

covered with an outer layer of cosmine, and grow by the addition

of new cosmine at the edge and new layers of bony tissue on the

inner surface. The second kind, the true Ganoid scale, grows by

the addition of new complete concentric layers, formed of cell-

less ganoine on the outer surface and bony tissue on the inner

surface. Cosmoid scales are found in the Dipnoi and Osteolepidoti

(extinct Crossopterygii), and in these only. True Ganoid scales

occur only in the Actinopterygii and Polypterini(which probably

belong to the Actinopterygii, 2). Moreover, it was further

shown that the Ganoid scales can also be distinguished into two

varieties—the Paleoniscoid and the Lepidosteoid. The former

is characterised by the presence of a middle cosmine-like layer,

and occw's only in the Chondrostei (Paleoniscide and their

allies) and in the Polypterini; while the latter variety—the

Lepidosteoid scale—is found in the Orders Amioidei (Protospondyli,

+ Pholidophoride, and Oligopleuridz) and Lepidosteoidei (Lepi-

dosteide and Aspidorhynchide). The lepidosteoid scale is easily

distinguished by the absence of the middle cosmine-like layer

and by the presence of a system of delicate tubules running

through and at right angles to the bony layers. The tubules have

been described by Reissner (5), Hertwig (3), and Nickerson (4)

STRUCTURE OF BONE IN FISHES. 81

im the seales of Lepidosteus, and by myself in those of various

extinct Amioids and Lepidosteoids (1, 2). They branch, as a

rule, only at their inner end, and pass outwards to the surface.

In the living tissue they are occupied by the long protoplasmic

processes of large cells on the surface of the scale, Probably,

these remarkable cells are merely modified bone-cells, which,

instead of becoming buried in the ostein matrix, remain outside

it while retaining their connection, by means of the long process, -

with the place they originally held. This interpretation is

illustrated in text-fig. 13.

Text-fig. Se

Diagram illustrating the structure and growth of lepidosteoid bone.

b.l., bony lamella; /.¢., lepidosteoid cell; o., osteoblast or bone-cell.

It follows that the Actinopterygii can be classified into two

large groups according to the structure of their scales: the first

is distinguished by the possession of paleeoniscoid scales, and

contains the Chondrostei (with which the Polypterini should

probably be placed, 2); the second group contains the Amioidei

(Protospondyli, Pholidophoride, and Oligopleuride) and the

Lepidosteoidei (AStheospondyli).

Now it might be expected that this striking difference in the

histological structure of the true ganoid scales would also be

found in the cranial plates and other dermal bones of these fishes,

and this is indeed the case. The dermal bones resemble the

scales not only in appearance, but also in microscopic structure.

Often the resemblance is so close that they cannot be

distinguished; but the dermal bones may, of course, lose the

covering of ganoine, as sometimes happens with the scales them-

selves in the more modified forms. Thus, whereas lepidosteoid

tubules are never found in any part of the skeleton of the

Polypterini or Chondrostei, they occur in the dermal bones of all

the recent and extinct Amioidei'and Lepidosteoidei I have been

able to examine, with a single possible exception ( Oligopleurus)

to be discussed later *.

* T am much indebted to Dr. A. Smith Woodward for the supply of most of the

material on which these researches were carried out, and to Miss R. Harrison for

the preparation of a large number of microscopic slides of the bone of various fishes.

Proc. Zoot. Soc.—1913, No. VI. 6

2 MR. E. S. GOODRICH ON THE

Not the dermal bones alone, however, show the lepidosteoid

structure, but the whole endoskeleton as well. The skull-bones,

the ribs, even the vertebral centra, are all provided with the

characteristic tubules traversing the bony lamelle, just as in the

Text-fig. 14.

Enlarged view of a section of the endoskeletal fin-ray of Lepidosteus osseus.

Lettering as in text-fig. 13.

Text-fig. 15.

b./.

Enlarged view of a section through the neural spine of Amia calva.

Lettering as in text-fig. 13.

scales (text-figs. 14-16). This remarkable and interesting fact

has not, so far as I am aware, hitherto been observed. It follows

that, from the examination of the minutest fragment of the

STRUCTURE OF BONE IN FISHES. 83

skeleton of a living or extinct species of fish, we can decide

whether or not it belongs to the Amioidei and Lepidosteoidei,

or to some other group. The histological structure of the bone

may therefore be of the greatest practical value for the identi-

fication of fragmentary specimens *. It also may prove of great

importance in the interpretation of phylogeny.

Text-fig. 16.

Enlarged view of a fragment of the vertebral centrum of Ophiopsis.

Lettering as in text-fig. 13.

We cannot as yet determine for certain which type of bone is

the more primitive; but I am strongly inclined to believe that

the lepidosteoid type is the more specialised form, some of the

osteoblasts having become modified to form tubules. This

conclusion is supported by the evidence of paleontology, since no

Amioid or Lepicosteoid is known below the Permian, while

Chondrosteans occur in the Devonian strata. In the absence of

decisive evidence we may suppose that the lepidosteoid structure

first appeared in the scales, then spread to the dermal bones, and,

finally, reached the deepest parts of the endoskeleton—this, of

course, is mere conjecture. At all events, since we find ordinary

bone in all the Osteolepidoti, Ccelacanthini, Polypterini, and

* The following is a list of the Actinopterygii examined :—Lepidosteoidei :

Lepidosteus, Aspidorhynchus. Amioidei: Eugnathide—Eugnathus, Caturus,

Heterolepidotus. Pachycormide—Pachycormus. Semionotidee—Lepidotus, Dape-

dius. Macrosemiide — Macrosemius, Ophiopsis. Pyenodontidz — Mesturus,

Gyrodus. Amiide—Amia. Pholidophoride—Pholidophorus. Oligopleuridax—

Spathiurus, Oligopleurus, Ginoscopus. Also Leptolepis, Thrissops, and a large

number of Teleosts.

84 ON THE STRUCTURE OF BONE IN FISHES.

Chondrostei, it would seem that the Amioidei and Lepidosteoidei

have been derived from a common ancestral form which developed

the lepidosteoid scale and bone, diverging in this and other

respects from the remainder of the Teleostomi.

It becomes now a matter of great interest to ascertain from

which group the Teleostei may have been derived. On general

anatomical grounds they would certainly be associated with the

Amioidei (2). But so far I have been unable to discover the

lepidosteoid structure either in the scale or in the skeleton of

any living or extinct species of Teleost, even after the examina-

tion of representatives of a very large number of families. In

the lower Teleostei (including the Leptolepide) the bone is of

the ordinary structure, similar to that of Osteolepis or Palconiscus ;

but, as is well known, in the higher forms it becomes generally

very much modified, chiefly owing to the loss of the bone-cells.

Only in the Fistulariidee does the structure of the bone recall

that of the Amioid. Even here, however, the resemblance is not

close, and I have not been able to convince myself that the fine

canals described by Stewart (6) are really homologous with

lepidosteoid tubules.

In connection with the phylogeny of the Teleostei it is

interesting to note that Oligopleurus vectensis A.S. W. has no

lepidosteoid tubules in its endoskeleton. Oligopleurus esocinus

Th. I have not had an opportunity of examining ; but Spathiwrus

and Gnoscepus, the only other genera belonging to the family,

have the typical lepidosteoid structure in the scales, dermal bones

and endoskeleton. Associated with a skeleton of Oligopleurus

vectensis in the British Museum is a plate, either a scale or a

dermal scale-like bone, of typical lepidosteoid structure; it

cannot, however, be made out for certain whether this bone

belongs to the skeleton or not. QO. vectensis certainly differs in

bone-structure from other Amioids, and probably should be

placed not with the Oligopleuride, but with the Teleostei *.

We may suppose that the common ancestor of all the Holostei

(Amioidei, Lepidosteoidei, and Teleostei) was some primitive

Actinopterygian with lepidosteoid scales, but in which the

lepidosteoid structure had not yet penetrated to the endoskeleton.

If this supposition is correct, ‘“ Oligopleurus” vectensis might

perhaps be a representative of such an unspecialised group.

Placed by some authors among the Amioids, and by A. Smith

Woodward among the primitive Teleosts (7), it certainly appears

to be a somewhat intermediate form.

As a provisional hypothesis, the view may be adopted that the

Amioids and Lepidosteoids on the one hand, and the Teleosts on

the other, diverged from a primitive group possessing lepidosteoid

scales and ordinary bone; and that in the former the lepidosteoid

structure spread inwards over the whole endoskeleton. The

extreme modification of the scales in the Teleost series would

* Mr. Regan informs me that O. vectensis is withont fulera, and resembles the

Leptolepid in the structure of the tail, and should be included in that family.

Py Aro, Weise

Percy Highley del.etlith C Hodges & Son imp.

MYZOSTOMA COSTATUM F.S.L.

a. te

ThA Se

wae

Poe, Se vie

Son imp.

Hodges &

lith

ey deLet

Percy Mighl

1. MYZOSTOMA RUBROFASCIATUM v.Graff

2-3. M.CROSSLANDI, Sp.n.

Dae. LOIS a eis

C.LB.&Hdel. :

Highley lith. C Hodges & Son imp.

MYZOSTOMA CROSSLANDI!, dSp.n.

If, Ze S. Ils, wl, Ale,

1. MYZOSTOMA CROSSLANDI. 2-4, M COSTATUM.

ON MYZOSTOMIDA FROM THE RED SEA. 85

sufficiently account for the loss of the tubules in these fishes.

Obviously much remains to be done before the full significance

of the structure of bone can be elucidated, and this paper has

been written with the object of drawing attention to the interest

and importance of this new line of research.

List of References.

1. Goopricu, E. 8.—‘ On the Scales of Fish, Living and Extinct,

and their Impovytance in Classification.” Proc. Zool. Soe.

1908.

2. Goopricu, E. $8.—‘‘ Vertebrata Craniata” first fascicle—

Cyclostomes and Fishes. Part 9 of ‘Treatise on Zoology,’

ed. by Sir Ray Lankester. London, 1909.

Herrwic, O.—‘* Ueber das Hautskelet der Fische.” Morph.

Jahrb. vol. v., 1879.

. Nickerson, W. 8.—‘‘ Development of the Scales of Lepid-

osteus.” Bull. Mus. Comp. Zool. vol. xxiv., 1893.

. RetssNer, E.—‘‘ Ueber d. Schuppen von Polypterus u. Lepid-

osteus.” Arch. f. Anat. u. Phys., 1859.

6. Srewarr, C.—‘“ Endoskeleton,” in Catal. Roy. Coll. of Surgeons,

2nd edit. vol. 1., 1900.

7. Woopwarp, A. 8.—Catalogue of the Fossil Fishes in the British

Museum, 4 vols., London, 1889-1901; and ‘ Outlines of

Vertebrate Paleontology, Cambridge, 1898.

By Se)

7. Report on the Myzostomida collected by Mr. Cyril

Crossland in the Red Sea in 1905. By CHaruxs L.

Boutenaer, M.A., D.Sc., F.Z.8., Lecturer on Zoology

in the University of Birmingham.

[Received November 12, 1912: Read November 26, 1912. |

(Plates V—VIIT. and Text-figures 17-23.)

INDEX.

Page

Anatomy of Myzostoma spp.. Be ate: 8S

'The “Suckers? of MM. costatum I. S. L.. Meee oer OO)

Ophiurid as the host of HW. costatwm ¥. Sha ae OS

Abnormalities in IW. costatum F.S.L. ............00..... 983

M. costatum F.S. UL. Jal gab Ramen chica ee co Rockne ESeeEn SMiEasIC

M. rubrofasciatum v. Graff the ements eine dake MOL

HUT CRROSSUGIIC Ng SUSUR) toabiaveshe ogodeccee nce ddbmosuemedenaasoemcaqocien ] MOM

List of References aeRO Ene Re NeA EASA oustcn paced oars! OOE

Explanation of the DIA GASIIC ie ee Dadian eee. TOR

The Myzostomida described in this report were collected by

Mr. Cyril Crossland on the Sudanese coast of the Red Sea im

86 DR. C. L. BOULENGER ON

1905 and were sent, together with the other Annelida, to Mr. F.

Potts, of Trinity Hall, Cambridge, who submitted them to me for

examination.

Mr, Crossland’s collection is of considerable interest, since very

little is known of the Red Sea Myzostomida, and the only two

species recorded from that locality were described from very scanty

material.

The material as I received it from Mr. Potts consisted of six

tubes containing about 170 specimens. Of these six belong to a

species hitherto unknown and which I take great pleasure in

dedicating to Mr. Crossland; two I have referred to v. Graft’s

Myzostoma rubrofasciatum, described from a single specimen in

1884; whilst the remainder belong to a species which I take to

be the old-established form J/yzostoma costatum, one of the first

three species of the group described by Leuckart so long ago as

1830.

The importance of a collection of this kind, however, is not to

be judged by the number of species represented in it. About

100 species of Myzostoma are on record and of these by far the

greater number have been described from external characters

only, ill-preserved or limited material having made a complete

examination impossible. Mr. Crossland’s specimens were in an

excellent state of preservation and I have been able to give a

fairly complete account of the anatomy of two of the forms men-

tioned above; moreover, the very large series of specimens of

A. costatum reveals a range of variation remarkable even for this

group of animals, and affords an opportunity of judging the

merits of various characters for systematic purposes.

In my descriptions of the species I have endeavoured to give as

complete an account as possible of the general morphology of the

various organs, especially of those which might be of taxonomic

interest. I have, however, refrained from attempting any histo-

logical decriptions, as I felt it useless to do so without material

specially preserved for such a purpose. Throughout this paper I

have retained as far as possible the nomenclature of parts used

by v. Graff (7-9) * and Nansen (15), and have not followed the

example set by some of the more recent workers on the

Myzostomida.

I desire to take this opportunity of thanking Mr. Potts for en-

trusting me with this valuable material; Prof. F. Jeffrey Bell

very kindly allowed me to examine some of the types of the

‘ Challenger’ collection of Myzostomida preserved in the British

Museum (Natural History), and I am also indebted to Mr. H. C.

Chadwick, of the Port Erin Biological Station, for information

concerning the Red Sea Echinoderms from which Mr. Crossland’s

specimens were obtained 7.

* The figures in brackets refer to the List of References on p. 107. ‘

+ I also wish to express my indebtedness to Mr. Cox, of the Zoological Depart-

ment of this University, for the excellent photographs reproduced on Pl. VIII.

MYZOSTOMIDA FROM THE RED SEA. 87

Myzostoma costatum F. 8. Leuckart. (Pl. V. & Pl. VIII. figs. 2-4.)

M. costatum Leuckart (10), p. 612; (11), p. 8, pls. 1.1.

5 v. Graff (7), p. 11, pl. i. figs. 13-14; (8), p. 32.

Myzostoma costatwm was one of the first three species of the

genus described by F.S. Leuckart in 1830 (10), his account being

based on specimens obtained in the Red Sea from Comatula multi-

radiata Lam.

A more complete description of these specimens was published

by the same author in 1842 (11), together with the following

brief diagnosis of the species :—

‘Char. specif.: Corpore depresso, ovali, margine crenulato,

dorso costato; acetabulis suctoriis et hamuliferis separatis, ace-

tabulis utrinque quatuor et hamulis in utroque latere quinque.

Hab. in mari rubro, Comatule multiradiate parasitus.”

The species was rediscovered by Semper, who obtained two

specimens from Bohol, in the Philippines, and these were de-

seribed by v. Graff in his well-known monograph published in

1877 (7). The two individuals, which measured 32 and 23 mm.

in length respectively, agreed closely with those described by

Leuckart. The dorsal surface presented the characteristic rib-

like elevations which gave the name to the species, and, more-

over, the animals possessed another feature noticed by Leuckart,

namely a small triangular incision at the anterior end of the

body, at the apex of which the mouth is situated. v. Graff’s

diagnosis of IM. costatwm was as follows :—“ Corpus ovale de-

pressum, incisura antica triangulari, griseo- aut nigro-brunneum,

supra costatum. Costa una longitudinalis mediana qua commu-

nicantes 10-14 cost transversales ad marginem product. Mar-

gine orientes inter has coste breviores secundarie et tertiare

illam longitudinalem non attingentes marginemque crenulantes.

Parapodia crassitudine uncinorumque forma J. glabrum, longi-

tudine V. cirriferum equantia. Acetabula elliptica. Os ad basin

incisure triangularis, ventrale, papilla cloacalis ventralis. Longit.

ad 32mm. In mari rubro Comatule multiradiate Lam., in mari

prope Bohol Actinometra solaris Lam. incola.” ;

A few years later v. Graff obtained another specimen, collected

by Haeckel at Tur, near Sinai, and described it in the * Challenger’

Expedition Report (8). This specimen was of a yellowish-brown

colour and measured nearly 2 mm. in length. The breadth was

rather greater than the length, and the anterior notch, as well as

the “ribs,” were not so distinct as in the specimens from Bohol ;

the marginal notches were, however, more marked.

As mentioned above, of the large number of Myzostomids col-

lected by Mr. Crossland more than 160 belong to a species which,

although differing somewhat from previous descriptions, I have

decided to refer to Leuckart’s M. costatum. The series of speci-

mens is a most interesting one, containing individuals of all sizes

between 0:5 and 4 mm. in length and presenting a wide range of

variation.

Mr. Crossland’s specimens were obtained from two localities :

88 DR. C. L. BOULENGER ON

(a) at a depth of 10 fathoms in Suez Bay on Antedon serripinna

Carpenter, (6) at Ul Shubuk (9 fathoms) from Antedon savignyt

Miller, and from Ophiurids * which lived with their arms twisted

round those of the Crinoid. This is not the first time an Ophiurid

has been described as the host of Myzostomids: H. L. Clark in

1902 (4) called attention to the occurrence of a speciest on

Asteroceras pergamena Lyman, and remarked that the position

of the worms upon the concave underside of the arms made it

clear that they had not become accidentally attached to the

Ophiurids since the latter were dredged~. With regard to

Mr. Crossland’s specimens, I think there can be little doubt but

that the actively moving Myzostomids had migrated from the arms

of the Antedon on to those of the commensal Ophiurids; it is

important to note that Antedon savignyi is most probably the

Comatula multiradiata Lam., from which Leuckart obtained his

type-specimens of Myzostoma costatum.

A typical adult individual (cf. Pl. V. figs. 1-5) has a flat body,

approximately oval in shape; the breadth is usually greatest

anteriorly, the posterior region narrowing slightly behind the

last pair of parapodia. There is no distinct transparent border,

the intestinal and uterine branches extending to the body-

margin.

On the dorsal surface a median longitudinal elevation is well

marked, running along the whole length of the animal; it is

broadest near the middle of its course. From this median eleva-

tion arise 6-8 (usually 7) pairs of primary coste which radiate

outwards and terminate on the body-margin. The second to fifth

pairs are frequently the most developed, the first and last two or

three being less distinct and often incomplete. Between the

primary coste are secondary and tertiary ones, as described by

v. Graff, their number and arrangement varying greatly. In some

individuals the coste are broken up so as to form rows of flat

tubercles (Pl. V. fig. 2). The body is thickest in the region of

the median longitudinal elevation and gradually thins down

towards the margins.

According to Mr. Crossland’s notes, the colour of the living

worms is ‘greenish white, with white dorsal ridge and two

longitudinal grey lines on either side of it.” The colour of the

preserved specimens varies considerably ; a few individuals kept

in a separate tube are of a greenish-grey colour, but the majority

are of a dark sepia-brown ; this coloration is probably artificial

and due to some dissolved pigment in the alcohol with which the

worms were preserved.

The margin of the body is indented so as to form a series of

broad, approximately triangular processes, of which in a well-

* Not yet identified.

+ This species has since (1906) been described by McClendon (13) under the

name of Myzostoma japonicum.

{ Another species of Myzostoma has since been recorded from Ophiacantha

vivipara (Koehler, Bull. Sc. France et Belg. vol. xli. p. 279), cf. also Fedotov (6).

MYZOSTOMIDA FROM THE RED SEA. 89

developed specimen there are 40-50, roughly corresponding to

the number of coste which terminate on them. These processes

are, of course, cirrl; they are better developed and more regular

in size in some individuals than in others, but in most cases they

present the appearance of typical cirri only on the anterior margin

in front of the mouth, where the body is thinner and more delicate

than posteriorly. The last pair of cirri is in most cases consider-

ably enlarged, forming a pair of caudal appendages into which

Text-fig. 17.

Myzostoma costatum.—Sketch of a mounted specimen 2°25 mm. in length. Caudal

appendages are not present and the individual is abnormal in possessing only

four parapodia and five suckers on one side of the body.

branches of the alimentary canal and uterus extend; in some

individuals these appendages reach a length equal to nearly one-

sixth of the length of the body. The penultimate and antepen-

ultimate pairs of processes are frequently enlarged also, but never

quite to the same degree.* The shape of the posterior extremity

* This characteristic enlargement of the posterior cirri was not described by

previous investigators of this species; v. Graft’s figure (7, pl. i. fig. 14a), however,

shows a distinct triangular incision at the posterior end of the body, indicating that

something of the kind occurred in the specimens examined by him.

90 DR. C. L. BOULENGER ON

of the body is subject to great variation, and it seems that the

development of caudal appendages bears no relation to the size of

the individual, these organs being absent in some specimens of

considerable size (cf. text-fig. 17), and, per contra, being well

developed in individuals under a millimetre in length.

The presence of caudal processes, as well as the number of these

organs, are characters which have been used to a considerable

extent for taxonomic purposes; the above description, however,

shows how unsafe it is to fix the limits of species by them.

The ventral surface is quite smooth; the five pairs of parapodia

have the usual radial arrangement, and their insertions are ap-

proximately equidistant between the centre of the body and the

periphery.

The structure of the parapodia is as described by v. Graff (7);

they recall those of M. cirriferum, although differing in being

rather more muscular. When fully extended they project as far

as the body-margin or even slightly beyond it, their extremities

being sometimes visible in a dorsal view of the animal.

The parapodia seem capable of movement in every direction,

and in his field-notes on this species Mr. Crossland remarks that

the little animals “ move their leg-like parapodia very actively,

like the legs of an insect, and can crawl actively about the arms

of the Ophiuroids and on a needle, but not on glass.”

The hooks are powerfully built, and their terminations strongly

bent, more so than in WV. cirriferum, being in this respect inter-

mediate between that species and J. glabrum.

Lateral to the third parapodium on either side is a large male

genital papilla; its insertion is close to the base of the para-

podium, and when completely protruded it can extend beyond

the margin of the body.

Both mouth and cloaca are ventrally situated close to the ex-

tremities of the body. Except in one individual the anterior

body-margin in front of the mouth is complete and without a

triangular incision. In the specimen figured on Pl. V. fig. 4,

such an incision is present. and in this respect it resembles those

described by v. Graff and Leuckart. I am, however, inclined to

consider this a post-mortem phenomenon due to the abrasion of

the delicate area which I mentioned above as occurring just in

front of the mouth.

In a ventral view of MW. costatum the “suckers” are quite con-

spicuous, and of these there are usually six pairs, all equally

developed and situated close to the periphery of the body. Four

pairs are in the usual position, that is to say in the interspaces

between the five pairs of parapodia, these are the second to

fifth pairs ; the first is situated in front of the first pair of para-

podia, close to the anterior margin, whilst the last or sixth pair of

suckers lies approximately halfway between the last pair of para-

podia and the posterior median termination of the body.

The suckers have much the same structure as those described

by Wheeler (17) in Jf. glabrwm, and when retracted appear as.

MYZOSTOMIDA FROM THE RED SEA. 91

spherical or oval bodies with walls so thick that the cavity

is almost obliterated or reduced to an irregular ramifying slit

between the folds of the wall.

Text-fig. 18.

Myzostoma costatum.—Sketch of an abnormal specimen 1°25 mm. in length. Only

two parapodia are present on one side of the body, and four on the other; the

number of suckers 1s normal.

The number of suckers in Mr. Crossland’s specimens is quite

constant, and, apart from some otherwise abnormal individuals,

there were only a few in which all twelve could not be made out

by means of the microscope. This is by far the most serious dis-

crepancy between my account of J/. costatum and that of previous

authors, and for some time I considered establishing a new species

to receive Mr. Crossland’s specimens. It seems probable, how-

ever, that the extra pairs of suckers have been overlooked ; even

when well developed these organs are difficult to make out on

unmounted specimens of Myzostomids, a fact which the majority

of workers on this group have commented on from time to time.

Leuckart’s description of MM. costatum was naturally incomplete,

and the few specimens seen by v. Graff seem to have been in a

rather poor state of preservation and much contracted ; no mounts

were made, and under the circumstances the extra suckers, if

present, might easily have remained unnoticed.

The presence of six pairs of suckers in a species of Wyzostoma

appears of considerable interest, since it necessitates some modifi-

cation in the definition of the group, as well as of the generally

92 DR. C. L. BOULENGER ON

accepted views on the morphology of the Myzostomida. W. M.

Wheeler, in his important paper on the sexual phases of these

animals (17), discusses the structure and homologies of the suckers

at some length. These organs (which he prefers to call segmental

sacs) he considers to be metameric, lying laterally or dorsally

to thew respective parapodia, like the ‘ Seitenorgane” of the

Capitellide, and for various reasons he believes them to be

homologues of these lateral line organs or segmental sacs. In

the course of his discussion of this subject we find the following

passage :—‘‘ The fact that in M/yzostoma there are five pairs of

pavapodia but only four pairs of segmental sacs, naturally leads to

the question as to what has become of the missing pair of sacs.

The answer to this question I believe we need not go far to seek ;

the third pair of the original five pairs of sacs has been converted

into the so-called penes. '‘lhese are more or less prominent

papille, lateral to and near the bases or the third pair of para-

podia. Each papilla is perforated by a ductus ejaculatorius, which

widens proximally into a vesicula seminalis. The latter receives

the mature spermatozoa from the vasa deferentia, and these in

turn from the ramifying testicular follicles. Bizarre as the de-

velopment of a male reproductive organ from a lateral line organ

may appear at first sight, I am nevertheless unable to see any great

difficulty in such a change of function. It is, in fact, easy to see

how the bottom of an eversible sac might acquire an opening into

the body-cavity under the pressure of a great accumulation of

spermatozoa ; the sac would then become reduced to a mere

conduit.”

A few years later von Stummer-Traunfels (16) investigated the

anatomy of the curious endoparasitie form J/. asterie Marenz.,

and showed that this species possesses a median sucker behind the

last pair of parapodia; whilst describing Wheeler’s view of the

homology of the penes as somewhat far-fetched, he considers this

extra sucker to represent the missing fifth pair, and his researches

on the innervation of this organ show that it must have been

formed by the fusion of two originally separate suckers.

Since the publication of my preliminary note (3) on the suckers

of WM. costatwm, Fedotov (6) has described a remarkable type of

Myzostomid, an endoparasite of the brittle-star Gorgonocephalus

ewcnenis Muller et Troschel; in this form, named Protomyzo-

stomum polynephris, five pairs of suckers are present, and these

are not situated between the pairs of parapodia on the ventral

side, but opposite to them and in a dorsal position. It is to be

hoped that further investigation of theanatomy of this new genus

may throw some light on the relations between suckers and

parapodia in the Myzostomids ; meanwhile, the occurrence of six

pairs of suckers in J/. costatum merely complicates the problem.

Granting the metameric nature of these organs, we must now

consider that we are dealing with six segments instead of five,

and it seems necessary to account for a missing sixth pair of

parapodia *.

* CF, also remarks under ‘* Neryous System” on p. 95.

MYZOSTOMIDA FROM THE RED SEA. 93

The above description of J. costatwm vefers especially to full-

grown specimens measuring between 13 and 4 mm. in length.

Young specimens under 14 mm. present a somewhat different

appearance; the body is thinner, and the cost on the dorsal

surface are less conspicuous than in the adult; though in some

eases very faint they are never absent. The marginal region is

more delicate and transparent than in the larger specimens, this

being due to the absence of ova in the peripheral branches of the

uterus ; 12-20 pairs of well-defined cirri are present, and between

them can be seen smaller processes which are about to give rise

to more of these organs. The last pairs of cirri show the same

tendency towards enlargement as in the adults. The suckers are

not very prominent, but the male papille are fully developed and

are extremely conspicuous.

Before concluding my account of the external features of this

species, I wish to call attention to the fact that quite a considerable

number of specimens in Mr. Crossland’s collection present abnor-

malities chiefly regarding the number of parapodia. In these

specimens one or more parapodia are missing from one or both

sides of the animal, and, although in some cases this may be due

to injury and incomplete regeneration of parts of the body, in

others the malformations must have been congenital. I have

figured some of these abnormal specimens; the sketch (text-

fig. 18) on p. 91, shows an extreme case, that of an individual

measuring 1°25 mm. in length, which possesses four parapodia on

one side of the body and only two on the other; the rest of the

animal seems normally developed, and the full number of suckers

is present. In some cases reduction in the number of parapodia

is accompanied by a similar reduction in the number of the

suckers (cf. text-fig. 17, p. 89).

Alimentary Canal.

As mentioned above, the mouth is situated on the ventral side

close to the anterior extremity of the body; through it can be

protruded a large and muscular pharynx provided at its distal end

with small papille, as described by v. Graff; these are very in-

conspicuous in all the specimens before me, and, so far as I can

ascertain, number four only.

The stomach is elongated, extending back as far as the level of

the last pair of parapodial bases, and constricted laterally at the

points where the intestinal branches arise, almost opposite the

male genital papille.

There are only two intestinal trunks on each side arising close

together from the stomach; these divide into from 7—9 primary

branches, which in their turn radiate outwards and give rise to

numerous diverticula, which end blindly on the body-margin.

The rectum is separated from the stomach by a distinct valve-

like constriction, and takes the form of a straight tube running

back from the level of the last pair of parapodia to the cloaca,

where it is joined by the oviduct.

94 DR. C. L. BOULENGER ON

Surrounding the base of the pharynx and the extreme anterior

end of the stomach is a thick ring of cells (cf. Pl. VIII. fig. 3)

resembling the ‘‘ multipolar” cells described by Nansen ((15),

p. 71) as occurring close to the cireum-cesophageal nerve-ring in

several species, and which he considered to be ganglionic cells

forming a kind of sympathetic system. Similar cells were found

by v. Stummer (16) in J/, asteriw, but this author showed that

Text-fig. 19.

Myzostoma costatum.—Longitudinal horizontal section through part of the

body to show the position of the ovaries and nephridia.

st. Stomach. rect. Rectum. iné. Intestinal trunk. ov. Ovary. neph. Nephridium.

their minute structure differed considerably from that of gan-

glionic cells, and that the presence of secretion granules and

capillary ducts indicated that they were of a glandular nature.

The cells surrounding the anterior end of the stomach of

M. costatwm very closely resemble those described by v. Stummer

(cf. (16) p. 551, pl. xxxvii. figs. 5 & 6); in this species, however,

they are more numerous and not so scattered; moreover, the

capillary ducts are all directed towards the gastric epithelium.

MYZOSTOMIDA FROM THE RED SHEA. 95

I think there can be little doubt but that v. Stummer was right in

considering these cells to be unicellular glands. In I. costatum

these glands may have a digestive function ; they certainly form

the only glandular tissue in connection with the alimentary canal

in this species.

Nervous System.

The best and most complete account of the Myzostomid nervous

system is that published by Nansen in 1885 (15); it was based

on investigations made on several species, especially W/. gigantewm

Nansen, WV. graffi Nansen, and WV. cirriferwm Leuckart. In these

forms, which can be regarded as quite typical, the nervous system

consists of a feebly-developed circumpharyngeal ring communi-

cating with a ventrally-situated unsegmented nerve-mass, from

which eleven pairs of nerves arise. Of these nerves five on each

side are large and supply the five parapodia, as well as the parts

of the body-margin on both sides of them, whilst the other six

are more slender and alternate with the larger nerves supplying

the parts of the body between the parapodia.

Text-fig. 20.

né

Diagram to show the origin of the paired nerves from the ventral nerve-mass in

Myzostoma costetum.

N!_N5, The five pairs of large nerves. !-n®, The six pairs of smaller nerves.

C. Circumpharyngeal commissures.

Nansen did not state the nerve-supply of the “suckers,” but

v. Stummer (16) believes these organs to be innervated by branches

96 DR. C. L BOULENGER ON

of the small nerves, at least in the species which he examined,

M. asterie Marenz.

I spent some time in attempting to reconstruct the nervous

system of M/. costatwm, as I hoped that the distribution of the

paired nerves might shed some light on the problem of the homo-

logies of the additional suckers in this species ; the diagram (text-

fig. 20) on p. 95 gives, I think, a tolerably accurate idea of the

nerves arising from the ventral nerve-mass.

It will be seen from the diagram that the nervous system of

M. costatum is remarkably similar to that described by Nansen

in other species, and especially to that of JZ. cirriferwm; the

number of nerves arising from the central mass is normal; there

are eleven pairs, of which five are large and branch chiefly to the

parapodia. I agree with v. Stummer in believing the suckers to

be supplied by branches of the smaller intermediate nerves; the

six pairs run almost straight out towards the six pairs of suckers,

but I was unable to trace the finer branches actually into these

organs.

It will be seen from my diagram that in J/. costatwm the origin

of the six pairs of smaller nerves differs somewhat from that in

the species described by Nansen; the first on each side seems to

originate from the circumpharyngeal commissure, whilst the re-

maining five are given off posteriorly from the bases of the five

larger nerves to the parapodia.

If we take for granted that the suckers in Myzostoma are

‘‘Seitenorgane” homologous with those of other Annelids and

therefore parts of the parapodia—and I think Wheeler and

v. Stummer have made out a very strong case for such a

homology,—then it becomes obvious from the above account that

each sucker belongs to the parapodium anterior to it. In the

majority of species the fifth pair of parapodia has lost its

‘“‘Seitenorgane,” these being retained in four species only —

M. asterie, M. costatum, M. mebianum*, and Protomyzostomum

polynephris. \n the latter form they have remained in the more

normal position, 7. ¢., lateral and dorsal to their respective para-

podia. ‘The innervation of the first of the six pairs of suckers in

M. costatwm from the circumpharyngeal commissures suggests

that these may belong to a peristomial segment from which, as

in other Annelids, the locomotory portions of the parapodia have

disappeared.

I wish again to emphasize the fact that I have been unable to

prove that ihe suckers really are innervated from the six pairs of

finer nerves, and that therefore the above conclusions are highly

speculative. I think, however, that they afford a satisfactory

explanation of the variation in the number of suckers in this

eroup.

Sexual Organs.

Like nearly all well-investigated species of M/yzostoma, MW. cos-

tatum is hermaphrodite, and all the larger specimens in

* Cf. Boulenger (8, p. 350).

MYZOSTOMIDA FROM THE RED SEA. om

Text-fig. 21,

CLP

Nb. ar aa

Myzostoma costatum.—Three transverse sections through the centre of the body :

as in text-fig. 19, only the outlines of the various organs are shown and the

testicular-follicles are omitted. The sections are slightly oblique, so that

right and left halves of the figures show different regions.

st. Stomach. int. Intestinal branch. meph. Nephridium. x. Ventral nervous

system. ov. Ovary. o. Anterior narrow end of the oviduct. w. Median

division of the uterine celom. wt. Branch of the uterus.

Proc. Zoou. Soc.—1913, No. VII. 7

98 DR. C. L. BOULENGER ON

Mr. Crossland’s collection are in what Wheeler (17) terms the

androgynous phase, both male and female organs being fully

developed and functional *. The smaller specimens, however,

exhibit a marked protandric condition; as already mentioned,

those under about 1:5 mm. in length possess a well-developed

male apparatus of relatively greater size than in the adults.

Closer examination of such specimens reveals that whilst ripe

spermatozoa are present in abundance, the female organs are but

poorly developed, and mature ova absent or rare in the uterus

and oviduct.

Male Organs.

The testes, as in other Myzostomids, are much branched; the

ramifications of each organ are for the most part restricted to

the ventral parenchyma. below the alimentary canal, but occasional

dorsally directed branches are given off between the intestinal

and uterine diverticula. There is a single vas deferens on each

side of the body, which widens out distally to form a vesicula

seminalis, usually packed full of ripe spermatozoa, at the base of

the genital papilla. A short ductus ejaculatorius lined with

ectoderm leads through the papilla to the exterior.

Female Organs.

M. costatwm possesses a single pair of ovaries situated near the

middle of the body, ventro-laterally to the stomach, and occupying

the spaces between the four main intestinal trunks (text-figs. 19,

21 A). In structure the ovaries resemble those of MW. glabrum as

deseribed by Wheeler (17), being irregular in shape; as a matter

of fact they are to be considered as formed by the proliferation

of the peritoneal epithelium at the terminations of ventrally

directed diverticula of a small ccelomic space, which occupies a

transverse position dorsal to the stomach in this region. The

uterus, as in other species, consists of a series of ramifying

celomic diverticula lying dorsal to the intestine and correspond-

ing roughly, both in number and arrangement, with the main

branches of this organ. ‘The uterine branches, like those of the

intestine, radiate outwards from near the centre of the body,

and arise from the lateral terminations of the transverse coelomic

space mentioned above (text-fig. 22). This space and its ven-

trally directed diverticula which connect it with the ovaries are

lined by a low epithelium of small ciliated cells; in the uterine

branches, which in adult specimens are filled with ova in various

stages of development, this epithelial lining is absent, bein

replaced by a ‘“ pseudoepithelium ” similar to that described by

von Stummer-Traunfels in MW. asteriw (16).

The oviduct is sharply marked off from the uterus; it lies dorsal

to the stomach and rectum in the middle line, and has the form

of a straight wide tube, narrowing somewhat posteriorly, and

* Cf. also Coventry (8).

MYZOSTOMIDA FROM THE RED SEA. 99:

much dorsiventrally compressed along its whole length (text-

fig. 21 C); the organ is iined throughout by a distinct epithelium

_ Text-fig. 22.

Diagram to show the relations between the uterus, oviduct, aud nephridia

in Myzostoma costatum.

Ut. Uterus. Ov. Oviduct. O. Opening of oviduct into the uterine eclom.

JV. Internal openings of the nephridia. C. Position of the cloaca.

of comparatively large ciliated cells. Anteriorly the oviduct

communicates with the central ceelomic space by means of a

100 DR. C. L. BOULENGER ON

short, narrow duct (text-figs. 21 B and 22, and Pl. VIII. figs. 2

and 4), also lined by a ciliated epithelium; the latter, however,

consists of very small cells with long cilia, and seems continuous

with the epithelium lining the ccelom. The diameter of this

tube is less than half that of a mature ovum such as one finds

in the uterus; it is, however, surrounded by transversely

arranged muscle-fibres, which no doubt allow the dilatation of

the tube necessary for the passage of such ova into the oviduct.

The above account shows that, at least in JZ. costatum, the

oviduct is quite a distinct structure, and I have therefore used

for it the name employed by the older investigators of Myzo-

stomid anatomy. I cannot follow Wheeler and v. Stummer-

Traunfels, who refer to this organ as a median posterior branch

of the uterus,

Nephridia.

The nephridia resemble those described by Beard (2), Wheeler

(17), and v. Stummer-Traunfels (16) in other species. They have

the form of a pair of bent tubes opening anteriorly into the dorsal

ceelomic space (text-fig. 22), and sloping outwards and backwards

towards the ventral surface so as to come to run parallel with the

alimentary canal (text-fig. 21); they open posteriorly by separate

openings into the commencement of the rectum (text-fig. 19).

Along the greater part of their course the nephridia are of

considerable diameter and are lined by large glandular cells with

long cilia; they, however, narrow considerably towards their

anterior ends, where the lining cells become much smaller and,

as a matter of fact, the nephridial epithelium passes gradually

into that of the celom. In this respect I am in agreement with

Maidl (12), who in a recent paper described much the same in

M. glabrum.

The nephrostomes are situated on either side of and very close

to the internal opening of the oviduct, although not quite in the

same plane as this structure, being slightly dorsal to it ; all three

openings can, however, be seen in one horizontal section.

The nephridia contain numerous immature and fragmentary

ova (text-fig. 21 C), but, like Wheeler and other investigators since

Nansen, I have never seen complete, ripe ova in this situation.

The similarity in structure and position between the nephridial

and oviducal openings in J. costatwm is very striking, and has,

perhaps, some bearing on the homologies of these organs. Nansen

(15), who described a very similar arrangement in JZ. carpenteri,

referred to them as three oviducts, and we must remember that

before his discovery of the “lateral oviducts” Beard (1) had

suggested that ‘the oviduct opening into the cloaca may perhaps

originally have been formed from two segmental organs, for the

cloaca is an epidermic invagination, and if it were obliterated the

oviduct would open on the median line.” That partial fusion of

a pair of segmental organs can take place has been shown by

Wheeler (17), who described the nephridia of J/. glabrum as

MYZOSTOMIDA FROM TILE RED SEA, 101

possessing a single nephrostome as well as an unpaired end-piece

opening into the rectum; a similar condition is to be found in

M. cryptopodium, where, however, the median end-piece opens

directly to the exterior on a papilla just under the cloacal orifice.

Moreover, according to Wheeler, in J/, belli the cloaca, oviduct,

and nephridial end-piece open separately on the posterior surface

by three ciliated apertures.

Beard’s suggestion as to the nature of the oviduct was forgotten

when the nephridia were discovered, and, whilst not wishing to

commit myself to this theory, I recall it as one to be borne in

mind. Perhaps Protomyzostomum polynephris will afford clues to

a number of such problems in the morphology of this difficult

group of animals.

MyzosToMA RUBROFASCIATUM Vv. Graff. (Pl. VI. fig. 1.)

M., rubrofasciatum v. Graff (8), p. 33, pl. u.

In Mr, Crossland’s collection are two specimens of a J/yzostoma

from Foreulch Barrier Island, obtained from a Comatulid which

unfortunately cannot now be traced.

The animals are of an oval shape, but, owing to the contraction

of the bodies being accompanied by a curling-up of the anterior

and posterior ends (¢f. Pl. VI. fig. 1), it is impossible to give exact

measurements. One specimen isa little larger than the other, and

must have had a length of about 24 mm., with a maximum

breadth of 2 mm.; the smaller specimen measures approximately

mm. by 1:75 mm.

The body is of considerable thickness ; on the dorsal side is a

very conspicuous median longitudinal elevation running from one

extremity to the other; narrow at each end, it is very broad

towards the centre of the body, where it reaches a breadth almost

one-fifth of that of the whole animal. From the sides of the

median elevation arise seven to eight broad radial ones extending

to the body-margin, where they end in short blunt processes.

Between these primary radial “coste” are a number of shorter

secondary and tertiary ones, these are also of considerable

breadth, and terminate in marginal processes.

The coste are so numerous near the edges of the body as to

come in contact with one another laterally, thus forming a thick

ribbed margin, a very conspicuous feature of this species. As

mentioned above, the body-margin presents a series of short blunt

processes formed by the terminations of the radial elevations ;

there are between 25 and 30 pairs of these processes, which are

very irregular in size and shape. In both specimens, however,

the last pair and, to a slightly less degree, the penultimate and

antepenultimate pairs, are considerably enlarged and thickened,

The coloration of the dorsal surface in the preserved specimens

is peculiar and very characteristic; the median elevation and the

coste are of a yellowish colour; the rest of the dorsal surface is

brownish, except where a pair of narrow light-coloured streaks

102 DR. C. L. BOULENGER ON

run longitudinally down the body, each about halfway between

the middle line and the margin. Although expressed somewhat

differently, it will be seen that this colour- -pattern is practically

identical with that described by v. Graff (8) in 1. rubrofasciatum,

a Myzostomid obtained from an unidentified Crinoid in the Red

Sea at Tur, near Sinai. I think there can be no doubt that

Mr. Crossland’s two specimens must be referred to the same

species.

v. Graff’s description was based on a single individual, which

was more brightly coloured than those I am describing, moreover,

at first sight it seems to differ considerably in shape from those

specimens. ‘This difference is, in my opinion, due entirely to the

mode of contraction, the body i in v. Graff’s specimen having the

sides bent ventrally so as to give it the form of a boat, the median

elevation representing the keel.

Owing to the state of contraction of his only specimen, v. Graff

was not able to describe the ventral surface at any length. In

the individuals collected by Mr. Crossland the ventral-surface is

yellowish in colour, and the central muscular mass prominent.

The five pairs of parapodia are well developed and, as in

M. costatum, without division into proximal and distal regions ;

they arise approximately halfway between the centre of the body

and the margin.

There are four pairs of suckers of relatively large size with

radially folded walls. They are in the normal position between

the parapodia, and are situated close to the periphery.

Both mouth and cloaca are subterminal. hk

The male papille are not conspicuous ;. they have the form of

short conical processes in the usual position lateral to the third

pair of parapodial bases.

I did not feel justified in cutting sections as I had only two

specimens of this species. at my disposal, and neither seemed likely

to yield good results. One individual was cleared in cedaiwood

oil, but owing to the thickness of the body rev ced practically

nothing of the internal organisation.

MyzosToMA CROSSLANDI, sp.n. (Pl. VI. figs. 2, 3; Pl. VIL,

& PMV aL tie. 41)

From various localities in the Red Sea Mr. Crossland obtained

four specimens of a large elongated Myzostomid (Pl. VI. figs. 2, 3)

which is evidently closely related to M. nanseni v. Graft, but suffi-

ciently distinct, in my opinion, to justify the creation of a new

species to receive it. This form, which I name M. crosslandi,

is In certain respects inter mediate between WM. nanseni and

M. folium v. Graft.

Of the four individuals one was found at Ul Shubuk on Antedon

savignyt, another in Suez Bay at a depth of 10 fathoms on

Antedon serripinna, whilst the other two were obtained from a

Comatulid which J have been unable to trace.

As mentioned above, this Myzostomid is of comparatively large

MYZOSTOMIDA' FROM THE RED SEA, 103

size, the lengths of the specimens measuring 85, 8, 8, and 5 mm.

respectively.

The bodies have a maximum thickness of about 1 mm.; owing

to the fact that all four specimens were contracted with the sides

incurved ventrally, it is impossible to give exact measurements of

the breadths, these, however, I estimate as varying between 3 and

3'5 mm, in the different individuals.

The body-dise is rounded off anteriorly and posteriorly ; it does

not terminate in a point, as in M/. foliwm. The thick body thins

out gradually towards the edges, and has a narrow translucent

margin which is finely notched, giving rise to a number ot incon-

spicuous marginal processes ending in fine points.

The dorsal surface is smooth; a median longitudinal elevation

is noticeable in three specimens, and is conspicuous only poste-

riorly in the region of the oviduct; on either side are feeble

elevations corresponding in position to the parapodial insertions.

Mouth and cloaca are to be seen on the ventral side, situated

on small] papilla and both close to the extremities, in this respect

differing from MW, nanseni, where the cloacal papilla lies at the

commencement of the last quarter of the body, A narrow median

ridge projects on the ventral surface, and marks the position of

the stomach and rectum.

The parapodia are well developed and arranged in two almost

parallel rows; the first pair lies close behind the mouth, but the

insertions of the last pair are, as in W/. nanseni and M. foliwm, at

a considerable distance from the posterior termination of the body,

this distance varying in the different specimens between a quarter

and a third of the total length of the animal, Hach parapodium

is strong and, as in the two species just mentioned, consists of a

broad muscular region, situated proximally, and of a narrower

distal region provided with a conspicuous longitudinal groove on

its ventral face.

The male papilla ave inconspicuous and have the form of short

conical processes in the usual position (text-fig. 23, p, 104), At first

I thought that suekers were absent, and I could detect none under

a low magnification; the use of a higher power of the micro-

scope, however, revealed up to four pairs of these structures

between the parapodia and close to the body-margin (text-fig, 23).

They are much smaller than those deseribed by v. Graff (9) in

MM. nanseni, and are, J think, in a vestigial condition.

The colour-pattern of WM. crosslandi is quite characteristic; the

dorsal surface in the spirit specimens varies from a greenish grey

to a brown colour, with seven or more pairs of very narrow lighter

bands running transversely across the body at regular inter vals.

T have no information as to the colour of fresh individuals,

Among a number of young specimens of J/. costatwm from

Ul Shubuk I found two small individuals which I take to be

young stages of WM. crosslandi. They measure 1:25 x 1 mm. and

1 x ‘75 mm. respectively. The last pair of parapodia hes much

closer to the posterior extremity of the body than in the specimens

104 DR. C. Le BOULENGER ON

just described, but, except for this and the proportions of

length to breadth, they resemble the adults in every respect.

Neither cirri nor suckers are better developed than in the latter,

and, as a matter of fact, I could not trace any suckers at all in

one of these young specimens,

Text-fig. 23.

‘

y eNOS

Myzostoma crosslandi, sp. n.—Anterior part of the body; sketch made from

a specimen cleared in cedarwood oil.

M. Mouth. P. Male papilla. S. Vestigial sucker.

No complete account of the anatomy of any of the large

elongated Myzostomids has yet been published. 1 therefore used

one of the larger specimens of JM. crosslandi for the preparation

of sections; when merely cleared in cedarwood oil they showed

little more than just the peripheral parts of the alimentary canal.

MYZOSTOMIDA FROM THE RED SEA. 105

A series of transverse sections was obtained, and they

show a very characteristic feature of this species, namely, the

great development of the ‘“ Hautmuskelschlauch,” the muscle-

sheath below the skin (Pl. VII., & Pl. VIII. fig. 1); it is best

developed on the dorsal side of the animal, where it is separated

from the outer epithelium by a cutis-like layer of connective tissue.

This cutis is absent from the ventral surface, and here the muscle-

sheath can be seen to consist of distinct outer and inner layers of

transverse and longitudinal fibres respectively.

Alimentary Canal.

The mouth leads into a muscular elongated pharynx, which,

when retracted, extends back in the body almost as far as the

level of the second pair of parapodia, The pharynx was in this

condition in all four specimens, and I was therefore unable to

make out whether terminal papille occur. ‘here is no distinct

esophagus, the pharynx leading directly into the stomach; this

part of the alimentary canal is comparatively narrow and of

considerable length ; slightly sacculated anteriorly, it gives off on

each side three intestinal trunks which arise close together at a

point just posterior to the level of the third pair of parapodial

insertions. The central of these three trunks branehes profusely

in the normal manner, but supplies only a small area of the body

with intestinal ceca; the anterior and posterior trunks, however,

continue in a straight course forwards and backwards respectively

to the two extremities and, keeping close to the middle line, supply

the rest of the body with numerous almost parallel branches given

off from their external sides only. This peculiar and very cha-

racteristic arrangement of the intestinal branches (text-fig. 23)

recalls that described by v. Graff in I. elongatwm (7); it seems

probable that it will be found to occur in all the elongated species.

The rectum commences a short distance behind the last pair of

parapodia and runs backwards as a straight narrow tube, which

opens on a small but conspicuous papilla (Pl. VIII. fig. 1) close

to the posterior margin of the body.

T could find no trace of salivary glands in connection with the

alimentary canal, and the peculiar glandular tissue described as

surrounding the anterior part of the stomach in UM. costatum is

very poorly developed in this species,

Reproductive Organs.

The single specimen sectioned proved to be in an androgynous

condition, and both sets of organs seemed equally developed.

Male Organs.

The testes (Pl. VII.) are embedded in the parenchyma, chiefly

ventral to the alimentary canal, but a certain number of follicles,

asin M, costatum, extend upwards between the intestinal branches

106 DR. C, L. BOULENGER ON

and come to occupy a dorsal position between or above the uterine

cxeca.,

The testicular follicles are in some cases enclosed in small

cavities which, in agreement with most recent workers, | consider

to be ceelomic ; these cavities are in places considerably enlarged,

and are then filled with large masses of mature spermatozoa

(Pl. VII. fig. 5); their size equals or exceeds that of the uterine

spaces, which they resemble except for the nature of their contents.

Some of these cavities seem to have lost their communications

with the rest of the male organs, and I am at a loss to explain

how the spermatozoa. within them can make their way to the

exterior,

Wheeler seems to have observed a somewhat similar condition

of the male organs in his species J/. cireinatum, but in this form

vesicule seminales and ‘ penes ” appear to be absent; in WV. cross-

landi the majority of the testicular follicles communicate with the

vasa deferentia, and through these with seminal vesicles which

open on to the short male papillae.

female Organs,

The female organs of JV. crosslandi are very similar to those

just described by me in J, costatum. The single pair of hollow

ovaries (Pl. VII, fig. 1) lies ventro-laterally to the stomach just

behind the point of origin of the first pair of intestinal trunks.

The cavities of the ovaries communicate with a pair of narrow

celomic tracts which run upwards along the sides of the stomach

to join the small median celomie space from which the uterine

branches are given off., The latter are numerous and relatively

very slender ; they follow the main ramifications of the intestine.

As in WM. ecostatwm, the oviduct communicates with the median

uterine ccelom by means of a narrow ciliated duct, which in this

species is rather long. The oviduct itself is very slender and

much compressed so as to appear crescentic in transverse sections;

it decreases in width considerably in the region of the rectum

and opens on the cloacal papilla by a very narrow terminal tube.

Nephridia,

The nephridia (Pi. VII. figs. 2-5) oceupy very much the same

position as in J/. costatum, and, as in that species, open posteriorly

into the sides of the rectum close behind the commencement of

that organ. Unlike J/. costatwm, the anterior ends of the nephridia

do not communicate directly with the median uterine cceelom, but

open separately into the narrow anterior end of the oviduct, so that

the three tubes which afford communication between the ccelom

and the exterior possess but a single internal opening.

The anterior terminal part of each nephridium has the form of

a transversely set narrow tube lined with long cilia ; this widens

out laterally when the organ follows a rather sinuous course,

backwards and downwards, and comes to form a broader tube

MYZOSTOMIDA FROM THE RED SEA. 107

lying ventral to the stomach. The nephridia run side by side

ventral to the stomach for a short distance, but in the posterior

region of that organ slope upwards again, finally lying lateral to

the rectum just before opening into it.

Nervous System.

I was able to make out very little of the nervous system of this

species, as the peripheral nerves were only very poorly stained in

my sections. ‘The central nerve-mass was, however, distinct and

occupied the same position as in other species, that is to say the

centre of the body, in the region between the second and fourth

pairs of parapodia.

This typical concentration of the central nervous system is a

point of interest, since it shows that the. elongated form of

M. crosslandi is probably not a primitive character ; the arrange-

ment of the intestinal branches and other organs also indicates

that this species is derived from some form possessing a more

radial symmetry.

_ This is probably also true of allied species, such as WM, nansenr

and WM. foliwm.

List oF REFERENCES.

1. Brarp, J.— On the Life-History and Development of the

Genus Myzostoma (F. 8. Leuckart).” Mitth. Zool. Stat.

Neapel, Bd. v. 1884, p. 544.

. “The Nature of the Hermaphroditism of Myzostoma.”

Zool. Anz. Bd. xvii. 1894, p. 399.

_ Boutencrer, C. L.—‘The ‘Suckers’ of the Myzostomide.”

Zool. Anz. Bd. xxxvii. 1911, p. 346.

Crarx, H. L.— <A new Host for Myzostomes.” Zool. Anz.

Bd. xxv. 1902, p. 670.

5. Covenrry, A. F..—“The Application of Mr. G. W. Smith’s

Theory of Dwarf Males to Myzostoma.” Ann. Mag. Nat.

Hist. ser. 8, vol. v. 1910, p. 378.

6. Feporov, D. —“ Protomyzostomum polynephris, eme neue

Myzostomidenart.” Zool. Anz. Bd. xxxix. 1912, p. 649.

7. Grarr, L. v.—‘Das Genus Myzostoma (F. 8. Leuckart).”

Leipzig, 1877.

E> fo

8. “Report on the Myzostomida collected curing the

Voyage of H.M.S. ‘Challenger’ during the Years 1873—

76.” Rep. ‘Challenger’ Exped. vol, x. Pt. 27, 1884.

She Supplement to the preceding Report. Rep. ‘Chal-

lenger’ Exped. vol. xx. Pt. 61, 1887.

10. Leucwart, F. S.—‘‘ Bericht tber die achte Versammlung

deutscher Naturforscher und Aerzte zu Heidelberg im

September 1829.” Oken’s Isis, 1830, Heft v. p. 612.

“ Zoologische Bruchstiicke. III. Helminthologische

Beitriige.” Programm zur Eroéffnung der Wintervorles

ungen in Freiburg i. B. 1842, p. 5,

ple

108 ON MYZOSTOMIDA FROM THE RED SEA.

12. Marv, F.—“ Ueber die Ceelomverhiltnisse von Myzostoma.”

Verh. zool.-bot. Ges. Wien, Bd. lx. 1910, p. 200.

13. McCuienpon, J. F.—‘‘ The Myzostomes of the ‘ Albatross’

Expedition to Japan.” Bull. Amer. Mus. Nat. Hist.

WO o-dil, UNO, Jaq 1.9).

‘“New Marine Worms of the Genus Myzostoma.”

Smithson. Inst. U.S. Nation. Mus. Proc. vol. xxxii. 1907,

p- 63.

15. Nansen, F. —“ Bidrag til Myzostomernes Anatomi og

Histologi.” Bergen, 1885,

16. Stummer-TRAUNFELS, R. y.—“ Beitriige zur Anatomie und

Histologie der Myzostomen. J. Myzostoma asterie

Marenz.” Zeitschr. f. wiss. Zool. Bd. Ixxv. Heft 4, 1903,

p- 49.

17. Wueeter, W. M.—‘The Sexual Phases of Myzostoma.”

Mitth. Zooi. Stat. Neapel, Bd. xii. Heft 2, 1896, p. 227.

14.

EXPLANATION OF THE PLATES.

-PuateE V,

Figs, 1-6. Myzostoma costatum F. S. Leuckart.—Series of specimens illustrating

the variation in the shape of the body and the development of the coste.

The individuals shown in figs. 2 and 3 possess an abnormal number of

parapodia (cf. p. 93). In all the figures a represents the dorsal, 6 the

ventral aspect of the animal. X12.

Pruate VI.

Fig. 1. Myzostoma rubrofasciatum v. Graff.—Dorsal view. X18.

Figs. 2-3. Myzostoma crosslandi, sp. n.—Two specimens, dorsal and ventral aspects.

X12.

Prate VII.

Figs. 1-5. Myzostoma crosslandi, sp. n.—Transverse sections to show the anatomy.

em. Central nerve-mass. int. Branch of intestine. m. Muscular

sheath. 2. Nephridium. o. Ovary. ov. Oviduct. 7. Rectun.

sp. Mass of spermatozoa occupying ccelomic space. s¢. Stomach.

ée. Testis. ut. Branch of the uterus.

Fig. 1. Section passing through the ovaries.

Figs. 2-4. Sections passing through the anterior portion of the oviduct and showing

the position of the nephridia,

Fig. 5. Section passing through the posterior parts of the stomach and oviduct.

Prate VIII.

. Myzostoma erosslandi, sp. n.—Photograph of a transverse section through

the posterior region of the body, passing through the cloacal papilla

(el.p.). Lettering as in Plate VII.

. Myzostoma eostatum K.S., .—Photograph of a longitudinal section showing

the principal organs. ph. Pharynx.

Fig. 3. Myzostoma costatum ¥.S. L. Photograph of a transverse section passing

through the anterior extremity of the stomach and showing the ring of

glandular tissue (gl.) developed round the alimentary canal in this

region.

Fig. 4. Myzostoma costatum ¥.S. L.—Part of the same section as fig. 2, under a

higher magnification. gl. Glandular tissue.

i i eae OI) oe ra

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43)

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: BEASTWOODZAS,

TALITRIATO

AMC SAU Ohes IE ASE ONO) DA.

P ZS Cie) Pa eae

i Mea

Fhath imp,

ON A NEW AMPHIPOD FROM THE TRANSVAAL. 109

8. Description of an Amphipod belonging to the Family

Talitridee, from the Woodbush, Transvaal. By Patt

A. Mrruugn, F.Z.S.

[Received July 7, 1912 ; Read November 26, 1912].

(Plates X. & XI.*)

INDEX,

Systematic : ; Page

I NTKEPUCIADIA, (XC 5 IONS onecar.coosso coooeonnacootoocosene: LO)

Ab GTGDOOL, (IVS 105 cooodn onsscenns sb odooce uno copeouene | LlKO)

A few months ago I received from Miss Audrey Eastwood

some Gammarids from streams near Mr. Kastwood’s farm, Clear-

water, in the Woodbush, Northern Transvaal. Seeing that

these are the first Gammarids recorded from the fresh-waters of

South Africa other than near the sea, they are of some little

interest.

The specimens sent belong to one species and to a family

whose members, though they are usually littoral marine forms,

have often been recorded from localities far from the sea and at

some considerable altitude in other countries.

Thus Valitrus sylvaticus has been found in Tasmania up to a

height of 760 metres on Mount Wellington, and Chiltonia in New

Zealand in mountain streams up to 450 metres and more. The

genus /yalella is restricted to fresh-waters, occurring according

to Stebbing (in ‘Das Tierreich’) “in depths above the sea-

level extending to 4053 metres”; thus in South America, Lake

Titicaca supports a number of deep-water forms.

We can probably derive most of the fresh-water species in

this family directly from marine ancestors. In the Cape some

littoral marine forms do actually ascend for a short way up streams

near the coast. But until this record from the Woodbush none

had been discovered at any distance from the sea.

I think we can safely say that this new species has been

independently derived from a marine form. Though a new

genus has been made for its reception, yet it is very closely

related to the genus Talitrus, in which genus it may have to be

included eventually.

The fact that there is an Amphipod in this country which

thrives in mountain streams at least in one locality is very

suggestive as to what might be done to increase its numbers and

range for the purpose of trout food; and I think no one would

dispute its value for this purpose.

Genus TALITRIATOR, gen. nov.

Diagnosis.—Like Talitrus except for the fifth side-plate and

the following characters :—

Antennule is slightly shorter than peduncle of antenna. The

* Wor explanation of the Plates see p. 112.

110 THE HON. P. A. METHUEN ON A

fourth joint of the maxilliped is present. Gnathopod I not so

long as gnathopod IT and not stronger; fifth joint strong and

swollen.

TALITRIATOR EASTWOOD, Sp.n. (Pls. X. & XI.)

Length, not counting antenne, with pleon bent 7:2 mm.; no

striking difference between the male and female.

Antennule (Pl. X. fig. 1).—The third joint slightly the longest.

On the surface each joint carries distally two fairly long bristles.

On the upper surface. opposite the ventral groups of sete are

groups of bristles, shorter than those borne ventrally, each group

either of two or three bristles; on the basal joint a single bristle.

The last segment, which is very short, carries two stiff bristles.

The number of segments is nine or ten.

Antenna (Pl. X. fig. 2).—Penultimate joint of peduncle is

nearly twice the length of the proximal joint; the distal nearly

twice the length of the penultimate. All the segments of the

flagellular part are provided distally with a whorl of bristles,

there being four groups generally to each segment, three bristles

to each group. The last segment ends in a compact group of

fine bristles.

Upper lip (Pl. X. fig. 3) rounded, with numerous hairs at

the extremity.

Lower lip (P1. X. fig. 4).—The lobes with long and short sete ;

the mandibular processes, which are fairly well developed, possess

minute sete.

Mandible.—Palp absent : otherwise normal,

Mawillula.—Much as in Talitrus, with nine claw-like toothed-

bristles—five larger and four smaller—furnishing the outer plate.

The inner plate with two setose spines, and a few hairs proximally

along the inner margin. The palp exceedingly small, with hairs

on outer margin; it is apparently two-jointed, the distal joint a

minute glabrous blunted spine.

Mawilla.—As for Talitrus locusta, normal.

Mawilliped (Pl. X. fig. 5).—Inner plates are remarkable in

that they are furnished with three glabrous acorn-shaped processes

on each side; they bear also a number of setose spines. The

palp is four-jointed.

Gnathopod I (Pl. X. fig. 6).--Coxal plate narrow, with a few

spines of unequal size on the ventral margin ; the other six joints

armed with a few spines on both margins, longest on the fourth

joint. Fifth joint swollen. Sixth joint with six stout, slightly

curved spines on the posterior margin and one slightly smaller

than the other six, together with small bristles at the base of

most of these spines.

Gnathopod II (Pl. X. fig. 7).—Coxal plate excavate behind

with conical projection; lower border armed with short stout

spines. The second joint is not swollen or expanded; it is much

longer than any of the other joints. The third joint is narrower

than the second, slightly swollen about the middle of anterior

NEW AMPHIPOD FROM THE TRANSVAAL. lel

margin; two small spines on posterior margin distally. The

fourth joint short, the shape of a rhombus, with setose cushion

on posterior margin and two stout bristles just behind the

cushion. The fifth joint longer than the third or sixth, broadest

at its distal base, and expanded also about the middle; a setose

cushion extending along nearly the whole of the free posterior

margin ; a thickened chitinous enlargement on the distal part of

the cushion; four bristles just behind the cushion placed subequi-

distant apart. The sixth joint nearly as long as the fifth; the

posterior margin also with minute sete. Behind the setose part

a number of stout bristles.

Pereiopod I.—Not quite as long as pereiopod II, the coxal plate

like that of gnathopod IT, excavate behind with conical process.

The second joint linear; the third joint shortest and convex

behind; the fourth joint linear, broadest distally ; the fifth and

sixth joints linear. The sixth joint longer and much narrower

than the fifth ; the last joint small, with claw.

All the joints with spines which are most numerous on the

anterior margin of the appendage.

The costegites are rather small lanceolate structures which are

twisted and bent inwards; each carries about ten sete.

Pereiopod IIT (P\. XI. fig. 8).—Coxal plate unlike the preceding,

bilobed, with a few spines. ‘The second joint swollen, posterior

margin convex; the third joint short; the fourth and fifth

joints subequal. The fourth joint broadest at its distal extremity

and broader than the fifth. The sixth joint linear, narrow, and

longer than the fifth. Spines most numerous on the anterior

margin of appendage.

Pereiopod IV (Pl. XI. fig. 9): together with pereiopod V con-

siderably longer than the other pereiopods. In general pro-

portion like pereiopod IIT. The coxal plate is small. The second

joint is expanded... ;

Pereiopod V (Pl. XI. fig. 10).—Coxal plate small and shallow ;

the second joint much’ expanded, the posterior border notched

behind each small spine. Third joimt very short. Fourth, fifth,

and sixth joints linear. Numerous spines, especially on anterior

margin of appendage.

The Pleopods (Pl. XI. figs. 11 & 12).—The first the longest ;

the third by far the shortest; the second is intermediate in

size.

The Uropods (P|. XI. figs. 13 & 14).—The first the longest ;

the second a good deal shorter, and the third minute.

The first and second uropods are armed with stout bristles ; on

the distal jomts these bristles are curved at their distal ends.

The distal end of the third uropod is armed with one stout and

one very small bristle ; it consists of two joints. The basal joint

with one large and one smaller bristle.

The Telson (Pl. XI. fig. 15).—Simple, slightly divided at base.

It bears two bristles on each side.

112 MR. G. W. SMITH AND DR. E. H. J. SCHUSTER ON

EXPLANATION OF THE PLATES.

Talitriator eastwoode, gen. et sp. nov.

All the figures have been drawn with the aid of a Camera lucida.

PraTE X.

Fig. 1. Antennule.

Antenna.

Upper lip.

Lower lip.

. Maxillipeds.

. Gnathopod 1.

. Gnathopod 2.

EOE wwe

Prats XI.

Fig. 8. Pereiopod 3.

9. Branchial appendage of Pereiopod 4.

10. Pereiopod 5,

11. Pleopod 3.

12. Sketch of pleon, showing comparative size of appendages .

13. Dorsal aspect of posterior part of body.

14. Uropod 3.

15. Telson.

9. The Genus Hngeus, or the Land Crayfishes of Australia.

By G. W. Smirx, M.A., Fellow of New College,

Oxford, and KE. H. J. Scuusrrer, M.A., D.Sc., F.Z.S.,

Fellow of New College, Oxford.

[Received August 8, 1912: Read November 26, 1912. ]

(Plates XII.—XXV.*)

INDEX.

Pages

Geographical Distribution and Taxonomy ............ 112-116

HM Ghologiy) ssa hesevae sas Mester dowsacocdaesaescostee ce torte see LG = 18

Systematic: Five new species of Hngeus .,.......... 118-126

IntrRODUCTION.

The existence of burrowing forms of Australian Crayfish,

which live in underground tunnels excavated in damp soil, was

first made known by Erichson, who described two species from

Tasmania and placed them in a new genus, Yngeust. Besides

the two Tasmanian species, a very large collection of these

burrowing Parastacidee from Victoria has gradually accumulated

in the collection belonging to the Melbourne Museum, chiefly

through the activity of Messrs. Kershaw and Fulton, and the

present memoir is founded on this. large collection and also on

specimens which one of us obtained in ‘Tasmania in 1907-8.

Before proceeding to the description and classification of this

* For explanation of the Plates see pp. 126, 127.

y Archiv f. Naturg. vol. xii. 1846, p. 102.

eee See Sis. Ee cle)

6 3 Ai

GiLis OF ASTACOPSIS AND PARACHAIRAPS.

i, % Se eis) 1 sam.

i ~

GILLS OF ENG4US CUNICULARIUS.

(ane) an fey

. ais OMFS

B.S. Sie ee Sone

BINGAG US a OSs ORs

Po So WENS), EL SOV,

Bale & Danielsson, 1.19

BNGAIUS FOSSOR.

Pee So MNS ell SVL,

Bale & Danielss orn ditt

BENGHUS FOSSOR.

B.S. LOLs, IPL, Savane:

eAk

Bale & Damelsson, td

aia HINCZnUS) 1 OS SOR. 28-295. 19, AIDIEMONIS).

PA. S. UNS), Il. SCvAulae.

ea, Bale & Danielsson, Lt

28, ENGALUS NMEENUNIS. a7, 428, 1, WIC TORTEINSIS..

IF, 4. S5 ISIS. Pl, Sab<,

Bale & Deadclecon Lil

22) GINGA S: VAICWVORMEINSIUS, SO), Sl, Eh PNA Ocmecus)

D, Bo So tess talk Sex

Bale & Damelsson, itd

82. SS, INGZzaI0)s: WVICTORIENSIS. 34. H. HEMICIRRATULUS.

BP. Z.-S.-1Sis), i! year

Bale & Danielsson, Lt

St. is PIENLL OCs RCUS.

35. ENGAZAUS HEMICIRRATULUS.

Po Ao Ss ISIS), EI, Decay

Bale & Damelsson, Lt?

Se) ja), CUINWO WIL ASUS.

37,38. ENGAUS HEMICIRRATULUS.

Zs Sie LONG IP NOCD,

AVS

Bale & Danielsson, L,t&

ENGAUS CUNICUEARTUS:

IP, Zo S. USNS}, I SOD,

IUS, (WARRAGUL, GIPPSLAND).

HENCATUS CUNIC

Po Zo 3, US, IPL, DORN,

Bale & Danielsson, pte

ENGAIUS CUNICULARIUS.

THE LAND CRAYFISHES OF AUSTRALIA. ; 113

material, it is necessary to correct an error of interpretation,

published in the account which one of us gave of the Freshwater

Australian Crayfishes in the ‘ Proceedings of the Zoological

Society’ for March 1912 (p. 144). This account did not pretend to

deal with the Land Crayfishes of the genus Hngeus except in a

very general manner, but in treating of the geographical distri-

bution and relationship of the Australian Crayfishes as a whole,

certain statements were made as to the probable derivation of

Engeus from the freshwater form Paracheraps bicarinatus,

which a more detailed study of Hngeus has shown to be entirely

mistaken. Since the rectification of this error has an important

bearing on the geographical distribution of the group, and clears

up certain puzzling and unsatisfactory features of the problem,

we feel it right to give it some prominence, in order to prevent

the misconception spreading any further. In the memoir referred

to, it is pointed out that there exist in Australia two distinct

groups of Freshwater Crayfishes, the genus Astacopsis occupying

Victoria, Tasmania,and New South Wales, 7. e. the South-eastern

portion of Australia, and the genus Cheraps occupying Western

and Northern Australia and New Guinea, but nowhere mingling

in its range with Astacopsis. In addition to Cheraps and Asta-

copsis with their distinct and limited distributions, there is a

single form, the common Paracheraps bicarinutus, which is closely

allied to Cheraps in all its anatomical features and is evidently

a derivative from Cheraps. This form has spread from the West

right into and across the Central Australian deserts, and is now

found all over the Australian continent, mingling with Astacopsis

in Victoria and New South Wales. It has not, however, pene-

trated into Tasmania or New Guinea, which is strong evidence

in favour of its being a comparatively modern derivative from

Cheraps.

As P. bicarinatus is accustomed to live in small water-holes in

the desert, it has taken on a wandering and burrowing habit,

and is frequently found walking about in fields in search “of some

moist ditch or water-hole.

Now, it seemed probable that the genus Hngcus, containing the

burrowing land forms, and confined in its distribution to Victoria

and Tasmania, had been derived from Paracheraps bicarinatus,

through an intensification of the habit of leaving the water and

burrowing in damp soil. This idea was further confirmed by the

curiously close resemblance which some of the Hngwus bear to

Paracheraps bicarinatus in external appearance. It was there-

fore too confidently stated in the memoir referred to that Hngeus

was probably a derivative of Paracheraps. Nevertheless, there

was a very puzzling feature pointed out, supposing this derivation

to be trye, viz., that whereas Hngeus is represented in Tasmania

by two species, the supposed parent form, Paracheraps bicarinatus,

is entirely absent from that island, so that it was necessary to

introduce one of two rather improbable hypotheses, either that

P. bicarinatus once existed in Tasmania and is now extinct, or

Proc. Zoou, Soc.—1913, No. VIII. 8

114 MR. G. W. SMI'YH AND DR. E. H. J. SCHUSTER ON

else that the two species of Hngwus have been somehow trans-

ported across Bass’ Straits by accidental means.

Now, it transpires as the result of a detailed examination of the

various species of Hngeus, especially in respect to the structure of

their gills, that the derivation of Hngceus from Paracheraps is

entirely false, and that the superficial resemblance of these forms

to one another is due to convergence. On the other hand, it

is abundantly evident from the gill characters that Hngcus is

a derivative of Astacopsis, and has nothing to do with either

Cheraps or Paracheraps. The reasons upon which this conclusion

is based are as follows. In Astacopsis the podobranchia do not

possess a broad ala or wing-like expansion of the stem of the gill ;

the ala is, on the contrary, reduced to a mere rudiment, as shown

in the transverse section (Pl. XII. fig. 1). Attached to this

rudimentary ala of the podobranchs are a few gill-filaments,

which are furnished at their tips with characteristically shaped

hooks (fig. 2). All the other gills in Astacopsis, exclusive of the

podobranchs, i. ¢. the arthrobranchs and pleurobranchs, have their

filaments entirely free from terminal hooks. The peculiar hooked

setee present on the bases of the podobranchs in Astacopsis also

have a constant and characteristic shape, the terminal hooks

being not sharply recurved (fig. 3). Now all the above characters

are absolutely constant for the various species of Astacopsis. In

Cheraps and Paracheraps, on the other hand, we have a totally

different series of gill characters which are just as constant and

characteristic for all the species of these two closely related

genera. In these forms the podobranchia possess a very broad ala

(fig. 4) which is furnished with numerous filaments, whose hooks

have a characteristic sickle shape (fig. 5). In all the other gills,

besides the podobranchs, numerous filaments are provided with

these terminal hooks. Finally, the hooked setz (fig. 6) on the

podobranchs have an entirely different shape compared with those

of Astacopsis, being sharply recurved at the ends.

Now, the distinctive character and absolute constancy of these

gill-structures force us to attach especial taxonomic importance

to them, far more importance than external appearance or other

characters which fluctuate from species to species, so that an

examination of the gills of Hngaus should give us the key to its

relationship with the other genera. The gills of Engzeus agree in

all the above particulars with those of Astacopsis, and differ entirely

from those of either Cheraps or Pavacheraps (Pl. XIII. figs. 7-10).

Thus the podobranchs possess a rudimentary ala (figs. 7, 8) which

carries a few filaments, and these filaments terminate in hooks

(fig. 9) shaped like those of Astacopsis. The gills other than

the podobranchs have their filaments unprovided with hooks;

the hooked sete (fig. 10) on the podobranchs are not sharply

recurved at the end, but are shaped as in Astacopsis.

Now, apart from the gills, Hngeus differs so widely from both

Astacopsis, Cheraps, and Paracheraps in the characters of its

bodily structure, appendages, etc., that these no longer serve as a

THE LAND CRAYFISHES OF AUSTRALIA. 115

guide, and the resemblance to Puracheraps, which proved at first

deceptive, consists merely in the absence of spines and ridges on

the body and the hairiness of the mouth appendages, which are

evidently due to convergence following on the similar mode of

life.

As a matter of fact, in a certain number of anatomical features,

besides the gills, Hngeus agrees rather with Astacopsis than with

the other two genera: thus the penultimate segment of the second

maxillipede projects nearly as far forward as the terminal segment,

and the vas deferens is situated on a short simple papilla.

We therefore have no hesitation in claiming that Hngeus is a

derivative of Astacopsis or of a close ancestral form of Astacopsis,

and is fundamentally distinct from the western Chewraps and its

widely distributed derivative P. bicarinatus.

It may be pointed out that this conclusion is far more intelli-

gible, on general grounds, than the original derivation of Hngeus

from P. bicarinatus, and it clears up all the puzzling features in

the geographical distribution of these forms. We can now clearly

explain why Hngeus occurs as two species in Tasmania, although

P. bicarinatus is absent, because Astacopsis occurs in Tasmania.

It was also rather unaccountable that P. bicar inatus, being a com-

paratively recent derivation from Cheraps, which had not been

able to penetrate into New Guinea or ‘Tasmania, should have been

able to give rise to several widely divergent species of Hngeus

distributed through Victoria and Tasmania. We can, however,

easily see that if Hngeuws is a derivative from the much more

ancient Astacopsis, that there has been ample time for its diver-

gence into several species, and its distribution in Tasmania and ~

Victoria alongside the parent dAstacopsis, though occupying a

totally different station, is perfectly intelligible.

To sum up the relationships of the Australian Crayfishes

as a whole. The western and northern genus Cheraps and

the south-eastern genus Astacopsis have been isolated from

one another completely since a very ancient date, and at the

present time they nowhere intermingle. Cheraps sent a northern

straggler into New Guinea and the Aru Islands at a time when

these islands were connected with the mainland (C. quadricart-

natus), and Astacopsis has a typical representative in Tasmania,

so that this genus in very much its present condition must have

been present in Southern Australia at any rate when Tasmania

was joined to the mainland. During this period, namely, when

Tasmania was joined to the mainland, Astacopsis gave rise to the

burrowing Land Crayfishes, Hngeus, which are now represented

by several species in Victoria and by two species in Tasmania.

Subsequently to the separation of New Guinea and Tasmania

from Australia, the western Chwraps gave rise to an offshoot,

Paracheraps bicarinatus, which, forsaking the rivers as a necessary

habitat, took to its wandering, pond and w. ater-hole frequenting

mode of life, and was thus enabled to spread across the desert

regions and invade the territory of Astacopsis in the south

116 MR. G. W. SMITH AND DR. E. H. J. SCHUSTER ON

and east. Thus P. bicarinatus:is now found everywhere on the

Australian continent, but it has not been able to reach New

i » Tasmania, si ries a ’ from the

Guinea or Tasmania, since these countries were cut oft fro

mainland when P. bicarinatus arrived.

The relationships of the Australian Parastacide should there-

fore be expressed as follows, in place of the diagram given in the

former memoir :—

Cheraps. | Paracheraps: Hnge@eus. Astacopsis.

va \ vy

\ /

= :

“, =

% SY

“y \

GS &

\ s

Common ancestor,

2. Structure, Habits, and Interrelationships of the

Species of Engeeus.

The several species of Hngcus are characterised by the great

depth of the carapace, measured dorso-ventrally, and by its arched

roof-shaped form. The eyes are small, and the abdomen tends to ©

be reduced in size. We also note a greater hairiness of the mouth-

parts and neighbouring regions, a feature probably correlated

with the necessity of filtering the water in their burrows, which

is usually very muddy. The gills, especially the arthrobranchs,

tend to be reduced in size, and though the gill formula may be

the same as in the other Australian Parastacide, yet in certain

forms the last pleurobranch is altogether suppressed. This entire

suppression of a gill, which is unknown in any of the other

Australian Crayfishes, is only one example of the very striking

morphological changes which the species of Hngwus may exhibit ;

and it may be truly said that the species of this genus are often

separated from one another by greater differences than those

which distinguish the genera of the Parastacidee from one another.

THE LAND CRAYFISHES OF AUSTRALIA. 117

Thus, in one series of forms, /. fossor. affiiis, victoriensis, phyllo-

cercus, fultoni, and hemicirratulus, the exopodite of the third

maxillipede may be rudimentary or entirely suppressed; in

another series the outer flagellum of the first antenna is absent

(Z, hemicirratulus), while the antennary scale and the uropods

undergo striking changes in certain species,

It is difficult to find the reason for these marked aberrations of

structure, but it may be pointed out that all of them are in the

nature of losses or suppressions, e. g. the loss of the pleurobranch,

the loss of the exopodite of the third maxillipede, the loss of a

flagellum on the antennule, the reduction of the antennary scale,

and the abortion of the eyes; and it may be plausibly argued that

the underground, burrowing habit which removes the animal

from active competition with other water forms has permitted

degenerative changes which have no special adaptive meanings.

Another interesting point is the extraordinary variability of the

large chele in these forms, and their frequent asymmetry. It

appears to us extremely doubtful whether these variations

indicate specific or only individual differences, since hardly two

forms coming from the same locality and identical in other

respects are ever quite alike in regard to their chele, and very

often they differ fundamentally in this respect, independently of

sex and size.

Unfortunately, we do not know very much about the habits of

these burrowing crayfish, and the only personal observations were

made by one of us in Tasmania on the smaller Tasmanian Lngeus,

EH. fossor.

This species is widely distributed in Tasmania, being found on

the southern mountain ranges, on the north coast, and in the

forests on the west coast. The burrows of the animal could be

recognised as little round holes in the damp ground, sometimes

near the banks of a stream or water-course, but very frequently

far removed from any water in the middle of the forest in some

damp situation. The burrow descends vertically into the ground,

or if it is in a bank it is frequently horizontal, and after passing

inwards for a foot or two it ends in a circular chamber which is

always full of water. Hngeus fossor, when freshly taken from its

burrow, has a rather soft whitish skin with brilliant blue patches

and an occasional tinge of red. In one place near the Magnet

Mine on the west coast of Tasmania, the banks of an artificial

water-course which was used in the mining operations were com-

pletely riddled by the burrows of Hngeus fossor, and the foreman

of the mine told me that these creatures were a source of continual

damage and danger in works of this kind.

It is stated in Erichson’s original memoir, on the authority of

Herr Schayer, who collected the specimens, that Hngeus if kept in

water soon dies. In view of the fact that there is always water

at the bottom of the burrows, and also from what I have heard

from people in Tasmania who have kept the animals in captivity,

this statement may be received with some scepticism,

118 MR. G. W. SMITH AND DR. E. H. J. SCHUSTER ON

Since the animals are never seen out of their burrows in the

day-time, very little is known as to their food or the means they

employ to obtain it, but it is probable that they are mainly

carnivorous in diet, as the remains of earthworms, insect larve,

and probably land Crustacea have been found in their stomachs.

As has been pointed out elsewhere (‘A Naturalist in Tasmania’),

the evergreen beech forests in Western Tasmania support a very

rich terrestrial fauna of land Amphipods (7'alitrus) which swarm

under the fallen beech leaves and timber, and numerous Myriopods

and insect larvee occur as well, affording abundant food in exactly

the situations which Hngceus chooses for its burrows.

The young are brought into the world and tended by the

female parent in the same way as in the ordinary crayfish, and I

have obtained females from their burrows with their young ones

still attached to the swimmerets under the abdomen. At this

stage the young ones have all the characteristic features of the

adult, showing the same vaulted carapace and reduced abdomen.

The interrelationships of the various species of Hngcws may be

gathered from the key for the determination of the species given

on p. 119.

The least modified and specialised species is #. cwnicularius from

Tasmania, Gippsland, and Victoria. This species has the general

form of the thorax and abdomen in a less aberrant condition than

the other species, and it is also more normal in its other characters,

possessing the last pleurobranch, the exopodite on the third maxilli-

pede, and the two flagella on the first antenna ina fully developed

condition. WH. fultoni from Victoria is closely related to this

species, but the exopodite of the third maxillipede is rudimentary.

The Tasmanian #. fossor and the Victorian species 1. affinis,

victoriensis, and phyllocercus are all closely related, and exhibit to

the full the peculiarly roof-shaped thorax and reduced abdomen ;

while 2. hemicirratulus is the most highly specialised, having Jost

not only the exopodite of the third maxillipede but also the

posterior pleurobranch and the inner flagellum of the first

antenna.

Genus ENG US.

Erichson, Archiv f, Naturg. vol. xii. p. 102 (1846); Huxley,

1B, Ze Se Wits jos (OY)

The gill-formula is the same as in Astacopsis, Cheraps, and

Paracheraps, except that the last pleurobranch may be entirely

absent. The posterior arthrobranchs are reduced in size.

The ala of the podobranchs (Pl. XIII. figs. 7 & 8 al.) is small

and inconspicuous as in Astacopsis, and it carries a few filaments

with terminal hooks which are shaped as in Astacopsis.

None of the other gill-filaments carry terminal hooks.

The hooked sete on coxopodites and podobranchs are not

sharply recurved, but resemble those of Astacopsis.

The mandibles (Pl. XV. figs. 16 & 17) have two prominent

THE LAND CRAYFISHES OF AUSTRALIA. 119

median teeth and one smaller tooth in front, and a row of smaller

serrations behind,

The first maxille (Pl. XVI. figs. 18 & 19) have the endopodite

without any trace of a flagellum.

The second maxillipede has the penultimate segment broad,

projecting nearly as far as the terminal segment (fig. 21).

There is an entire absence of filaments from the epipodite of

the first maxillipede,

The third maxillipede may have the exopodite normal, reduced,

or absent. There is a great development of filtering apparatus

of bristles and hairs on this and on all the mouth-parts.

The great chele show an immense range of variation, being

symmetrical or asymmetrical, serrated or smooth, hay or

comparatively hairless,

The vas deferens is situated on a simple short papilla.

The rostrum is reduced, and its lateral keels are either smooth

or feebly tuberculated, There are no keels on the carapace, nor

are there any spines on the carapace or abdomen, The abdomen

is reduced, sometimes markedly so, and it may be hairy,

The eyes are reduced in size.

The carapace is very deep (Pl. XIV. fig. 12) measured in a

dorso-ventral direction, and the distance from the tip of the

rostrum to the cervical groove is always longer than that from

the cervical groove to the posterior border of the carapace.

Key to the Species of the Genus Kngeus,

(1) Last pleurobranch present, two flagella on Ist antenna.

(A) Exopodite of 3rd maxillipede absent or rudimentary.

Autennal seale rounded at end without spine ................ EH. fossor,

Antennal scale pointed, last pleurobranch large ............ EL. affinis.

Antennal scale pointed, last pleurobranch reduced ......... H. vietoriensis.

Uropods produced into point:d apices ...................... H. phyllocercus.

Rostrum straight and large with prominent keels ......... H. fultoni,

(B) Exopodite of 3rd maxillipede well developed .....,...... H. cunicularius.

(2) Last pleurobranch and exopodite of 3rd maxillipede absent

and only one flagellum on first antenna..................6. EH, hemicirratulus,

Enemas Fossor Erichson, (Pls, XIV—XVII, figs, 11-22.)

Astacus (Engeus) fossor Evichson, Arch. f. Naturg, vol, xi.

p. 102 (1846).

Astacus fossor Von Martens, Monatsber, Akad, Wiss. Berlin,

1868, p, 618,

Engeus fossor Haswell, Cat, Australian Museum, Stalk- and

Sessile-eyed Crustacea, p, 178 (1882),

The posterior pleurobranch is present, and is larger than the

others. The anterior pleurobranch is very small,

The posterior arthrobranchs are all very small, consisting of a

stalk bearing not more than four or five filaments.

The exopodite of the third maxillipede is reduced to a very

small tubercle (Pl. XVII. fig. 22).

120 MR. G. W. SMITH AND DR. E. H. J. SCHUSTER ON

The antennal scale is rounded at the end and carries no terminal

spine (Pl. XV. figs. 14 & 15).

The inner flagellum of the first antenna is slenderer than the

outer one and about two-thirds its length.

Tubercles on propodite and ischiopodite of chela very incon-

spicuous,

The great chela always has a row of tubercles on the upper

border of the propodite, the lower border being generally smooth

and rounded. There is one prominent tooth on the inner surface

of the dactylopodite, while there are three teeth on the lower part

of the pincer. In some specimens the chele are equal in size and

shape, in others one chela is more slender and hairy, and with the

pincer more delicate and elongated.

In the female there is a pair of sperm-receptacles with con-

spicuous openings on penultimate segment (Pl. XV. fig. 13).

The eyes are small, and the rostrum is short with rather

prominent keels. Contour of forehead shown in fig. 12 (Pl. XIV.).

The thorax is laterally compressed and highly arched, It is

fairly free from hairs, but the abdomen is rather hairy.

The tail-fan is rounded without conspicuous spines or ridges.

Colour. Ground-colour ivory with blue and red blotches

ivregularly disposed.

Length 38 mm.

Locality. Magnet Mine, West Coast of Tasmania. Also

reported to occur on the southern ranges. Confined to Tasmania.

Local Variety. Some specimens obtained from burrows near

Muddy Creek, Bridport, on the north coast of Tasmania, while

agreeing essentially with the above description, show certain

variations. ‘The whole body and limbs are more hairy, and this

is especially marked on the chele. The propodites of the great

chele are slightly tuberculated on their lower as well as on the

upper border, and the chele tend to be slightly more elongated

than in the type specimens. The keels on the rostrum are

slightly more pronounced, and the eyes are a little larger.

ENGA&US AFFINIS, sp.n. (Pls. XVII. & XVIII. figs. 23-26.)

All the.pleurobranchs are of approximately the same size, the

posterior pleurobranch being well developed.

The exopodite of the third maxillipede is absent.

The antennal scale ends in a well-developed terminal spine and

an inner lobe which is not markedly produced (Pl. XVII. fig. 25).

First antenna is formed as in Z. fossor.

_ There are two conspicuous rows of tubercles on the inner

surface of the carpopodite of the great chela, anda row of marked

tubercles on the meropodite. There is a row of tubercles on the

upper border of the propodite, the lower border being smooth

and rounded.

There is, as usual, great variability in the size and shape of the

chele, in some specimens the two chele being similar and equal

in size, while in others one chela, either the right or left, is

THE LAND CRAYFISHES OF AUSTRALIA, 121

greatly enlarged. It is a general rule that the enlarged chela

(fig. 24) in this species has a broad and short propodite if com-

pared with that of the next species, H. victoriensis (Pl. XX.

fig. 33).

There are no sperm-receptacles with conspicuous openings in

the female.

The eyes are rather larger than in £, fossor, and the rostrum

is longer and ends in an upwardly directed spine, The contour

of the forehead (Pl. XVIII. fig. 26) is much less steep than in

EH. fossor (P]. XIV. fig. 12).

Thorax, abdomen, and tail-fan much as in LZ. fossor.

Length. Specimen figured from tip of rostrum to end of telson,

60 mm.

Localities :—

1. Several specimens from’ Warburton, Victoria, 15. x1, 05,

One specimen with equal-sized claws figured (fig. 23). Four

other specimens similar to this one in the matter of shape of

claws, though varying greatly in hairiness. One specimen with

one very large and stout chela (fig. 24), not hairy; the other

chela of this specimen was unfortunately missing.

2. Two specimens from the Upper Yarra, collected by Mr,

Williams in 1869 and 1871. In both the chelze are equal in size

and rather slenderly built,

3. Two specimens from the top of Black Spur, Fernshaw, 1880,

In both these specimens the right chela is much larger and more

massive than the left, which is elongated and narrow. One other °

small specimen from Fernshaw has the chele similar and equal.

Another specimen, simply labelled ‘* Victoria,’ resembles the

last-named.

4, One very large specimen from Healsville, Victoria, 5. vi. 82,

has the right chela enlarged and massive, the left chela narrow

and elongated.

ENGUS VICTORIENSIS, sp.n. (Pls, XVIII.-XX. figs. 27-29,

32 & 33.)

The penultimate pleurobranch is more than double the size of

the last pleurobranch, the latter being greatly reduced in size.

The exopodite of the third maxillipede is absent.

The antennal scale ends in a well-developed terminal spine

and an inner lobe which is not markedly produced.

First antenna (PI. XVIII. fig. 28) as in LH. fossor.

The chele resemble those of H#. affinis, save that when one is

enlarged the propodite is not so broad and short, but is rather

more “elongate in shape than in £#. affinis. There is, aS usual,

great variability in size, shape, and symmetry of the claws.

The rostrum resembles essentially that of 4. affinis, but the

contour of the forehead (fig. 27) is a little steeper.

The abdomen of both sexes is rather broader than in #. affinis,

Length of specimen figured from tip of rostrum to end of telson,

65 mm.

122 MR. G. W. SMITH AND DR. E. H. J. SCHUSTER ON

Localities :—

1. One specimen from the top of the Dandenong Ranges,

Victoria (Kershaw, 1. 72) (Pl. XIX. fig. 29). Left chela enlarged,

right slender.

2. One specimen, 8. Gippsland, July 1891, exactly similar to

above.

3. One specimen, Boxhill, near Melbourne, has right chela

enlarged, left slender.

4. Two small specimens, Emerald, Victoria (2. Jarvis, viii. 04).

Left chela enlarged, right slender, the other with chele subequal.

5. Several (small) from Fern Tree Gully, Victoria. Some with

right chela enlarged, some with left.

6. One specimen from Croydon, Victoria (/’. P. Spry, 1. xi. 04).

Right chela enlarged.

7. One specimen from Ringwood, Victoria (#. H. Hennell,

xi, 90). Left chela enlarged.

8. Several specimens, simply labelled “ Victoria.” Some with

right chela, some with left enlarged; a few with chele subequal

or equal.

9. One very large specimen from the Launching Place, Victoria

(J. Coghill, 15.1. 07), with left chela enlarged.

Remarks on the above two species, HK. affinis and victoriensis.

These two species, which are very widely distributed in

- Victoria, are clearly distinguishable from the Tasmanian Z. Jossor

by a dunes of chansons, and from the succeeding species,

Li. phyllocercus, by the remarkable shape of the uropods in the

latter form, But the characters distinguishing #. affinis and

victoriensis are very slight, and it may be possible at some time

to merge them together in one species, victoriensis. The only

really satisfactory character is the size of the last pleurobranch,

which is reduced in victoriensis and well developed in affinis.

The only other character which is of use in separating the two

forms is the shape of the enlarged chela, which is shorter and

broader in affinis than in victoriensis; but since the enlarged

chela is not always developed, this character is an unsatisfactory

one.

ENGZUS PHYLLOCERCUS, sp. n. (Pls. XIX. & XXI. figs. 30, 31,

& 36.)

All the pleurobranchs are of approximately equal size, the last

one not being reduced.

The exopodite of the third maxillipede is reduced, but is

clearly to be seen, and it is tipped with several plumose hairs

(Pl. XX. fig. 36).

The antennal scale ends in a spine which is less elongated than

in the preceding two species.

The first antenna is similar to that of preceding species.

The great chela has both the upper and lower borders of the

THE LAND CRAYFISHES OF AUSTRALIA. 128}

propodite serrated. There is a single row of tubercles on the

upper and inner border of the carpopodite, and a row of tubercles

on the upper border of the meropodite. The shape of the chele

is somewhat elongated, and one is generally enlarged, the other

remaining slender.

The rostrum ends ina rather blunt upeurved spine, which is

shorter and blunter than in the two preceding species.

The uropods exhibit a peculiar and highly characteristic modifi-

cation, in that both endopodite and exopodite are produced

distally into pointed apices, giving the tail-fan a leaf-like appear-

ance (Pl. XIX. fig. 31).

Length of specimen figured from tip of rostrum to end of telson,

59 mm.

Localities. Narracan River, Thorpdale, Trafalgar. All these

are Gippsland localities to which the species appears to be

confined.

Variety. The rudimentary exopodite of third maxillipede varies

in size in the different specimens, and in one specimen from a

small stream near Thorpdale, Gippsland, collected by Mr, Kershaw

in March 1890, the exopodite is reduced to a small papilla.

ENGUS HEMICIRRATULUS, sp.n. (Pls. XX.-XXII. figs. 34, 35,

37, 38.)

The posterior pleurobranch is entirely absent,

The exopodite of the third maxillipede is absent.

The antennal scale ends in a prolonged curved and blunt spine,

and the inner lobe is small (Pl. XXII. fig. 37).

The first antenna has only one flagellum, the inner flagellum

being entirely absent (fig. 38).

The upper and under and posterior borders of the propodite of

the chela are studded with marked tubercles, as are also the

borders of the carpopodite, so that the chela as a whole has a more

tubercular appearance than in any of the other species. One

chela is generally more stoutly developed than the other, some-

times on the right side and sometimes on the left. The pencils

of hairs on the chelze are more conspicuous than in preceding

species.

There are no sperm-receptacles with conspicuous openings in

the female.

The eyes are not markedly reduced. The rostrum has a con-

spicuous rounded humpat its base, but is flat and keelless distally

and ends in a blunt spine. Just proximal to the point of the

spine is a conspicuous bunch of hairs. The contour of the fore-

head is not at all steep.

The thorax is vaulted and deep as in the preceding species ;

the abdomen is reduced and hairy. The tail-fan is normal in

shape, but the pencils of hairs upon it are particularly conspicuous,

Colour. (From coloured figures by McCoy.) Carapace dull

purple. Chele reddish. Legs and abdomen brownish grey.

Length 51 mm.

124 MR. G. W. SMITH AND DR. E. H. J. SCHUSTER ON

Localities :—

1. Three female specimens and one male from a hill near Thorp-

dale, Gippsland (W. Kershaw, 111. 90), with left chela larger than

right. These range in size from 70 mm. to 31 mm. Four small

specimens, ranging in size from 30 mm. to 18 mm., had the chelz

equal or subequal.

2. Several specimens from Warragul, Gippsland. A large

female, 80 mm. in length, with right chela enlarged, 3. ix. 92;

and several specimens, some with right chela enlarged, others

with left, and some with equal chele (Prof. B. Spencer).

3. A large male, measuring 65 mm., with equal chele, froin

Moyarra, near Oultrim, 8. Gippsland (Avtson, 1905).

4, A male, measuring 50 mm., with right chela enlarged, from

Kongwak, near Jumbanna, 8, Gippsland (Avtson, 1902).

Remarks. 'Vhis species was partly described and figured by

McCoy in MS. as Hemicirratulus hystrix. Since it does not fit

in with the scheme of this memoir to place this species in a

separate genus, we have retained McCoy’s MS. generic name,

hemicirratulus, as the specific name. ‘The species is a constant

and easily recognised one; the chele, as usual, are variable in

development, and the rostrum differs somewhat in the different

specimens. ‘In the specimens from 8. Gippsland the rounded

hump at the base of the rostrum extends rather further along

the rostrum than in the specimens from other parts of Gippsland.

ENG &Us CUNICULARIUS Hrichson, (Pls. XXII.-X XV. figs. 39-47.)

Erichson, Archiv f, Naturg. vol, xii, p, 102 (1846); Von

Martens, Monatsber, Akad. Wiss, Berlin, 1868, p,619; Haswell, Cat.

Australian Mus., Stalk- and Sessile-eyed Crustacea, p. 179 (1882).

The posterior pleurobranch is present, and is the largest of all

four pleurobranchs, which increase in size from before backwards.

The third maxillipede has a well-developed exopodite (fig. 46).

The scale of the second antenna has a short terminal spine and

a well-developed inner lobe (fig. 47).

The two flagella of the first antenna are nearly equal in length,

the outer flagellum being slightly longer than the inner.

The upper or inner border of the propodite of the chela has a row

of five tubercles ; the lower or outer border is smooth. There is

a row of feeble serrations on the inner border of the carpopodite

and on the upper border of the meropodite, but the chela on the

whole is very feebly tuberculated and comparatively free from

hairs, ‘There is. however, great variability in the chele.

There are no conspicuous sperm-receptacles in the female.

The eyes are large and comparatively unreduced.

The rostrum is straight, well-developed, and ends in a blunt

spine. There are two highly-developed tumid keels on its sides,

which are continued a little way back on to the carapace.

The thorax is less highly vaulted and more normal in shape

than in the foregoing species.

THE LAND CRAYFISHES OF AUSTRALIA. 125

The abdomen is very large and unreduced ; the telson is broad

and the tail-fan rounded, well developed, and normal in shape.

The abdomen and tail-fan are not at all conspicuously hairy.

The colour varies from reddish brown to olivaceous grey, while

parts of the chela and thorax may be picked out in bright blue

and red.

Localities :—

1. Tasmania. In one specimen, a male, both chele are large

and equal in size and shape. There is a peculiar compound tooth

on each jaw of the pincer (Pl. XXII. fig. 39). This specimen comes

from Glenore, near Hageley, Tasmania (Lartholomew, 29. v.89).

In one male specimen from Launceston, Tasmania (Bartholomew,

1890), the left chela is enlarged and resembles that of the above

specimen; the right chela is toothless, small, and slender, A

similar specimen, but a female, comes from Mundan Farm, Long-

ford, Tasmania (Bartholomew, 1889).

2. Warragul, Gippsland. Three male specimens (Kershaw,

1887, 1888), two with left chela enlarged, one with right. There

is a prominent peg-like tooth on the dactylopodite part of the

enlarged pincer; the compound tooth on fixed jaw of pincer is

reduced (Pls. XXII., XXIIT. figs. 41 & 42).

3. Near Lake’s Entrance, Gippsland. One female specimen

(Kershaw, 1887) with two similar chele without teeth (fig. 40).

‘This type of chela is intermediate in character between the large

and small chela of the specimens from Warragul (figs. 41 & 42).

4, Derby River, Wilson’s Promontory (Aershaw, 1905). Three

specimens, all with similar and equal chele. Teeth are present

in the pincer, but are reduced in size. * The chele are rather

elongated in shape.

A note appended to these specimens states that they build

conical mud towers, about 8 to 10 inches high, on the flats.

5. From banks of Fraser Creek, Oberon Bay, Wilson’s Promon-

tory (Kershaw, 1905). Similar to the last mentioned, but pincer

tends to be less elongated.

6. Male specimen from Croydon, Victoria (/ulton, 1907), with

left chela enlarged. Interior of pincer is hairy and without any

enlarged teeth.

7. Female specimen from Fern Tree Gully, with left chela

enlarged and small teeth in pincer.

8. Numerous specimens from Croydon, Victoria, and unnamed

Victorian localities, characterised by the presence of a thick pad

of fine downy hairs on the inner surface of the chela (Pl. XXV.

fig. 45), and also a thick covering of downy hairs on the third

maxillipedes. This constant character is possibly of specific value,

and it might be advisable to separate these forms as a distinct

species; but they do not apparently show any other differences to

distinguish them from the typical #. cwnieularius. It is a re-

markable fact that an exactly similar downy pad occurs in some

species of Cheraps.

126 MR. G. W. SMITH AND DR. E. H. J. SCHUSTER ON

ENGZUS FULTONI, sp. n.

This species agrees with H. cunicularius in all its characters,

save that the exopodite of the third maxillipede, instead of being

large and normally developed, is reduced to a small papilla as in

E. phyllocercus. The teeth in the pincer of the chela are not

fused to form a compound tooth, but there are several, about

three, small teeth on the lower jaw of the pincer and two on

the upper. ‘The lower border of the propodite of the chela has a

distinct serrated keel.

Localities :—

1. Two specimens from Fern Tree Gully, Victoria, one being a

female measuring 53 mm., with both chele large and equally

developed ; the other a small female measuring 35 mm., with the

left chela enlarged and the right slender.

2. One male from Cape Otway Forest, Victoria (collected by

W. Fulton, 28. v.07), with both chele similar, fairly stoutly built,

and markedly hairy.

EXPLANATION OF THE PLATES.

PuateE XII.

Fig. 1. Semidiagrammatic cross-section of podobranch of Astacopsis. al. = ala.

Only a few of the gill-filaments are shown.

. Terminal hook from gill-filament on ala, taken from the podobranch of

Astacopsis.

. Ends of hooked sete from podobranch of Astacopsis.

. Semidiagrammatic section of podobranch of chela in Paracheraps.

Terminal hook from gill-filament of Paracheraps.

. End of hooked seta from podobranch of Paracheraps.

Aap xr

Puate XIII.

7. Podobranch of great chela from Hngeus cunicularius. Az.

8. Semidiagrammatic transverse section of do. X12. al. = ala.

Fig.

. Tips of hooked gill-filaments from ala of do. Xx 280.

. Tips of hooked sete from podobranch of do. 260.

Puate XIV.

Fig. 11. Engeus fossor. From dorsal surface. X13.

12. 3 45 Side view, with limbs removed. 13.

Pate XV.

Fig.13. Engeus fossor. Sperm-receptacles on penultimate segment of thorax in

the female. X23.

14, e a Basal portion of right antenna from above. X63.

15. i x Scale of antenna. X12.

16. 3s 4 Left mandible from above. X63.

17. ae , Do. trom below.

Puate XVI.

Fig. 18. Engeus fossor. First maxilla (left) from above. 63.

19. i a End of exopodite of first maxilla. 65.

20. i *s Second maxilla (left) from above. 6}.

21. 3 53 Second maxillipede (left) from below. 63.

0g”

24.

20.

ig. 26

27.

28.

BB]

Phy

THE LAND CRAYFISHES OF AUSTRALIA. 127

.22. Engeus fossor.

23.

Prate XVII.

Basal portion of third maxillipede (left) from above. X64.

affinis, 2. Dorsal view. X13.

. Engeus affinis.

bb)

Great chela. X24.

Scale of antenna (left). 64.

Prate XVIII.

Contour of forehead. X24.

victoriensis. Contour of forehead. X24.

bb)

First antenna (left). 6.

PuatE XIX.

Fig.29. Engeus victoriensis, 2. Dorsal view. X1}.

phyllocercus, 2. Dorsal view. X13.

Fig.

Vig.

Fig.

30.

31.

3”

Telson and uropods. X2.

PuatTeE XX.

32. Engeus victoriensis. Telson and uropods. XQ.

Great chela. X23.

hemicirratulus. “ Face.”

PLaTE XXI.

. Engeus hemicirratulus. Dorsal view. X14.

phyllocercus, Third maxillipede (left) from below.

Prate XXII.

.37. Engeus hemicirratulus. Base of right first antenna from below. X63.

First antenna. X63.

bb)

cunicularius. Chela of male from Glenore, near Hageley, Tas-

mania (fig. 43). X24.

Right chela of female found near Lake’s Entrance,

Gippsland. X2.

Right chela of male specimen from Warragul, Gipps-

land. 2.

Prate XXIII.

42. Engeus cunicilarius. Left chela of male specimen frotn Warragul, Gipps-

43.

”%

land. Companion to fig. 41.

6. From Glenore, near Hageley, Tasmania: 14.

PuatEe XXIV.

44. Engeus cunicularius, §. From Warragul, Gippsland. X14.

bP)

»”

PuatTE XXV.

. Engeus cunicularius. Left chela of specimen from Croydon, Victoria:

46.

47.

Third maxillipede (right) from above. X44.

Scale of left antenna. X62.

128 MR. B. F, CUMMINGS ON THE

10. On some Points in the Anatomy of the Mouth-parts

of the Mallophaga. By Bruce F. Cummrnes*.

[ Received October 17, 1912: Read November 26, 1912.]

(Text-figures 24-32.)

INDEX.

Structure or Morphology : Page

Palpi of theplisehnocerayewa-pa-esestes tase eS

. BEAD IV Geral Sasych cae tase cee cen ee eer 128

The esophageal sclerite and glands......... ........... 129

130

Varieties of sclerite and glands.....................2...2.

Distribution of sclerite and glands ...................... 133

Descriptions of special cases .............:0.: cee. 183

The maxillary forks 139

Taxonomy :

Conclusion BOE c at see Man eee aaeeeode 1D

The Mallophaga are for the most part such small parasites

that the dissection of the mandibulate mouth is often a matter of

considerable difticulty. It seems scarcely surprising, therefore,

that a good deal of misunderstanding has arisen in regard to

the mouth-parts, and in the literature which deals with this

subject a variety of opinion will be found on such questions as

the palpi, the cesophageal sclerite, and so on.

The Palpi of the Ischnocera.

In a paper in the P.Z.S. 1909 (1), appear figures and de-

scriptions of the Ischnoceran species Goniodes tetraonis Denny,

from the Grouse, in which a pair of very minute one-jointed

appendages is shown to exist on the labium below the para-

zlosse. In G. falcicornis N., I can find no trace of these minute

appendages, which perhaps represent the palpi of the second

maxilla. In the same place, the second maxille of G. tetraonis

are described as rounded with certain sete or hairs, whence it

appears that in this species they are less degenerate than in the

other Ischnocera so far examined, where the second maxille are

usually obsolete, flat lobes without palpi, thinly chitinized (or not

chitinized at all), and situated within the mouth-cavity behind

the mandibles. From the figures of both G. tetraonis and

another Nirmus cameratus N. they are omitted, and instead, the

paraglossee of the labium are labelled first maxille and the

minute appendages second maxille.

The Palpi of the Amblycera.

The other suborder, the Amblycera, is distinguished by the

presence of a pair of jointed palpi absent from the Ischnocera.

The view of Nitzsch, who first considered them in 1818 (2), may

at length be considered as sound, namely, that they belong to the

first maxille and not to the labium. Grosse (1885) (8) assigned

* Communicated by the SECRETARY and published by permission of the Trustees

of the British Museum.

MOUTH-PARTS OF THE MALLOPHAGA, 129

them to the labium. Snodgrass (1896) (4) followed suit, but

in 1905 (5) retracted in favour of Nitzsch, as the result of

examining preparations of Ancistrona gigas P. and Leemobo-

thrium gypsis Kell., in which the stipes of the palpus was seen

to be separate from the labium. C. O. Waterhouse (1904) (6)

had already illustrated the fact that in Z. titan P. the palpus

was connected with the maxilla by a delicate band of connective

chitin. I have discovered similar slips not only in Z. titan,

but in Ancistrona procellarie Westwood (text-fig. 25, p. 132)

and in Miteschia pulicaris N.

The Esophageal Sclerite and Lingual Glands.

These structures are unique in the comparative anatomy of

the insect mouth. They occur in their typical form in the

Ischnocera. In the other suborder it is customary to say that

they are either modified or absent, but, as I show below, they

are really present in all the Amblycera with the possible

exception of Latwmcephalum, which was not available for exami-

nation. The esophageal sclerite and glands in the Mallophaga

were first discovered by Snodgrass (4). They are unusually

difficult of dissection in the Amblycera on account of their

delicacy and minuteness and their position below the cesophagus,

having regard to the flatness of the head in the Mallophaga as a

whole.

The sclerite in the Ischnocera is of a densely chitinous character,

usually quite visible through the integument lying in the lower

wall of the esophagus behind the labium. There are two

anterior cornua and sometimes a posterior pair, but these are

absent in LZ. feroa G. (text-fig. 24, p. 131). Lying longitudinally in

the dorsal wall of the esophagus immediately above the sclerite, I

find in Goniodes falcicornis and in Lipeurus ferow a long, narrow

chitinous splint. Towards this, the posterior cornua in G, falci-

cornis curl up the sides of the cesophagus.

The lingual glands are hard, flat, oval pieces of chitin in which

no glandular structure can be detected, though they still await

histological examination. In LZ. ferox and G. falcicornis, if not

in most other Ischnocera, the anterior ends of the two glands are

encompassed by a small compound plate of chitin (text-fig. 24, E),

narrowing towards the edge of the labium. A curious duct,

cross-barred like a trachea, arises from the sclerite and runs

forward, where it bifurcates, one ramus (or ‘ bronchus”) entering

each of the glands. My own dissections lead me to agree with

Mjoberg (1910) (7) and Grosse (8), who regard the “glands” as

chitinous and as part of the sclerite, the whole to be regarded

as a compound hypopharynx*. Normally the hypopharynx lies

* A central canal is almost certainly present in the ducts of some, but its

meaning remains problematical, unless we suppose that a kind of chitinous

sclerosis has overtaken real glands and ducts, until finally, in such genera as

Ancistrona and Trinoton (text-figs. 26 & 29), the original structures have become

obliterated and replaced by chitin.

Proc, Zoor: Soc,—1913, No, IX. 9

130 _ MR. B. F. CUMMINGS ON THE

immediately above the labium in front of the mouth and not

below the esophagus, and on this account Snodgrass gives the

term hypopharynx only to certain setose lobes he found in

Lemobothrium gypsis which actually overlie the labium. A

similar arrangement of lobes is seen in JZ. titan, where, as the

anterior area of the hypopharynx, they are in close relation with

the sclerite and ‘“ glands”—the elaborate posterior area. The

Ischnocera possess an anterior area—the true hypopharynx of

Snodgrass—in the small encompassing plate mentioned above,

which is much reduced in correspondence with the fact that .in

the Ischnecera the whole labium has been shifted backwards so

as to leave the large mandibles a free field. In some genera of

the Amblycera such as Boopia or Heterodoxus the anterior area,

in which, as in Lemobothrium, the “ducts” are incorporated, is

so large as to extend not only as far as above the labium but

even beyond it as two lobes protruding from the mouth (cf

Mjoberg (7), p. 22). The two areas run into each other without

any dividing line.

Varieties of Sclerite and Glands.

I shall now discuss the various forms the hypopharynx assumes

in the different genera of the Amblycera. Modification begins

with Lemobothrium (see diagram, text-fig. 25, p. 132), in which

the sclerite is elongated and the anterior cornua very long,

extending almost up to the bifurcation of the ‘ducts.” The

“‘olands” are less developed. The posterior cornua are present.

In Gyropus the main “nucleus” of the sclerite has disappeared,

the anterior cornua are fused at the base, and the posterior

cornua are longer and the “glands” very much reduced. In

Trinoton, the “glands” as such have quite disappeared, but may

here be represented by the lateral pieces, the bifurcating sclerite

being the broadened “duct.”* The anterior cornua are entirely

fused together. In Ancistrona procellarie the anterior cornua

are quite distinct but adpressed closely against the broad rami;

each of the rami bears in front a rounded plate with a strongly

serrate edge corresponding with the anterior area of the hype-

pharynx. The diagram is intended to show the probable

evolution of the structure in the Ischnocera and Amblycera.

The presence of cesophageal sclerite and “glands” in the

Psocide as well as in the Mallophaga points to the antiquity of

the structures. Several features in the anatomy of the Psocide

* In Dochophorus sphenophorus (text-fig. 27), an Icshnoceran, the ducts are so

minute and delicate as to suggest atrophy. In front, they are accompanied and

supported by a forked piece of chitin, and in other species and genera the duct

is clearly seen to be accompanied along its course by chitin, so that in those

forms where the ducts gwa ducts are absent their former course along the bottom

of the pharynx may be traced in the track of persisting chitin which they leave

behind. It is this forked piece of chitin, rather than the duct itself perhaps, which

is homologous with the bifureating sclerite in Trinoton (text-fig. 29, B) and other

genera.

MOUTH-PARTS OF THE MALLOPHAGA. Sil

and Mallophaga indicate their affinity, and a plausible theory

has been suggested that the latter are Psocids which have

undertaken a parasitic existence. In any case the “glands”

and sclerite must be regarded as part either of their Psocid or

pre-Psocid inheritance, so that the typical cesophageal sclerite

and “glands” of the Ischnocera are older than the modifications

Text-fig. 24.

Hypopharynx and lingual glands of Lipewrus ferox, X 186.

A, esophageal sclerite; B, anterior cornu; C, duct; D, gland;

E, anterior compound plate,

described in the Amblycera, in spite of the fact that the Ambly-

cera by the possession of jointed palpi, by the form of the

alimentary canal, by the position of the mouth-parts which have

not shifted backwards, and in several other respects appear to be

the less specialised of the two suborders,

132 MR. B. F. CUMMINGS ON THE

Text-fig, 25,

Wi iry

gs mIPIRN

a Sy %

DraGRam to illustrate the evolution of the hypopharynx.

(1) Lipeurus. (2) Lemobothrium. (3) Hypothetical. (4) Ancistrona.

A, edge of labium; B, anterior compound plate; C, esophageal sclerite; D, “duct”;

E, rami; F, ‘‘ glands” or lateral pieces; G, anterior cornu.,

MOUTH-PARTS OF THE MALLOPHAGA. 133

Distribution of the Sclerite and “ Glands.”

A revised list of the distribution of the sclerite and “ glands’

in the Mallophaga must now stand :—

and glands more or | and glands modified. | and glands absent. and glands present

less typically or absent.

developed.

—

| IscHNOcERA. AMBLYCERA. IscHNOCERA. Dochophorus ?

Goniodes, Boopia, Ornithobius ? Lipeurus ?

Goniocotes, Nitzschia, Akidoproctus ? Menopon f

Onchophorus, Heterodoxus, Nirmus?

Trichodectes, Trinoton, AMBLYCERA.

Eutrichophilus, +Trimenopon, Eureum ? i

Eurymetopus, Ancistrona,

Giebelia, Latumcephalum ?

*Ornicholax, Gliricola ?

*Kelloggia.

IsCHNOCERA.

AMBLYCERA. Dochophorus |

Colpocephalum, sphenophorus, |

Lemobothrium, Tetrophtalmus,

Physostomum, Pseudomenopon.

Gyropus.

Genera with sclerite | Genera with sclerite | Genera with sclerite | Genera with sclerite

All the species examined of Virmus, Menopon, Lipeurus, and

Dochophorus in the collection of the British Museum possess

sclerite and glands. Snodgrass records them as absent from

NV. signatus, L. picturatus, and L. longipilus, D. icterodes, and the

following species of Menopon :—distinctum, malleus, persignatum,

precursor, rediculosum, robustum, titan, and trvidens. The sclerite

and glands are certainly present in titan and in tridens, though

they are considerably modified and require careful dissection.

They are described below. From a summary inspection of

Dochophorus sphenophorus N. the structures appeared to be absent,

but here again on dissection I found them present in a modified

form (text-fig. 27, p. 135).

Descriptions of Special Cases.

Ancistrona procellarie Westwood (from Procellaria capensis).—

The hypopharynx was incorrectly described and figured by

Westwood (1874) (8) as a rod bifurcating in front. The lateral

pieces (text-fig. 26, D) were omitted. In all probability the

* The typical sclerite and glands where they occur are plainly visible through

the integument, and, according to figures of these two genera, are present in them.

When modified, the sclerite and glands cannot be discovered without dissection,

and this, perhaps, accounts for the third column being so Jong in the list given by

Snodgrass.

} Gen. noy. about to be described in the Bulletin for Entomological Research.

134 MR. B. F. CUMMINGS ON THE

lateral pieces became detached while the preparation was being

made, as they are very delicately connected. Snodgrass in his

account of A. gigas, a closely allied species, does not mention

any lateral pieces, figures the hypopharynx asa bifurcated rod

and calls it the epipharynx. But the presence of ‘ maxillary

forks” is recorded in A. gigas, and these I have been unable to

find in A. procellarie. If they are absent, it might very well be -

that Snodgrass has mistaken the detached lateral pieces for

maxillary forks. They are alike in shape. The maxille are

attached to the lateral pieces (text-fig. 26, E).

Text-fig. 26.

Hypopharynx of Ancistrona procellaria, X 202.

A, cesophageal sclerite; B, ramus; C, anterior hypopharyngeal plate;

D, lateral piece; E, 1st maxilla.

-Dochophorus sphenophorus N. (from Platalea leucorodia).—This

species presents an interesting transition between the glands and

sclerite typical of the Ischnocera and the conditions found in the

MOUTH-PARTS OF THE MALLOPHAGA, 135

Text-fig. 27.

Hypopharynx of Dechophorus sphenophorus, X 360.

A, esophageal sclerite; B, duct; C, bifid sclerite; D, lateral piece;

E, anterior cornu; F, posterior cornu.

Text-fig. 28.

Hypopharynx of Tetrophtalmus titan, X 204.

ap esophageal sclerite; B, the two rami; C, lateral: Deas,

~ D, posterior cornu; E, maxillary fork.

136 MR. B. F. CUMMINGS ON THE

Amblyceran genera Frinoton, Ancistrona, etc. The glands are

almost completely transformed into lateral pieces (text-fig. 27, D),

but may still be represented by an indistinct circular impression

on each lateral piece in front where it is joined by the delicate

ducts which are supported by a forked piece of chitin, homologous

probably with the forked piece in Trinoton and others. The

cesophageal sclerite is of a peculiar form, quite different from

that of other Dochophori and Ischnocera. It is not visible

without dissection.

Tetrophialmus titan Piag. (formerly M. titan) [from the

Pelican |.—The rami of the broad “duct” bifurcate far forward

at the front margin of the labium. No traces of the “ glands”

(text-fig. 28, p. 135). In Pseudomenopon tridens (formerly

M. tridens), the hypopharynx is very similar, but the lateral pieces

are narrow in front where they run into the rami.

Text-fig. 29.

Hypopharynx of Trinoton luridum, X 246,

A, esophageal sclerite ; B, fused rami; C, anterior lobes of hypopharynx ;

D, lateral piece ; KE, maxillary fork ; F, posterior cornu.

Trinoton luridwm N. (from the Duck).—The anterior cornua

have disappeared—probably fused in the central rod, which is

MOUTH-PARTS OF THE MALLOPHAGA. 137

unusually broad here. The anterior area of the hypopharynx

contiguous with the curving rami of the “duct” consists of two

lobes with a few minute hairs (text-fig. 29).

Text-fig. 30.

aa---7-" tween,

---

tet tt tas

Hypopharynx of Nitzschia pulicaris, X 570.

A, esophageal sclerite; B, lateral piece; C, posterior cornu.

Nitzschia pulicaris N. (from the Swift).—The lateral pieces are

broad and thinly chitinized. The ‘ducts” are broad, and there

are teeth on the anterior area (text-fig. 30).

138 - MR, B. F. CUMMINGS ON THE

« Heterodoxus macropus Le Souéf (from the Wallaby).—Numerous

setee on the anterior area. The lateral pieces exhibit no traces of

glands. There is a projection on the inner side of each lateral

Text-fig. 31.

Hypopharynx of Heterodoxus macropus, X 500.

A, esophageal sclerite; B, rami; C, anterior lobe of hypopharynx ;

D, lateral piece. cared

piece which almost meets its fellow from the other side (text-

fig. 31). Boopia is similar, with an elegantly curved hypopharynx

thickly clothed in front with sete. : OW heen aed

MOUTH-PARTS OF THE MALLOPHAGA. . 139

-- Physostomum -sp.? mystax.—The “glands” are present but

appear to underlie the ducts, with which I cannot find their

point of attachment. Only one rather poor specimen formed

my available material for dissection in this species. The anterior

cornua are very long and curl round in front (text-fig. 32).

Text-fig. 32.

Physostomum sp., X 219.

A, esophageal sclerite; B, glands; C, rami; D, anterior cornu.

The Maxillary Forks.

These problematic “forks” are delicate chitinous rods un-

connected with the rest of the mouth-parts, but lying within the

mouth, one on either side of the hypopharynx. They have been

recorded from 4. gigas P., Goniodes dissimilis N., Lemobothrium

gypsis Kell. I find them present in Tetrophtalmus titan P. (text-

fig. 28, E), Lem. titan P., Trinoton luridum N. (text-fig. 29, F).

On account of their fragile nature they may easily be overlooked,

and therefore perhaps exist in many other species. It is tempting

to look upon these “forks” as the maxillule or super-lingue of

the hypopharyngeal or fifth segment of the insect head. A

third pair of maxille are present and well developed in many

Apterygota, and recently Prof. G. H. Carpenter and Miss Mabel

MacDowell have made a further contribution to the question of

the serial homologies of the insect head by a paper (1912) (10) on

the mouth-parts of certain beetle larvee, where the maxillule are

represented, it is suggested, by the side pieces of the hypopharynx.

This question in the Mallophaga is worth investigation. An

alternative suggestion is that the maxillary forks are the inner

140 ON THE MOUTH-PARTS OF THE MALLOPHAGA.

lobes of the first maxille, the maxillule being the glands and

rami (or the lateral pieces as the case may be). This view

has been actually put forward on behalf of the “glands” by

Dr. Giinther Enderlein (1903) (11).

Conclusion.

Dissections of the head, particularly in those species of the

large genus Menopon where the hypopharynx has been reported

to be absent, ought to yield useful data for systematic work. It

is curious that the case of Dochophorus sphenophorus from which,

at first, the glands and sclerite appeared to be absent, should on

careful examination be found to present a hypopharynx con-

stituting so abrupt a deviation from the normal form in the

Ischnocera, because this species is a typical Dochophorus differing

in external character but little from the group to which it

belongs. Now that schemes of classification, instead of being

merely arbitrary modes of arrangement for the convenience of

systematists, are expected to indicate phylogenetic relationships

whenever possible, it becomes necessary to include in descriptive

work internal as well as external characters. If the hypopharynx

were an external character, its peculiar form in D. sphenophorus

would, with some students of the order, be sufficient reason for

instituting a new genus for its reception.

The cesophageal sclerite is probably present in all Mallophaga,

and in breaking up many of the unwieldy genera such as Docho-

phorus and Menopon, it should prove to be of great assistance.

Of the list of species of Menopon in which Snodgrass discovered

no sclerite present, one, M/. tridens, has already been separated

out as Pseudomenopon tridens, and another, JW. titan, as Tetro-

phtalmus titan. Both these new genera are now found to possess

an cesophageal sclerite, though in a modified form. These facts

are suggestive.

Literature.

(1) Surptey, A. E.—Ectoparasites of the Red Grouse. P.Z.S.

1909, pp. 310-820.

(2) Nrrascu, C. L.—Der Familien und Gattungen der Thierin-

sekten (Insecta epizoica) als Prodromus einer Natur-

geschichte derselben. Germar’s und Zincken’s Magazin

fiir die Entomologie, Bd. iii. 1818, Halle.

~ (3) Grosse, Franz.—Beitriige zur Kenntniss der Mallophaga.

Zeitschr. Wissensch. Zool. 1885, xlii. pp. 536-540.

(4) Snoperass, R. E., in Kellogg’s ‘‘New Mallophaga, II.”

Proce. Calif. Acad. Sci. ser. 2, vi. 1896, pp. 484-457.

(5) Snoperass, R. E—A Revision of the Mouth-parts of the

Corrodentia and the Mallophaga. Trans. Amer, Ent. Soc.

1905, pp. 297-305.

(6) WaterHouss, C. O.—Mouth of Lemobothrium titan. Ent.

Soc. of London, 1904, Proc. pp. 5-6.

ON DEATHS IN THE GARDENS DURING 1912, 141

(7) Msopere, Eric.—Studien tiber Mallophagen und Anopluren.

Arkiv for Zoologi, Bd. vi. no. 13, 1910.

(8) Westwoop, J. O.—Thesaurus, 1874.

(9) Snoperass, R. H.—The Anatomy of the Mallophaga. Occas.

Papers Calif. Acad. Sci. 1899, vi. pp. 148-150.

(10) Carpenter, G. H. & Manet MacDowetni.—The Mouth-

parts of some Beetle Larvee (Dascillide and Scarabeidee),

with special reference to the Maxillule and Hypopharynx.

Q. J. M.S. 1912, vol. Ivii. part 4.

(11) EnpEriern, G.—Ueber die Morphologie, Gruppierung und

systematische Stellung der Corrodentien. Zool. Anz.

xxvi. pp. 424-428, 1903.

Postscriptum.

Dr. W. T. Calman has kindly drawn my attention to a memoir

in the ‘ Bollettino della Societa di Naturalistiin Napoli’ (vol. xxiv.

ser. li. vol. iv. anno xxiv.), published in 1911, and entitled

‘“ Contributo allo Studio dei Mallofagi, Observazione sul Menopen

pallidum.” This paper unfortunately arrived in this country too

late for consideration. The author, Euclide Armenante, investi-

gates M. pallidum and finds that the “ glands” and “ducts” are,

as I suppose, chitinous. The hypopharynx is not typical in this

species, but lends support to the homologies indicated above, and

appears to stand somewhere between Lamobothrium and such

forms as T’rinoton, Nitzschia, ete.

1i. Report on the Deaths which occurred in the Zoological

Gardens during 1912, together with the Blood-Parasites

found during the Year. By H. G. Purmmer, F.R.S.,

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

[Received and Read February 4, 1913.]

On January Ist, 1912, there were 885 mammals, 2180 birds,

and 518 reptiles in the Zoological Gardens ; and during the year

506 mammals, 1346 birds, and 648 reptiles were admitted,

making a total for the year of 1391 mammals, 3526 birds, and

1166 reptiles.

During 1912, 375 mammals, 817 birds, and 347 reptiles have

died: that is, a percentage of 26°9 for mammals, 23-2 for birds,

and 29°8 for reptiles.

633 deaths out of the total of 1539 for the year occurred in

animals which had not been six months in the Gardens. It has

been found that after six months’ residence in the Gardens the

percentage falls rapidly, so that it is assumed that by that time

the new animals have got over their journeys, or have died from

any diseases they may have brought with them, or have got quite

142 MR. H. G, PLIMMER ON DEATHS

used to their newenvironment. 144 of these 633 were mammals,

317 were birds, and 172 were reptiles; and if these be deducted

from the above the percentage appears as 16°6 for mammals,

14:2 for birds, and 15 for reptiles.

The following tables show some of the facts ascertained in

outline. Table I. summarizes the actual causes of death in the

three groups specified. Under Reptiles are included Amphibia.

Tas.e I.—Analysis of the Causes of Death.

Reference

Diseases. Mammals.| Birds. |Reptiles.| to Notes

following.

1. Microbie or Parasitic

Diseases.

Tuberculosis ..............-..0-00 0: 14 79 11 1

IMYcOsist erence seeteee ae eae 12 72 2 2

iBneumomiay | ese wanes eee 45 98 | 124 3

Septitcsemiianer re seaseeree eee aes 2 oF ds

Pericandubisumese sence eeereeeeeeeeee 5 1 as

Stomatitisiis..005. ices sscse- te te - 4

Peritomitisysscteseds.-c-ek scenes 7 age es

IADSCESS sare Ss Pater ie ete eee 1 1

JOTI VAUNE) gosoodooo cde esosendsc ane dae il

Hydatids > ).0. 28 See 4

Wroorins bc dtee nek nem tate 4, 1 4

Dermatitis (sarcoptic) ............ 1

2. Diseases of Respiratory

Organs.

IBronchitisieemessseeeseee eee ae 12

Broncho-pneumonia ............... 30 sae oe

Congestion of lungs ............... 14 108 22

Artelectasist¢oc..0:.2accenseentonmaes: 2 ae a

3. Diseases of the Heart and

7essels.

ATNCUTISDU ess rssh cece ote se eee ee eoes a5 1 wae 5

4. Diseases of the Liver.

Fatty degeneration ............... 1 7

Hepatitis), cSijcccecesosseees goueseers ses 14

5. Diseases of the Alimentary

Tract.

Gastritis .. : sauiheer Stee oa 1 a 6

Gastric ulceration ......-sce+s.-:. 8 Bae sos

Gastro-enteritis .. L cicasle aise 11 1 3 ; 7

Enteritis... Py aaaaier 38 154 25

Intestinal obstruction ............ 1 ae ue

6. Diseases of Urinary and

Generative Organs.

INephritis decsssesccutcnesgeeteeseee 89 104 9 8

Cystic kidneys ..................05 1 a ae 9

Stone ........ Ea area 1 bi

Inflamed oviduct . ia. i we 1

Sloughing uterus .................. 1

7. Various.

@ancinomayescaeesbeeeee eke 4 aus ee 10

Sarcoma . Eee 2 1 2 11

Senile Decay .. at 1 ae

Injuries discovered _ Post 3 5 8 1

TOOPEGTO seAnddotonced san voancecco

IN THE GARDENS DURING 1912. 143

Besides those tabulated above,

59 mammals, 100 birds, 2 reptiles, were killed by order

or by companions,

3 Be Oye WARE ier died from malnutrition

or starvation,

6 - Soi epee were too stale for de-

i tailed ination,

these completing the total. alled examination

In Table I. the classification is made of those diseases which

actually caused death. Table II. summarizes the other diseases

from which the animals were suffering; and if this Table be

taken in. conjunction with Table I., a much more accurate

estimate of the amount of disease in the Gardens will be

arrived at.

Taste I1.—Other Diseases found in the animals tabulated

in Table I,

| Reference |

Diseases. Mammals.| Birds. |Reptiles. to Notes |

| following.

Tuberculosis

Mycosis SO nan Eaa raacontad Ruaosemaee ths

PNEUMONIA ee Rey aa ees

IRB LCE REINS) Bataan odeasnebas dbo Reaatosocebos:

Peritonitis Wee aes cerca AL

CULiti Ss eee cee ace Soe ee nO:

Malaria SAA On ANSE BARS Sere cron

JO ENG Wie Saar Gee, oteeet ramen Wea mee teres 4 Prue Sento

WVOTTITS Rea srece deter cctioe eon area more: pp

Heaemogregarines............006000ceeeee eee BN SS 32 14

TA DETOSONNGS 0.0, c00000 000 s00 200 094 Sad o0a ono see a 11 ite 15

StOmaGIbISH cree eee cere ore oe ae ees ie Bas 4,

Abscess On Ae eM RRA ete Mas

GOCCTATOSIShY serine soma eee ee cee

37 esi Dy Ne

WE HOw

BrOnchitisy aoc ecccsisassoe wa sanaee asec wack: 8

Broncho-pneumonia ....................--.- 12 ner, ee

Congestion of lungs ...................0005- 34 63 10

(iidemajotlungsy ese eee 1 42 21

Wilatedtheart! vase vasnccncckece tena seee aces 5

HAE erOMa see. ceee ee ee on ee eee 7 1

Nat tyslivercekacteseea. casas eases 20

Hepatitis tp csscesenaeacannec etnaencacteuae ae 3 2

Cirrhosis of liver Les ee oc ash Eas 2

Gastritisteee Sie ec ee ok a aT 1

Gastric ulceration ..,............0...0e cece 18

Gastro-enteritisimy ce steeatctee eee sees 3

ITIGCLIGISN roe ee eee eee eee eee 34 6

Intussusception ............... cc ccc cence eee 3

Intestinal obstruction ..................... 2

INfep hiritis): 3 eesterstee acne ue nelaec een nak 52

Cystitiste tree ren eee 2

Inflamed oviduc Reet eee Dean eret || bs

ICKL Aiisrc canna eyes Aca eh 17

Ast ritistes ccc eee ah a CE eel ae

Chylous ascites ...........0.0 cee cece ee 1

3 te :

Seo: ee eb:

144 MR. H. G. PLIMMER ON DEATHS

Table III. shows, in still further detail, the distribution of

diseases amongst the various orders of nammals.

Tass III.—Showing the Distribution of Diseases causing Death

amongst the principal Orders of Mammals.

| | | la

| . | . =

a | 2 a iit Ol] arte

1 = ov = | te = i) =

Diseases. = Es hee 4 = | =}

= Bll egy Seay eh

olor elleneaanl (a iaicd vl etn

Peele alte ute S.. pie lervie| Museaael é es sens

| Tuberculosis 5 6 | Zea geal fanaa |

Mycosis SHER EP Ok a WAI aed laa Bees tlh ae ses 10 |

Bneumoniayceseprer eee eae 15 10 4 5 3 |

Septiceemiayy anes ce ences soeene asec eure 1 ity Meteo

IADSCOSS a ia c Ste etn Ae iis ae seal ae al ae |

Peri Can Gitlsy tn ose coe pee eee eet ese ces 2 1 aes Me leGenek a |

FPOrivOnitis: sue MaMa toads at cmesmoca inners Seay ak vedere

Empyema Re rh bacon Se Ae BL ie ee wel |

WWermatinis (SaECOptiC) ese ee -seee| eee See Wpteal |

lipeliy deatiidl’s eats... cena toon ue ice cea|io taal Risto 3. | |

IWVORINS ene eceeicees ose oe. vesbee meee 2 ee pena at

| |

| | | |

Bronchitisteecnece cetacean sce eeeee alee 6 pa eee Te ll poses al |

| Broncho-pneumonia ...................../ 11 hig avec arse 8) Pe, al

Congestion of lungs ..................5:. 7 Li 6 He | |

WNtelectasiss 2as.: secereeseee nc: Geometers. Sa pal ee 2 erste)

Fatty degeneration of liver ............ | Ae 1

Gastritiste Perse eon. eeoe es qoaenmsateee ado. ats 1 |

Gastro-enteritis <2. .2.......25cee0--- 2-0 ++: 1 Te IN a 2) |

Gastric ulceration ........................| 3 1 2 2 | |

Bintberibi see nn hee CRG a Real Glia, Th ey ue) (ec «|

Intestinal obstruction ...............-.. Da ie | Ea eae Damen: | il

IN/GIOI AIM Coassodeococonoondo bes scuaandna cos) | ues 31 | 15 LSe i e25G| eae

Cystic kidneys Ane a eee A ese ee ere ne ce enya |

SCONE Hise ie Beta ete enapoue seanceseremen a \iuemamau!

Wan cere tigi Besse. cotaen annem aaa anaes ee 3 | al

ISENRSONTE), | Goaonn enciona nepeeoscead ocuenb aceon 2 | |

Notes on the foregoing Tables.

1. There has been a general decrease in the deaths from

tubercle during the past five years, which has been most marked

in mammals. In 1908, 59 mammals died from tubercle, last

year only 14. The percentage of deaths of the total number of

animals is 1 per cent. for mammals, 2°2 for birds, and 1 for

reptiles. Of the 14 mammals, 9 had not been six months in the

Gardens; and 9 were pet animals which had been presented,

1 shared burrows in the squirrel’s enclosure with the rats (which

we know are infected), 1 had been in captivity in the Hast for

some months before reaching London, and 1 was a tame animal

which had been reared by hand. So that only 2 of the old

mammal inhabitants died last year from tubercle.

IN THE GARDENS DURING 1912. 145

Of the bird cases, 52 were generalised tubercle, and 6 were of

bovine type.

2. All the mould-diseases have been grouped under mycosis.

Ten of the mammal cases were in Wallabies, and of the same type

as that I have previously described, affecting primarily the jaws ;

the other 2 were in Cercopitheques in which the infection was

localised in the intestines and kidneys. There isaslight decrease

in the number of deaths from mycosis in birds, but it is still

large, and is 2 per cent. of the total number of birds.

3. There is an increase in the deaths from pneumonia in birds

and reptiles. The percentage on the total number of animals is

3-2 for mammals, 2:7 for birds, and 10 for reptiles. It is

amongst the reptiles that the increase has been most marked,

from 4 in 1908 to 120 in 1912 (these are pneumococcal cases and

do not include those due to irritation from worms’ eggs and

embryos).

4, “ Worms” is used in a comprehensive sense; two of the

mammals were Indian Buffaloes with an enormous trematode in-

fection of the stomach.

5. This was a ruptured aneurism of the ascending aorta in a

Pigeon.

6. In a Duck, after swallowing wire.

7. The percentage of gastro-enteritis is still high: 3°5 for

mammals, 4°3 for birds, and 2°4 for reptiles, on the total number

of each. It has been noticed during the last year in connection

with enteritis in mammals that there has often been an associated

condition of gingivitis, not bad enough to call pyorrhcea, which

possibly may stand in causal relationship to the enteritis. In

2 mammals it was of coccidial origin, and in 2 others it was due

to worms. Of the bird cases 72 were hemorrhagic and 9 were

due to foreign bodies. It has much decreased amongst the

reptiles, and 6 of the cases in this class were due to worms.

8. Nephritis has increased amongst mammals and birds, having

caused the deaths of 6°4 per cent. in the former and 2°9 in the

latter. Of the cases in mammals 60 were acute, 10 in condition

of “large white” kidney, and 19 in condition of “ contracted

granular” kidney. In birds it is nearly always chronic, about

one-third of the number being of the contracted granular kidney

type. A conjunction of climatic conditions with exposure would

seem to be answerable for many of the cases in mammals, and

40 out of the 60 acute cases had lung lesions, from congestion to

broncho-pneumonia.

9. In a Bay Duiker, the left kidney was converted into

multiple cysts containing stones, and there was one large cyst in

the right ; there was also a hemorrhagic cystitis.

10. The carcinomata were all visceral, and occurred in one

Kangaroo and three Gazelles. In all the cases the initial growth

was in the stomach, and there were atrophic changes in the

sexual glands.

Proc. Zoou. Soc.—1i913, No. X. 10

146 ; MR. H. G. PLIMMER ON DEATHS

11. The sarcomata occurred in a Binturong (visceral), a Bear

(kidneys), an Owl (heart), and a Sternothere (heart).

12. The diseases grouped under the term malaria were due in

31 instances to Hemoproteus danilewskyi and in 6 instances to

Plasmodium precox.

12, 13, 14, 15. See the section on blood-parasites.

During the year the blood of every animal which died has

been examined, with the result that parasites have been found in

140 cases ; in 80 species for the first time.

They have been distributed as follows :—

Filarie. In 3 mammals ; in 2 for the first time.

34 birds ; in 24 species for the first time.

2 reptiles.

Trypanosomes. In 11 birds; in 6 species for the first time.

Hemoproteus danilewskyi. In 33 birds; in 15 species

for the first time.

Plasmodium precox. In 7 birds; in 6 species for

the first time.

Leucocytozoon. In 1 bird for the first time.

Hemogregarines. In 48 reptiles; in 15 species for the first

time.

teen TES ON Te reptile for the first time.

richomonas type ...

Malaria.

The following Tables show the occurrence of the blood-parasites

in detail :-—

BiLoop-PARASITES FOUND IN 1912.

Embryo Filarie found in the blood of Mammals.

HaBitTat. TYPE.

Pinché Marmoset (Leontocebus edipus)... Colombia. Long.

Found in the following for the first time:

Clouded Tiger (Felis nebulosa) ............ Malay. Long, thin, large capsule.

S. American Night-Mouse (Nyetomys C. America. Short, thick capsule.

sp. inc.).

Embryo Filarice found in the blood of Birds.

Black-throated Hangnest (Icterus gu- S. America. Short, straight.

laris).

2 Blue Birds (Sialia sialia) ................ N. America. | Short, thick.

Whydah Bird (Urobrachya albonotata). E. Africa. Long.

Occipital Blue Pie (Uvocissa occi- India. Short, thick.

pitalis).

Green-billed Toucan (Ramphastos di- Guiana. Short, thick.

colorus).

Lawes’ Bird of Paradise (Parotia New Guinea. Long, thick.

lawesi).

White-throated Jay Thrush (Garrulaa India. Long, striated.

albigularis).

Lanceolated day (Garrulus lanceolatus). India. Short, pointed. ©

IN THE GARDENS DURING 1912.

Found in the following for the first time :

Crested Black Bunting

melanicterus).

Green Cardinal (Gubernatria cristata) .

Gouldian Grass-Finch (Poéphila gouldie)

Mynah (Acridotheres tristis) :

Yellow Sparrow (Passer luteus)... ........+

Crossbill (Leaxia curvirostra) ads

Grey Thrasher (Towostoma cinereum) ...

(Melophus

White-bellied Thrush (Turdus albi-

ventris).

Red Jungle-Fowl (Gallus gallus) ......

2 Blue-crowned Hanging Parrakeets

(Loriculus galgulus).

White-cheeked Coly (Colius erythro-

neelon).

Rutous-tailed Pheasant (Acomus ery-

throphthalmus).

Woodhouse’s Jay (Aphelocoma wood-

housei).

Hermit Thrush (Hylocichla

pallasi).

Burrowing Owl (Speotyto cunblendlce Ba

hypogea).

Little Owl (Athene noctua) Haile

Purple Grackle (Quiscalus purpur olas

Mexican Jay (Xanthura luxuosus)

guttata

Grey-headed Ouzel (Merula castanea) ...

Rufous-necked Weaver-bird (Hyphan-

tornis textor).

Green Bulbul (Chloropsis aurifrons) ...

Drongo (Dissemurus paradiseus) .........

Red-headed Weaver-bird (Foudia mada-

gascariensis).

Scops Owl (Scops iv) ......csessesseveess

HABITAT.

India.

S. America.

Australia.

India.

E. Africa.

Europe.

Lr. California.

Demerara.

Malay.

S. Africa.

Malay.

N. America.

C. America.

S. Europe.

N. America.

S. America.

India.

Gambia.

India.

Madagascar.

N. Europe.

147

TYPE.

Long, thick.

Short, pointed.

Short.

Long and thin.

Long.

Long, thick.

Short, thick.

Long, thick.

Short, thick.

Long, striated.

Long, thin.

Short, thick.

Long, thick.

Long, pointed, and with

very large capsule.

Short.

Very short, stout.

Long, thin.

Short, thick.

Long.

Long, striated.

‘Embryo Filarice found in the blood of keptiles.

Edible Frog (Rana esculenta)

Pine Snake (Pitwophis sayt) ......s00.cc00

S. Europe.

N. America.

Short, thick.

Long.

Trypanosomes found in the blood of Birds.

Dial Bird (Copsychus saularis) ... 1.0... 44.

India.

Found in the following for the first time:

Blue Bird (Sialia sialia)

Whydah Bird (Urobrachya Likenomeane

Gouldian Grass-Finch (Poéphila gouldie)

2 Grey Thrashers (Towxostoma cinereum).

4 Blue-crowned Hanging Parrakeets

(Loriculus galgulus).

Wandering Tree-Pie (Dendrocitta vaga-

bunda).

N. America.

E. Africa.

Australia.

Lr. California.

Malay.

India.

These were all of the ordinary type of bird Trypanosomes,

148

MR. H. G. PLIMMER ON DEATHS

Hemoproteus danilewskyi fownd in the blood of Birds.

Yellow-winged Sugar-Bird (Ocreba

cyanea).

11 Blue-crowned Hanging Parrakeets

(Loriculus galgulus).

Cape Sparrow (Passer arcuata)

Crossbill (Loxia eurvirostra)

Little Ow] (Athene noctua)

HABITAT. Tyre.

S. America.

Malay.

S. Africa.

Kurope.

Europe.

Found in the following for the first time :—

2 Pratincoles (Glareola pratincola) ......

3Silver-eared Mesias (Mesiaargentauris).

De Philippi’s Meadow-Starling (Starnella

defilippi).

Whydah Bird (Penxthetria laticauda) ...

Jerdon’s Accentor (Tharrhaleus jerdoni).

Banded Parrakeet (Paleornis fasciatus).

Crested Black Bunting (Melophus melan-

icterus).

Yellow-throated Sparrow (Gymnorhis

flavicollis.

White-throated Jay-Thrush (Garrulax

albigularis).

Yellow-headed Reed-bird (Ageleus ictero-

cephalus).

Rufous-necked Weaver-bird (Hyphant-

ornis textor).

Red-headed Weaver-bird (Foudia mada-

gascar ee

Wandering Tree-Pie (Dendrocitta vaga-

bunda).

Swainson’s Francolin (Pternistes

swainsoni).

Red Jungle-Fowl (Gallas gallus)

India.

Chili.

E. Africa.

India.

Mexico.

Gambia.

Madagascar.

India.

S. Africa.

Malay.

Plasmodium precox fownd in the blood of Birds:

in all for the first time.

Grey-headed Bunting (Emberiza fucata).

Crested Black Bunting (Melophus melan-

icterus).

Raven (Corvus coraz) .. seeks

2 Crossbills (Loxia eurvir a ay :

Pied Bush-Chat (Pratincola euprata

Grey Thrasher (Loxostoma cinereum) .

India.

Europe.

India.

Lr. California.

Leucocytozoon found in the blood of the following Bird

Sor the first time.

Scops Owl (Scops giu) ..........

N. Europe.

Hemogregarines found in the blood of Reptiles.

4 Common Boas (Boa constrictor)

2 Indian Pythons (Python molurus)

S. America.

India.

Long, host cells enlarged.

Medium, cells deformed.

IN THE GARDENS DURING 1912,

HasirTat.

8 King Snakes (Coronella getula) ...... N. America.

Teguexin (Tupinambis teguexin) ........._ S. America.

Eyed Lizard (Lacerta ocellata) ............ S. Europe.

4 Dark Green Snakes (Zamenis gemon- S. Europe.

ensis).

Russell’s Viper (Vipera russelli) ......... India.

2 Rat-Snakes (Zamenis mucosus) ........- India.

Indian Cobra (Nata tripudians) . India.

Tuberculated Iguana (Lguana beds S. America.

lata).

Indian Eryx (Hrya# johni) ......... India.

4 Diamond Rattlesnakes (Orotalusa days N. America.

2 Pine Snakes (Pitwophis sayi) ............ N. America.

Vivaceous Snake (Zarbophis fallax)

Green Mamba (Dendraspis viridis) ......

European Pond-Tortoise (Hmys orbi-

cularis).

Long-nosed Viper (Vipera ammodytes) .

S. Europe.

W. Africa.

S. Europe.

Found in the following for the Jirst time :—

Robust Lizard (Gerrhosaurus

lineatus).

Angulated Tortoise (Testudo angulata) .

Schott’s Snake (Philodryas schottt) ......

nigro-

S. Africa.

S. America.

Copper-headed Viper (Ancistrodon con- Texas.

tortrix).

Common Krait (Bungarus candidus) ... India.

Hamadryad (Naia bungarus) ............ India.

W. Africa.

C. America.

Calabar Snake (Calabaria reinhardti) ...

Blood-stained Terrapin (Cinosternwm

cruentatum).

Indian River-Snake (Tropidonotus India.

piscator). :

Baska Water-Tortoise (Batagur baska). Perak.

2 Phayre’s Tortoises (Lestudo envys) ...... Malay.

2 Thick-necked Tree-Boas (Hpicrates ‘Trinidad.

cenchris).

Carpet-Viper (Echis carinatus) ......... N. Africa,

Water-Viper (Ancistrodon piscivorus)... N. America.

Amboina Box-Tortoise (Cyclemys am- Malay.

boinensis).

Cape Colony.

149

TYPE.

Small, short.

Large, granular cells

enlarged.

Long, thin, host-cells

enlarged.

Small, host-cells enlarged.

Long.

Medium sized.

Long.

Short.

Long.

Large, host-cells enlarged.

Long, cells enlarged.

Medium.

Large.

Short, thick.

Large.

Large, granular, host-

cells enlarged.

Large, irregular.

Small, short.

Long; snake had been

12 years in Gardens.

Short and long.

Large, stout.

Medium.

Long.

Long, thick.

Stout.

Long, cells enlarged.

Large, cells enlarged.

Long.

Thick, cells enlarged.

Short, thick.

Intestinal Organisms found in the blood of Reptiles.

Found for the first time in the following :—

Amboina Box-Tortoise (Cyclemys am- Malay.

boinensis).

Trichomonas.

150 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

EXHIBITIONS AND NOTICES.

February 4,.1913.

Sir Joun Rose Braprorp, K.C.M.G., F.R.S., Vice-President,

in the Chair.

THE SecRETARY read the following report on the additions made

to the Society's Menagerie during the months of November

and December, 1912 :—

NOVEMBER.

The registered additions to the Society’s Menagerie during

the month of November were 161 in number. Of these 65 were

acquired by presentation, 66 by purchase, 11 were received on

deposit, 9 in exchange, and 10 were born in the Gardens.

‘The total number of departures during the month, by death

and removals, was 229.

Amongst the additions special attention may be called to the

following :—

1 Humboldt’s Saki (Pithecia monachus), from Iquitos, purchased

on November 20th.

1 Kusimanse (Crossarchus obscurus), from Axim, Gold Coast,

presented by R. W. Brent, Esq., on November 8th.

2 Black-spined Porcupine Anteaters or Echidnas (Zaglossus

nigroaculeatus ?), from Charles Louis Mountains, New Guinea,

new to the Collection, deposited on November 2/th.

DECEMBER.

The registered additions to the Society's Menagerie during

the month of December were 79 in number. Of these 40 were

acquired by presentation, 4 by purchase, 17 were received on

deposit, 7 in exchange, and 11 were born in the Gardens.

The total number of departures during the month, by death

and removals, was 168.

Amongst the additions special attention may be called to the

following :—

2 Rufted Lemurs (Lemur varius) 3 9, received in exchange on

December 16th.

1 White-tailed Gnu (Connochetes gnu) 2, purchased on

December 10th.

3 Mahali Weaver-birds (Plocepasser mahali), from South Africa,

new to the Collection, received in exchange on December 17th.

1 Pink-browed Rose-Finch (Propasser rhodochrous), from the

Himalayas, new to the Collection, presented by W. H. St. Quintin,

Esq., F.Z.8., on December 19th.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 151

1 Nepalese Eagle-Owl (Huhua nipalensis), from the Maikola

Valley, Hast Nepal, purchased on December 4th.

1 Bushmaster (Lachesis mutus), from Trinidad, presented by

the Baron Leijonhufvud, F.Z.8., on December 10th.

Mr. D. Sers-Surry, F.Z.S., Curator of Birds, exhibited a

female Carolina Duck (Lampronessa spoisa) which had partially

assumed the male plumage. It had shown no sign of this con-

dition before the last moult. It now resembled the male in most

respects, although the colours were duller and the characteristic

markings less clearly defined, but it retained the brown colouring

of the eye and the yellow skin and white patch of feathers sur-

rounding it, which were characters of the female.

Mr. E. G. Boutencer, F.Z.S., Curator of Reptiles, exhibited a

blue specimen of the Edible Frog (Rana esculenta) which he had

obtained from Tuscany. The blue colour was attributed to the

absence of yellow pigment, which is present in large quantities

in green frogs. This aberration is not very unfrequent in the

European Tree-Frog (Hyla arborea), but very rare in the Kdible

Frog, and this was the first occasion on which such a specimen

had been exhibitéd in the Gardens. Blue Edible Frogs have

been previously reported from France, Western Germany, and

Switzerland.

Mr. BouLEeNnGeER also exhibited a living specimen of the remark-

able Lizard Pygopus lepidopus, a species which is rarely seen

alive in this country, and which had not been represented in the

Society's Collection for some considerable time.

February 18, 1913.

Prof. KE. A. Mtncutn, M.A., F.R.S., Vice-President,

in the Chair.

THe Secrerary read the following report on the additions that

had been made to the Society’s Menagerie during the month of

January, 1913 :—

The number of registered additions to the Society’s Menagerie

during the month of January last was 120. Of these 38 were

acquired by presentation, 70 by purchase, 1 was received on

deposit, 1 in exchange, and 10 were born in the Gardens.

The total number of departures during the same period, by

deaths and removals, was 189.

152 DR. MITCHELL ON THE DORSAL BAND IN A LIVING TREE-HYRAX.

Amongst the additions special attention may be directed to :—

1 American Bison (Bison americanus) 9, from N. America,

received in exchange on January 16th.

2 Barbary Apes (Macacus inuus) $ 9, from Gibraltar, pre-

sented by Major C. E. P. Fowler, R.A.M.C., on January Ist.

2 Arabian Baboons (Papio hamadryas), from Arabia, 1 Anubis

Baboon (Papio anubis), from W. Africa, 2 Mandrills (Papio

maimon) 6 @, from W. Africa, 1 Egyptian Mongoose (Mungos

ichneumon), from Egypt, 1 Indian Green Barbet (Thereiceryx

zeylonicus), from India, 1 Iceland Falcon (Hierofalco islandus),

from Iceland, and 2 Black-necked Crowned Cranes (Balearica

pavonina), from Nigeria, presented by W. O. Danckwerts, Esq.,

K.C., F.Z.8., on January 22nd, 24th, and 28th.

3 Cuban Black Bullfinches (Melopyrrha nigra), purchased on

January 11th.

5 South-African Ostriches (Struthio australis), bred in Australia,

purchased on January 6th.

———

Mr. E. G. Boutencer, F.Z.S., Curator of Reptiles, exhibited

spines of a hedgehog-like Madagascar Insectivore, Hriculus setosus,

which had been found by Mr. C. R. Walter in the excrement of

a Boa madagascariensis, this being probably the first instance on

record of a Snake swallowing a spiny mammal.

The Srecrerary, Dr. P. CHatmers Mircuett, F.R.S., F.Z.S.,

exhibited a series of lantern-slides showing the opening and

closing of the patch of white hairs forming the dorsal band in a

living Tree-Hyrax (Dendrohyrax dorsalis).

No. 114.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

November 26th, 1912.

Dr. A. Smira Woopwarp, F.R.S., Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

Mr. Ouprietp Tuomas, F.R.S., F.Z.S., exhibited and made

remarks on a peculiar Amazonian Monkey (Cailimico goeldi)

which showed characters intermediate between those of the

ordinary American Monkeys (Cebidw) and of the Marmosets

(Callitrichide).

Mr. C. Tate Reean, M.A., F.Z.S8., exhidited male and female

examples of Cynolebias bellottii, a Cyprinodont Fish from the

La Plata, to illustrate the remarkable sexual characters.

A paper entitled ‘‘On the Structure of Bone in Fishes: a

Contribution to Paleohistology,” was read by Mr. E.8. Goopricn,

M.A., F.R.S., F.Z.8. He stated that a microscopic examination

of the bone of the Actinopterygian Fishes showed that in those

groups which are provided with lepidosteoid ganoid scales

[| Amuoidei (Protospondyli) and Lepidosteoidei (Aetheospondyli) |

the characteristic lepidosteoid histological structure extended

throughout the endoskeleton as well as the dermal bones. No

other fishes are known to have this lepidosteoid structure, either

in the scales or in the skeleton,

A paper on the Land Crayfishes of Australia was received from

Messrs. GEorrrey W. Suiru, M.A., and HE. H.J.Scuusrer, M.A.,

D.Se., F.Z.8.

This paper dealt with the genus Hngeus, a group of Victorian

* 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 Stx Shillings per annum, payable in advance.

and Tasmanian Crayfishes, which have forsaken the water and

excavate burrows in damp soil. In certain mining districts on

the west coast of Australia they do much damage to the artificial

water-courses by riddling through the banks and dams and

causing them to collapse.

Although the tunnel leading to the heart of the burrow is free

from water, there is always water in the circular chambers at the

end where the Crayfish lives.

In a former paper it was suggested that these Crayfishes of the

genus Hngeus are derived from the genus Paracheraps, which

has spread from Western Australia into the desert regions of the

centre, and is now found in all parts of continental Australia ;

but conclusive evidence is brought forward in this paper to show

that Engeus is derived from the South-eastern and Tasmanian

genus Astacopsis, and that its superficial resemblance to Para-

cheeraps is due to convergence owing to similar habits.

A striking peculiarity of the eight species of Hngceus is the

very great range of structural differences which they exhibit

compared with other Crayfishes, differences which are far greater

and concern more important characters than those which

distinguish genera or even families of other Crayfishes. Thus in

some species a gill may be entirely suppressed, in others half a

limb (e. g., the exopodite of the third maxillipede) is absent, im

others the flagellum of the antennule. All these characters are

of the nature of degenerations, apparently incident on the sub-

terranean mode of life. Apart from these differences, the

species are obviously closely related and represent a monophyletic

group.

A paper was communicated by Dr. C. L. Bounnezr, M.A.,

F.Z.8., dealing with the Myzostomida collected by Mr. Cyril

Crossland in the Red Sea in 1905, and containing descriptions of

three species, of which one is new. A complete account of the

anatomy of this form is given, as well as that of J/. costatum

F.S. L. The latter species is described as possessing six pairs of

ventral “ suckers,” the largest number recorded in any member

of the group.

The Hon. Paut A. Muruven, F.Z.8., contributed a paper con-

taining the description of a new Amphipod, belonging to the

family Talitride, which had been obtained in the Woodbush

district of Northern Transvaal.

A paper by Mr. Bruce F. Cummines, communicated by the

SECRETARY, and entitled “*‘ On some Points in the Anatomy of the

Mouth-parts of the Mallophaga,” dealt with some peculiar struc-

tures in the floor of the mouth-cavity of the Mallophaga, or the

biting bird-parasites, and known as the wsophageal sclerite and

lingual glands. The “ glands” are determined as chitinous, and

both sclerite and glands together are regarded as a compound

hypopharynx, Variations of the structures in various genera of

the two suborders of the Mallophaga are described and figured

and the homologies of the parts indicated. Attention is drawn

to the value of the hypopharynx as a character in splitting up

unwieldy genera. The writer further describes the problemati 1¢

Gi mreeillary forks’ ’—delicate chitinous splints lying unattached

within the mouth, one on either side of the hypopharynx,—and

suggests that they are the third pair of maxille, known as maxillule

or super une rene of the hypopharyngeal or fifth seement of the insect

head, well developed i in many primitive insects such as the Aptery-

gota, and prababl ya also in certain beetle larvee.

The next Meeting of the Society for Scientific Business will

be held on ‘Tuesday, February 4th, 1913, at half-past Hight

o’clock P.M,, when the following donornncnica ties will be made :—

F. KE, Beppagp, M.A. ; B.S., i, 4. Ss,

Chae ier 46 a eae ystematic Arrangement

of the Cestoidea.—VIII, On some Species of Jchthyotenia and

Ophidotenia from Ophidia.

(Ble UR OO Sey Es as

Report on he ae — aiiciey in the Zoological

Gardens during 1912.

ie iL] Hawsins, M SC. ie ‘G.I 8.

The ingen Same um of Hchinocardium cordatum Penn.,

and the Origin of Compound Plates in Hechinoids.

G. P. Farrayn.

Plankton from Christmas Is!and, Indian Ozean.-—II, On

Copepoda of the Genera Oithona and Paroithona.

The following papers have been received :—

H. B. Preston, F.Z.8.

Diagnoses of new Species and Varieties of Agnathous

Mollusca.

R. LypEK«eEr, F.R.S., F.Z.S.

The Dwarf Buffalo of Southern Nigeria: with a Revision of

the Dwarf Buffaloes of Western Africa.

A few Notes on the Habits of certain Reptiles in the

Lagos District.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Soctery or Lonpon,

Reeent’s Park, Lonpon, N.W.

December 3rd, 1912.

No. 115.

ABSTRACT OF THE PROCEEDINGS

ZOOLOGICAL SOCIETY OF LONDON

February 4th, 1913.

Sir JoHN Rose BrapForp, K.C.M.G., F.R.S.,

Vice-President, in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secretary read a Report on the Additions made to the

Society’s Menagerie during the months of November and

December, 1912.

Mr. D. Sers-Suirn, F.Z.S., Curator of Birds, exhibited a

female Carolina Duck (Lampronessa sponsa) which had partially

assumed the male plumage. It had shown no sign of this con-

dition before the last moult. It now resembled the male in most

respects, although the colours were duller and the characteristic

markings less clearly defined, but it retained the brown colouring

of the eye and the yellow skin and white patch of feathers sur-

rounding it, which were characters of the female.

Mr. HK. G. Boutencrr, F.Z.8., Curator of Reptiles, exhibited

a blue specimen of the Edible Frog (Rana esculenta) which he had

obtained from Tuscany. The blue colour was attributed to the

absence of yellow pigment, which is present in large quantities

in green frogs. According to Mr. G. A. Boulenger, this aberration

is not very unfrequent in the European Tree-Frog (Hyla arborea),

* 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,

but very rare in the Edible Frog, and this was the first occasion

on which such a specimen had been exhibited in the Gardens.

Blue Hdible Frogs have been previously reported from France,

Western Germany, and Switzerland.

Mr, Bovutencer also exhibited a living specimen of the re-

markable Lizard Pygopus lepidopus, a species which is rarely seen

alive in this country, and which had not been represented in the

Society’s Collection for some considerable time,

Dr. F, E. Bepparp, M,A,, F,R.S., F.Z,8., Prosector to the

Society, read a paper, the eighth of the series, on the Anatomy

and Systematic Arrangement of the Cestoidea, in which he dealt

with a number of new species of Jchthyotcenia and Ophidotenia

obtained from the gut of Serpents that had died in the Gardens.

Mr, H, G. Puimmer, F,R.S., F.Z.8,, Pathologist to the Society,

presented his annual report on the deaths which occurred in the

Society's Gardens during the past year, together with a list of

the Blood Parasites found during the same period. An examina-

tion had been made of the blood of every animal that had died,

with the result that parasites had been discovered in 140 cases,

and in 80 of these for the first time.

Mr. Hurpert L, Hawxiys, M.8c., F.G.S,, read a paper, com-

municated by Dr. F. A. Barumr, M.A., F.R.S., F.Z.S., on “The

Anterior Ambulacrum of Hchinocardiwm cordatum and the Origin

of Compound Plates in the Echinoidea.” It contained the de-

seription of a new method of exposing sutures in recent Echinoids

suitable for photographic purposes, the process combining staining

with etching, and the description of the complex plating of

ambulacrum JII, in Lehinocardiwm cordatum. The origin of

ambulacral ‘“ plate-crushing,” founded on a brief survey of the

phenomenon in all groups of Hchinoids, was discussed. Mechanical

growth-pressure was regarded as the cause, with the growth of

tubercles (Lambert’s hypothesis) as a secondary and merely modi-

fying agent.

Mr. G. P, Farran presented a paper, communicated by Dr. W.

T. Cauman, F.Z.8., entitled ‘“‘ Plankton from Christmas Island,

Indian Ocean.—II. On Copepoda of the Genera Oithona and

Paroithona.” This collection, made in 1908 by Sir John Murray,

K.C.B., F.R,S., and Dr. C. W, Andrews, F.B.S., contained eleven

species of Ozthona and one of Paroithona, ov rather more than

half the known species, the total number of known species of

Oithona being eighteen and of Paroithona two, This indicated

the great richness in species of collections made in tropical waters.

rn)

Seven of the species of Oithona and the one Paroithona appeared

to be new to science. A diagnostic table of all the known species

of both genera was included in the paper.

The next Meeting of the Society. for Scientific Business will

be held on Tuesday, February 18th, 1913, at half-past Eight

o'clock p.M., when the following communications will be made :—

EXHIBITIONS AND No'rIcEs.

H. B. Prusron, F.Z.8.

Diagnoses of new Species and Varieties of Agnathous

Mollusca from Equatorial Africa.

R. Lypexxer, F.B.S., F.Z.8.

The Dwarf Buffalo of Southern Nigeria: with a Revision of

the Dwarf Buffaloes of Western Africa.

W. A. Lamborn, M.R.C.S.

A few Notes on the Habits of certain Reptiles in the Lagos

District.

(Ee mT TOT Ee ay ALE Ue Dig Dee eS OAs.

On Two British Hntomostraca belonging to the Orders

Copepoda and Ostracoda.

R. Broom, M.D., D.Se., C.M.4Z.5.

On the Gorgonopsia, Suborder of the Mammal-like

Reptiles.

The following papers have been received :—

Arruur Witey, M.A., D.Sc., F.R.S., F.Z.S.

Notes on Plankton collected across the Mouth of the St. Croix

River opposite to the Biological Station at St. Andrews, N.B.,

in July and August, 1912.

Frank E. Bepparp, D.Sc. M.A. I-RS., BZS.

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.—IX. On a New Genus of Ichthyotzniids.

Epita KE. BAMForD.

Variations in the Skeleton of the Pectoral Fins of Polypterus.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpoy,

Reeent’s Park, Lonpon, N.W.

February 11th, 1913.

No. 116.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

February 18th, 1913.

Prof. E. A. Mincury, M.A., F.R.S.,

Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

The Srcretary read a Report on the Additions made to the

Society’s Menagerie during the month of January, 1913.

Mr. E. G. Boutencrr, F.Z.8., Curator of Reptiles, exhibited

spines of a Madagascar Hedgehog (Hriculus), which had been

found by Mr, C. R. Walter in the excrement of a Boa (B, madagas-

cariensis), this being probably the first instance on record of a

Snake swallowing a spiny mammal.

The Secretary, Dr. P. Coatmers Mircuety, F.R.S., F.Z.5.,

exhibited a series of lantern-slides showing the opening and

closing of the patch of white hairs forming the dorsal stripe in a

living Tree-Hyrax (Dendrohyrax dorsalis).

Mr. H. B. Preston, F.Z.8., read a paper entitled ‘ Diagnoses

of new Species and Varieties of Agnathous Mollusca from Equa-

torial Africa.” He drew attention to the enormous field for

conchological research awaiting the student of this very fruitful

region, and stated that in many parts each range of hills

appeared to have, to a certain extent, its own special molluscan

* 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,

fauna, often characterized by certain local and peculiar phases

common not only to the species but also to the genera occurring

in that particular locality.

Mr. W. A. Lamborn, M.R.C.S., presented a paper, communi-

cated by Prof. E. B. Pouuron, F.R.S., F.Z.S., containing some

notes on the habits of certain Reptiles in the Lagos district.

It contained an account of the habits of the Lizard Agama

colonorum, especially relating to courtship, polygamous practices,

and combativeness, and of native superstitions in regard to

Chameleons. Observations were also recorded on a batch of eggs

of a Crocodile, probably Crocodilus niloticus, on their hatching,

on the behaviour of the newly-hatched young, and on the native

beliefs as to the habits of the mother crocodile.

A paper presented by Dr. R. Broom, M.D., C.M.Z.S., entitled

“On the Gorgonopsia, a suborder of the Mammal-like Reptiles,”

contained the descriptions of a new genus and two new species of

Gorgonopsids, based on well-preserved skulls discovered by

Mr. 8. H. Haughton and the Rev. J. H. Whaits. The Gorgon-

opsia were re-established as a distinct suborder of the Therapsida,

and a list of the characters distinguishing the Gorgonopsians

from the Therocephalians was given.

A second paper from Dr. Broom dealt with the South African

Rhynchocephaloid Reptile Huparkeria capensis. A detailed

account of this species was given, and its affinities with allied

forms discussed. The evidence at present seemed to show that

Euparkeria was to be regarded as a member of an order of

generalised Rhynchocephaloid Reptiles, and might be taken as

the type of a most important suborder of this group containing

the ancestors of the Dinosaurs, the Pterodactyles, and the Birds.

Mr. R. Lypexxer, F.R.S., F.Z.8., described the heads of a

male and female Dwarf Buffalo shot by Lieut. A.W. Hunt, R.N.,

in Southern Nigeria, for which the name Bos caffer hunit was

suggested. This race agrees with the Gambian B. c. planiceros

in that the adult bulls are darker than cows, but is of smaller

size, with the orange band on the throat narrower. Mr. Lydekker

also proposed the name B. c. beddingtoni for a mounted bull of a

Red Dwarf Buffalo from Ashanti, mainly on the ground that it

is cut off from the Red Congo B. c. nanus by the above-mentioned

Nigerian race.

Dr. G. Srewarpson Brapy, D.Sc., F.R.S., C.M.Z.S8.. contributed

a paper containing the descriptions of two British Entomostraca

apparently new to science—one a Diaptomus obtained abundantly

in Loch Ness many years ago but hitherto unnoticed; the other

an Ostracod, of which one specimen only was found in brackish

water in Sussex. The latter formed the type of a new genus, and

possibly also a new family.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, March 4th, 1913, at half-past Hight o’clock p.m.,

when the following communications will be made :—

EXHIBITIONS AND Notices.

Frank E. Bepparb, D.Sc., M.A., F.R.S., F.Z.S.

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.—IX. On a New Genus of Ichthyoteniids.

W. A. Cunnineton, M.A., Ph.D., F.Z.S.

Zoological Results of the Third Tanganyika Expedition

conducted by Dr. W. A. Cunnington, 1904-1905.—Report on

the Branchiura.

WitiiAm Scuavs, F.Z.S.

New Species of Rhopalocera from Costa Rica,

ArtHur Witiery, M.A., D.Sc., F.R.S., F.Z.S.

Notes on Plankton collected across the Mouth of the St. Croix

River opposite to the Biological Station at St. Andrews, New

Brunswick, in July and August, 1912.

The following paper has been received :—

Epiro EK. BAamMrorp.

Variations in the Skeleton of the Pectoral Fins of Polypterus.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZooLoeicaL Socrery or Lonpon,

Recent’s Park, Lonpon, N.W.

February 25th, 1913.

LIST OF PLATES.

1913, Part I. (pp. 1-152).

' Plate

ie i :

tr, jf Falkland Island Spiders ..-........ssesseecere erence eee?

III. 1. Barbus spurrelli. 2. Barilius macrostoma

IV. Structure of Bornean Dragonflies :

WF) -Malzostoma CoStatind, ie terice ene is s/c he nie etayshoenerstas otete atereuel

VI. 1. Myzostoma rubrofasciatum. 2,3. M. crosslandi ....

VE MG) Ost Onan ROsSuarea the ae taptNa lies Se Ceicisn hre sak ads) Ss sce) wt

VIII. 1. Myzostoma crossiandi. 2-4. M. costatwin....-. .....000s

TXGS Nestamacde bys @rane-utaligi als ctsetee les site occier eee nce sees

al Talitriator eastwood@ |....00.eseuces ereeichwere rates Sie

XII. Gills of Astacopsis and Paracheraps ....ceecccesesveeeece®

MIL. Gills of Hngeus cunicularius 2... 0scres. cece an veces see =

XIV.

Exe | Ragen PORUP Go oR co On deS POOL Seat aa Onpaiea a ees ac Ta 8

XVI. naa Wegs . .

XVEL: 22. Bngeus fossor. 23-29.) HOGiis: 2. eae eee ae es es

XVIII. 26. Engeus afinis. 27,28. H.victoriensis 2.10552. .0-2220-

XIX. 29. Engeus victoriensis. 30, 31. EL. phyllocercus..........-.

XX. 32, 33. Hngeus victoriensis. 34. EH. hemicirratulus..........

XXI. 35. Engeus hemicirratulus. 36. E. phyllocercus ...... 21.40.

XXII. 37, 88. Engeus hemicirratulus. 939-41. E. cunicularius ......

XXIII. ;

XXIV. | Lg CUS CH ILACUUC TUS Nei nka's\ Se staletate) ee ah elet\=\sielatss Jefeies) a ete &

XXYV.

109

)

NOTICE.

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,

so that the complete reference is now P. Z. 8.1913, p.... The Distribution

is as follows :—

Part I. issued in March.

nS Ophelia se June.

a) Oogl a F nea September,

Pi itl Vayins crane December.

‘ Proceedings,’ 1912, Part IV. (pp. 757-918), were published on

November 27th, 1912, ~

The Abstracts of the ‘ Proceedings,’ Nos. 114—116, are

contained in this Part.

PAPERS.

age

1. Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.—VII. On

Six Species of Tapeworms from Reptiles belonging to tne Genus Jchthyotenia (s. 1.).

By Frank E. Bepparp, D.Sec., M.A., F.R.S., F.Z.8., Prosector to the Society. (Text-

figs. 1-10.) Cece Ob oO Ose ec ete e sere eh sces Feet erase rese ee eect ce ee oe eeccoe ne eee 4

2. Some Falkland Island Spiders. By H. R. Hoae, M.A., F.Z.8. (Pls. 1. & IL) ..... Pt

8. Descriptions of three new Fishes discovered in the Gold Coast by Dr. H. G, F.

10.

Spurrety, M.A., F.Z.8. By G. A. Bounenenr, F.R.S., F.Z.8. (Pl TIL) .......... di

. On some Parasites of the Scoter Duck (demia nigra), and their Relation to the

Pearl-inducing Trematode in the Ndibie Mussel (Mytilus edulis). By UW. Lysrmr

Jameson, M.A., D.Se., Ph.D., and Witttram Nicour, M.A., D.Sc, M.D. (Text-figs.

UR ep aR BiGrinG oA Ok TD Ea ea non oiin atalino so nCOS HAA uBpocdog ad uo riaghe5 doo0 os 53

. Contributions toa Study of the Dragonfly Fauna of Borneo.—Part I. The Corduliine :

The Genus Amphicnemis: The Legion Protoneura. By FP. F. Larpuaw, M.A., F.Z.S.

(GRICE Visas aieitt Siva era's te ape aivawsceat a aitieicteainee srnaamme Seals cio mae oes meme alee occas sere estate eileen 63

. On the Structure of Bone in Fishes: a Contribution to Palzohistology. By Epwiy §.

Goopricu, M.A., F.R.8., F.Z.8., Fellow of Merton College, Oxford. (Text-figs. 13-16.) 80

. Report on the Myzostomida collected by Mr. Cyril Crossland in the Red Sea in 1905.

By Ouarres L. Boutmncer, M.A., D.Sc., F.Z.8., Lecturer on Zoology in the University

of Birmingham, (Pls. V.-VIII. and Text-figs. 17-23.) ........ ne ai arsielee ebaaale Mee 85_

. Description of an Amphipod belonging to the Family Talitridz, from the Woodbush,

Transvaal. By Paun A. Muravuen, F.Z.8, (Pls. X.& XL) sc ccesssesceceeees aoe

. The Genus Hngeus, or the Land Crayfishes of Australia. By G. W. Smru, M.A,,

Fellow of New College, Oxford, and E. H. J. Scuusrmr, M.A., D.Sc., F.Z.8., Fellow of

New College, Oxford. (Pls. XEL—XXYV,) [SWerg.. 2s ccs eve vn: esses iecreieleteieustelers 112

On some Points in the Anatomy of the Mouth-parts of the Mallophaga. By Brucu FE.

Cummines. (Text-figs. 24-32.) ..... apes shulesBus weno) vcagstecats tele SHAG HN Ooh aS oevoods eoekee

Report on the Deaths which occurred in the Zoological Gardens during 1912, together

with the Blood-Parasites found during the Year. By H. G. Punimur, F.R.S8., F.Z.8.,

Pathologist to the Society, A. eres sles sees Sue ae nee Ree nate cu iereieses San

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

| | ZOOLOGICAL SOCIETY

OF LONDON.

LOLs.

PART’ II.

CONTAINING Paces 153 to 337, witH 24 PuaTEs

AND 35 TEXT-FIGURES.

JUNE 1913. x

}

Me vf yee

Dee C

PRINTED FOR THE SOCIETY,

SOLD AT ITS HOUSE IN REGENT’S PARK.

LONDON :

ii MESSRS. LONGMANS, GREEN, AND CO

| PATERNOSTER ROW.

Si [Price Twelve Shillings.}

LIST OF CONTENTS.

1913, Part IT. (pp. 153-337).

EXHIBITIONS AND NOTICES.

The Secretary. Exhibition of a lantern-slide of an abnormally marked Donkey ........ 241

Mr. E. G. Bourencer, F.Z.8. Remarks on the Lung-Fish, Protopterus eihiopicus ...... 241

Dr. S. F. Harmer, M.A., F.R.S. Exhibition of a Hair-ball from Madagascar .......... 241

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

of Hepruary VOUS Wisse. s «6s a pbs Tenele sien eth ele o ananhc team mene cals eee Ae et

“

PAPERS.

12. Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.— VIII. On

some Species of Ichthyotenia and Ophidotenia from Ophidia. By Frank HE. Bepparp,

M.A., D.Sc., F.R.S., F.Z.S., Prosector to the Society. (Text-figs. 33-88.) .......... 153

13. The Anterior Ambulacrum of Echinocardium cordatum Penn., and the Origin of Com-

pound Plates in Echinoids. By Hersurr L. Hawkins, M.Sc., F.G.S., Lecturer in

Geology, University College, Reading. (Pl. XXVI. and Text-figs, 39-41.) .......... 169

14, Plankton from Christmas Island, Indian Ocean.—II. On Copepoda of the Genera

Oithona and Paroithona. By G. P. Farran, (Pls. XXVII.-XXXI.) ..............

15. Diagnoses of New Species and Varieties of Agnathous Mollusca from Equatorial

Africa. By H. B. Preston, F.Z.S. (Pls. XXXII.-XXXYV.) .........-..6-- see vrewod

16. Notes on the Habits of Certain Reptiles in the Lagos District. By W. A. Lamzorn,

-M.R.C.S. eeoeece Bece se ce ener eos Oe a c.0 8 0.0 .6,0,0 10 0 oe ine:s 6 600 2 ¢ 0.0 © ©4658 0) e)6/ seb! > ele eee 218

17. On the Gorgonopsia, a Suborder of the Mammal-like Reptiles. By R. Broom, M.D.,

D.Se., C.M.ZS8.. (Pls. XXXVI. GXAKV EE) seis ic ele aierels ce oe eee ee.» 225

18. On Two British Entomostraca helonging to the Orders Copepoda and Ostracoda. By

G. Srewarpsoy Brapy, M.A., LL.D., D.Sc., F.R.S., C.M.Z.S. (Pls. XXXVIII.-XL.) . 231

Contents continued on page 3 af Wrapper,

THE ZOOLOGICAL SOCIETY OF LONDON.

Turs Society was founded in 1826 by Sir Sramrorp Rarrius,

Mr. J. Sastnz, Mr. N. A. Viegors, and other eminent Naturalists,

for the advancement of Zoology and Animal Physiology, and for the

introduction of new and curious subjects of the Auimal Kingdom,

and was incorporated by Royal Charter in 1829.

Patra.

HIS MAJESTY THE KING.

COUNCIL.

HIS GRACE THE DUKE OF BEDFORD, K.G., FBS, President,

Str Joan Roszr Braprorp,

Ke CMEC MED. DSc: RRS.

Vice-President.

Ricnarp H. Burne, Ese., M.A.

Lr.-Cot. Sre R. Havenocx

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

Tae Rr. Hon. Tar Ear. or

Cromer, P.C., G.CB.,

Ge@3Mie Ge; cKe CS bn

Vice-President.

F. G. Dawrrey Drewrrr, Ese.,

M.A., M.D.

Coartes Drummonp,

Treasurer.

Tisa.,

Str Epwarp Duranp, Br., C.B.

Freperick Giniett, Esa.

Tre Lorp GLenconneEr,

F. Do Cann Gonman, Esea.,D.C.L.,

Str Watrrer Roper Lawrence,

Breen Gre Clea:

Sir Epmunp G. Loprr, Br., Vice-

President.

Ernest W. MacBripe, Ese.,

IME AG DESC) Helse aularce=

President.

Prorrssor Epwarp A. Mrncury,

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

P. Caatuers Mircnett, Esea.,

Whale, IDGSi IGN ID) IMIR Sie.

Secretary.

W. Rt. Ocitvie-Grant, Esa.

Appian D. W. Pottocg, Esa.

OxprFietp Tomas, Ese., F.R.S.

Antony H. Wriyertetp, Esa.

Henry Woopwarp, Esa., 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 the collection

of living animals, by its Library, and by its Scientific Publications.

The Office of the Society, Regent’s Park, N.W., where all com-

munications should be sent, addressed to “The Secretary,” is open

from Ten till Five, except on Saturdays, when it closes at OnE P.M.

The Library, under the superintendence of Mr. Henry G. J. Peavot,

is open daily (except Sunday) from ten a.m. till five p.m.; on

Saturdays, ten a.m. till two p.m.

The Library is closed from Good Friday to Easter Monday, and

upon all other Bank Holidays. It is also closed annually for

cleaning purposes during the whole month of September.

The Meetings of the Society for General Business are held in the

Meeting Room at the Society’s Office on the third Wednesday in

every month of the year, except in September and October, at half-

past Four o’clock p.m.

The Meetings for Scientific Business are held in the Meeting

Room at the Society’s Office fortnightly on Tuesdays, except in

July, August, September, and December and January, at half-past

Hight o'clock p.m.

The Anniversary Meeting is held on the 29th. of April, or the

nearest convenient day, at Four p.m.

The Society’s Gardens are open daily from Nine o’clock until

Sunset. Mr. R. I. Pocock, F.R.S., F.L.S., is the resident Super-

intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator

of Birds and Inspector of Works, and Mr. E. G Boulenger is

Curator of Reptiles. The Prosectorium for Anatomical and Patho-

logical work is under the charge of Mr. Frank E. Beddard, M.A.,

D.Sc., F.R.S., Prosector, assisted by Mr. H. G. Plimmer, F.R.S.,

M.R.C.S., Pathologist to the Society.

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bution of £3, due on the Ist. of January, and payable in advance,

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No person can become a Frrrow until the Admission Fee and

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Ladies or Gentlemen wishing to become Fellows of the Society

are requested to communicate with ‘‘ The Secretary.”

P. CHALMERS MITCHELL,

Secretary.

Regent’s Park, London, N.W.,

June, 1913,

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

FOR

SCIENTIFIC BUSINESS:

1913

IPronasoylns donno oe aN Wiha in ord.

Hg OCTOBER, Miccnn sean 28th.

2 UNCVEMBER seo see ilitheand atm

The Chair will be taken at half-past Inght o'clock in the Evening

precisely.

ZOOLGGICAL SOCIETY OF LONDON:

THE ZOOLOGICAL RECORD.

[ete object of the Zootogicat Recorp is to give, by means of an

annual Volume, complete lists of the Works and Publications

relating to Zoology in all its branches that have appeared during

the year preceding the issue of the Volume, together with full

information as to the points they deal with, arranged in such a

manner as to serve as an Index to the literature of Zoology in all

parts of the globe, and thus to form a repertory that will retain its

value for the Student in future years.

The ‘ Zoological Record’ haying been amalgamated with the

International Catalogue of Scientific Literature, Zoology, Volumes

from 43 onwards can now be obtained only from Messrs. Harrison

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Under the scheme of amalgamation, Fellows of the Society, and

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following.

The Society is able to supply complete sets of the Record on the

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The prices for separate volumes are as follows :—

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Inpex Zootoeicus. 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

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Record.’ London, 1902. Price to Fellows, 18s.; price to the

public, 20s., or if sold with a set of the ‘ Zoological Record,’ 10s.

Inpex Zoonoercus, No. IJ. An alphabetical list of names of

genera and subgenera proposed for use in Zoology, as recorded in

the ‘ Zoological Record,’ Vols. 38-47 inclusive (1901-1910), and

the Zoology volumes of the ‘ International Catalogue of Scientific

Literature, Annual Issues 1-10. Compiled (for the Zoological

Society of London) by Caoartes Owen Warernouss, I.8.0., and

edited by Davip Suarp, M.A., F.R.S., Editor of the ‘ Zoological

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to the public, 15s. net., or if sold with a set of the ‘ Zoological

Record,’ 10s.

Divisions of Vols. 39 to 42 of the ‘ Zoological Record’ can be

supplied by the Society, but those of Vol. 43 onwards can be had only

of Messrs. Harrison & Sons, 46 St. Martin’s Lane, W.C.

[P. T. Oo,

SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.

Divisions of the ‘ Zoological Record, Vols. 39-42, containing

the literature of the years 1902-1905, may be obtained separately

as follows :—

No?

List of abbreviations of journals, etc.

Special Records, viz. :—

I. Generai Subjects .

Il. Mammalia

III. Aves ai ges ate uleei

IY. Reptilia and Batrachia..

V. Pisces

VII. Mollusca

VIII. Brachiopoda . .

IX. Bryozoa

X. Crustacea

XVI.

XVII.

XVIII.

. Arachnida

. Myriopoda

. Insecta -.

. Echinoderma

. Vermes..

Ceelenterata ..

Spongize

Protozoa

VI. Tunicata Pree ARE | Ls Neat ce 1

COC CcCoomoeooaoeoaooeonaocoooeoaem ons

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ON OPHIDIAN TAPEWORMS. 153

PAPERS.

12. 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 November 4, 1912: Read February 4, 1913.]

VIII. On some Spsctus oF Ichthyotenia anv Ophidotenia

FROM OPHIDIA.

(Text-figures 33-38.)

INDEX.

Page

Introductoryiecscens cree ae eee a NR

Ichthyotenia gabonica, sp. n. ......... 0c. cece eee eee. 153

Ophidotenia russelli, sp. ....................--....-.... 160

TehthyGuentd Spay mrnsret eee eee eee Ld

In the present communication I lay before the Society the

result of an investigation into the anatomy of a number of

species of tapeworms which I have obtained during the last

three years from the gut of serpents, and all of which are

members of the family Ichthyoteniide.

The parasites of serpents are obviously but incompletely known

at present. The only well-known genera which have been seen

in those reptiles are Solenophorus from certain pythons *, a few

species of Jchthyotenia + from various species of serpents, Cre-

pidobothrium = from boas, Ophidotenia § from the Indian cobra,

Oochoristica || from Zamenis viridiflavus.

Ichthyotznia gabonica, sp. n.

I obtained on June 21, 1912, a number of tapeworms from

Bitis gabonica, which I refer to a new species and name as above.

I am not able to give the exact length of the worm, but the

largest piece which I measured was 150 mm. Weare safe, there-

fore, in allowing its length to be not less than 160 mm., and

probably rather more. -The scolex is intermediate in size between

that of such species as Crepidobothriwm gerrardi, which has a

particularly large scolex, and such species as Ophidotenia naie,

which has a particularly small one. The transverse diameter is

about two-thirds of a millimetre. As the greater part of the

scolex is taken up by the suckers, each of these is fully a quarter

* Bronn’s ‘ Thierreichs,’ Bd. iv. Abth. 1 B, and literature therein cited.

+ v. Linstow, “ Helminthen von Java,” Notes Leyd. Mus. xxix. 1908, p- 85.

{ Monticelli, Atti Soc. Nat. e Mat. Modena (4) i. 1899, p. 9.

§ Beddard, P. Z.S. 1913, p. 25; also Schwarz, “‘ Ichthyotenien d. Rept.,” Inaug.-

Diss. Basel, 1908. : ’

|| Zschokke, “‘ Das Genus Oochoristica,” Zeitschr. wiss. Zool. lxxxiii. 1905.

Proc. Zoo. Soc.—1913, No. XI. 1l

154 DR. F. E. BEDDARD ON

of a millimetre across. The rostellar region of the scolex is

very small in the contracted condition, as is the case with other

Ophidian Ichthyoteniids. The suckers look outwards and rather

upwards. The length of the scolex is not more than half its

breadth. Immediately after the scolex there is a neck in which

no segmentation is visible; it is as wide as the scolex, and the

body rapidly attains to its greatest width, there being thus no

long and thin anterior region.

The segments become elongated as they mature, and attain toa

length of four or five times their breadth, or even perhaps rather

more. In this species, as in others of the genus, the genital pores

alternate in position from side to side of the body, and the rela-

tive positions of the cirrus-sac and vagina also alternate.

The calcareous bodies in this species are very abundant; they

extend into the neck region, where they are very plain, in trans-

verse sections, forming a layer in the cortical parenchyma, not

very far below the subcuticular layer. They are also apparent in

the scolex. I have frequently observed in posterior segments of

the body that the centre of the calcareous bodies is deep black,

due to pigment. In sections through the scolex the four suckers

are seen to occupy nearly the whole of the area available, there

being but little space between them. The thickness of the

suckers is much greater actually and relatively than in the allied

Acanthotenia, which I have recently described *. An examina-

tion of these sections failed to show any spiny covering of the

body round or in the suckers such as is so conspicuous in Acan-

thotenia. Iam convinced that such a spiny covering is entirely

absent.

Sections through the scolex also show the slender muscular

fibres which effect the movements of the latter. These fibres do

not form bundles, but pervade singly the region lying between

the suckers. I have pointed out in a previous communication T

that the stout muscles of the suckers in the genus Acanthotenia

concentrate themselves in the neck into very marked and thick

bundles of rather thick fibres, which are continued back for a

short distance only. It appeared to me when comparing that

genus with Jchthyotenia, that Acanthotema was to be probably

distinguished from /chthyotenia by, inter alia, this presence of a

thick layer of longitudinal muscles in the neck and by the absence

of such a layer in the trunk region, this latter layer being present

in Ichthyotenia. The examination of the species which forms

the subject of the present memoir confirms that opinion; for in

Ichthyotenia gabonica I have not been able to detect a longi-

tudinal layer of fibres in the neck region; the slender fibres

already referred to which effect the movements of the suckers

do not become concentrated in the neck into a thick series of

bundles as in Acanthotenia; nor indeed could I discover any

layer at all of such muscles in this part of the body.

* P.Z.S. 1913, p. 8, text-fig. 1.

+ P.Z.S. 1913, p. 9, text-fig. 2.

OPHIDIAN TAPEWORMS. 155

In the posterior part of the body, however, (see text-fig. 33)

behind the neck a: longitudinal layer of muscles was very clear in

Text-fig. 33.

Link:

Transverse section through part of proglottid of Ichthyotenia gabonica.

lm. Longitudinal muscles. c.c. Calcareous bodies,

Ichthyotenia gabonica as in other Ichthyotenias from serpents,

including my genus and species Ophidotenia naie*, This layer

* P.Z.S, 1913, p. 25.

11*

156 " DR. F. E. BEDDARD ON

divided off the cortical. from the medullary parenchyma and lay

therefore at a distance from the outside of the body of about

one-third of the entire vertical diameter on each side ; that is to

say, the medullary parenchyma in this species is of about the

same diameter as the cortical layer. The fibres are strong and

are disposed in a layer which varies from one to three fibres

thick. They stand out so conspicuously from the general paren-

chyma of the body that it would be impossible to miss them. In

Ophidotenia this layer, although in reality plain enough, does not

strike the eye so forcibly. I am indeed reminded by their appear-

ance of the figure of the same muscles in Palaia varani given in

his account of that species by Dr. Shipley*; and it will be

remembered that I have myself ventured to consider that

Palaia may be really identical with Ichthyotenia, or at least

near to it 7.

This layer of longitudinal muscles does not form a complete

layer surrounding the medullary parenchyma. It is interrupted

at its two ends by the nerve-cord which is partly within and

partly without the medullary region. Dr. Schwarz has laid some

stress upon the fact that the transverse layer of muscular fibres

in the genus Jchthyotenia at least occasionally runs within the

medullary parenchyma, occupying the greater part of that region,

and the figure already quoted from Dr. Shipley’s memoir shows

something of the same kind in Palaia varani. There is no doubt

that in Ichthyotenia gabonica the medullary parenchyma is per-

vaded by slender muscular fibres running transversely and

scattered through it fairly uniformly, but not anywhere very

close together. I imagine that this is the same appearance that

has been seen and described by the two authors quoted. These

fibres are definitely muscular fibres, and not the usual paren-

chymal meshwork arranged in a more markedly transverse

fashion. The last point in the general structure of the body to

which I shall refer is the subcuticular layer. This layer is

several cells thick; but it is by no means so conspicuous as in

Acanthotenia and Ophidotenia and apparently in some other

species of Ichthyotenia, since in these three genera the indi-

vidual cells are much larger than in Ichthyotenia gabonica.

In more mature segments (see text-fig. 34) the layer of

muscular fibres lying between the cortical and medullary

regions is not at all defined in transverse section. Instead of

a clear cut row of fibres, a denser layer of parenchyma seems

alone to divide the cortical and medullary regions. In longi-

tudinal sections, however, this dense layer is seen to be traversed

by delicate longitudinal fibres one or two deep; these are not

apparent at all in transverse sections unless the latter happen to

be cut rather obliquely, in which case the fibres become visible.

This difference between the longitudinal muscular layer in

* “ Zoological Results. . . from New Britain, New Guinea, etc.,” by Arthur Willey,

Cambridge, pt. v. 1900, pl. lv. fig. 18.

+ PB. ZS. loc. cit.

OPHIDIAN TAPEWORMS. 157

various regions of the body of Jchthyotenia gabonica is not at all

due to the differences in the mode of preservation and staining.

The pieces of worm from which the various sections described

above were cut had been treated in precisely the same manner.

Text-fig. 34.

A series of sections of Iehthyotenia gabonica to show rudimentary uterine pores.

e. Cortical layer. e¢.d. Calcareous bodies. m. Medulla. ¢. Testis.

u. Uterus.

The medullary parenchyma also changes its character in different

regions of the body. Anteriorly it is dense thr oughout; later

the cortical region is composed of a lax parenchyma while the

158 DR. F. E. BEDDARD ON

medullary parenchyma is more dense. In the fully mature

segments towards the end of the body both the eortical and

the medullary parenchyma are equally lax in structure. This

renders it very difficult to follow the excretory tubes the calibre

of which is not greater than that of the spaces between the fibres

of the parenchyma.

This species also contrasts with some others (as is shown in

text-figs. 33 & 34) in the large and very conspicuous calcareous

corpuscles.

The ¢estes are numerous and show the usual arrangement met

with in this genus. In ripe segments I counted as many as ten

testes in one transverse row, five on either side of the median

uterus. It appeared to me that in this species the entire genera-

tive system was ripe at the same time. That is to say, the testes

were mature in segments in which the uterus was filled with

eggs; and, on the contrary, more anteriorly, where the uterus

was only represented by a slender median cord, in which a lumen

was hardly discernible, the testes were also immature. They

are at first represented by patches of nuclei in the medullary

parenchyma. Later they are in the form of sharply marked

cavities, loosely packed with the testicular cells, and when fully

mature the masses of spermatozoa nearly fill the cavities. The

sperm-duct forms the usual coil close to its opening into the

cirrus-sac. This coil appeared to me to be denser than in

some other species, and the lumen of the sperm-duct not to

be quite so wide as is often the case. The coil of the sperm-

duct and the cirrus-sac together occupy from one-third to

one-half of the diameter of a proglottid, and lie at about its

middle, sometimes rather anterior, at times rather posterior, to

the middle line.

The cirrus-sac is, as a rule, almost spherical in shape in the

mature proglottids. Its walls are thin. The cirrus itself lies

coiled up within it, and not much room is left between the cirrus

and the walls of the sac in which it lies. Everywhere the cirrus

is surrounded by a layer of glandular cells like those to which 1

have referred in Ophidotenia naie*. When the cirrus-sac is

examined in an entire proglottid mounted in glycerine, the

anterior and outside region of the sac is seen to be occupied by

the distal part of the cirrus, which is of much greater calibre

than the rest and lies almost straight. The rest of the sac is

occupied by the close coils of the narrower region of the cirrus.

In such preparations the cirrus is of a golden yellow colour, and

thus stands out conspicuously from the rest of the tissues of the

worm. I have never observed the cirrus in a state of protru-

sion; but, on the other hand, I have seen the whole sac itself

partially protruded in a way commented upon by Schwarz 7 in

other species of this genus. I found no spines upon the cirrus

anywhere. The vagina runs at first straight and parallel with

* P.Z.S. 1913, p. 29.

+ “Ichthyotenien d. Reptilien,” Inaug.-Diss. Basel, 1908. .

OPHIDIAN TAPEWORMS. 159

the cirrus-sac; in this region it is wider than posteriorly. It

then bends towards the median line of the segment and passes

straight down the middle line until near to the ovary. The rest

of the vagina, both in front of and behind the ovary, forms &

close coil. I did not find any conspicuous shell-gland ; but the

“ Schluckapparat ” was quite conspicuous. The absence of the

shell-gland, or at any rate its inconspicuous size if present, 1s to

to be accounted for, as it appears to me, by the structure of the

uterus, to which attention will be called immediately. The

ovary of Ichthyotenia gabonica is a single organ, the two wings

being continuous with each other across the median line. The

ovary is flat and lies on the ventral side of the body, touching

the border-line between the cortical and medullary parenchyma.

It extends to near to the lateral boundary of the medullary

layer. The vetelline glands show no peculiarities, and are as

usual lateral.

The uterus, however, is in some ways remarkable. When

immature it is a solid rod running throughout the body.

This is soon excavated, and even in ripe segments parts of the

uterus are still simply a narrow tube, while elsewhere it is

dilated and full of eggs. In the most fully mature segments

which I have seen the uterus seems to fill the greater part of the

medulla from side to side.

There are no very marked lateral diverticula of the uterus such

as occur in other species of Ichthyotenia, and which are so very

markedly differentiated from the median stem in my genus

Ophidotenia. But there are here and there outgrowths on each

side, which are shown in transverse sections of a ripe proglottid

by a trabecula dividing the cavity of the uterus. The walls of

the uterus where it is narrow, and as a rule though not invari-

ably without any eggs, are thick and of glandular appearance.

These walls are darkly staining and granular, and seem to consist

of a glandular epithelium which very possibly secretes the shell

of the egg. But it must be remarked that this layer of tissue

is not like the numerous stalked glandular cells which form the

cells of the diverticula of the uterus in Ophidotenia naire. The

drawing (text-fig. 34, p. 157) presents the appearance which is

shown by the uterine walls of this species. Where the uterus

is wider and full of embryos the walls are much thinner ; but

this appearance may be due simply to the stretching of those

walls and not due to any difference of real structure. When the

uterus is widened out it occupies the middle of the body and is

in close contact with the poundary-line of the medullary and

cortical regions above and below.

This is not, however, always the case: in some parts of the

same proglottid the uterus extends further towards the outside

of the body ventrally than dorsally. In sections of the uterine

tube which have a narrower calibre it is plain that that tube

occupies a ventral position, There is more medullary paren-

chyma above than below it. I have observed in such places a

160 DR. F. E. BEDDARD ON

state of affairs which is represented in text-fig. 34. It will be

there seen that the uterus gives off a slight diverticulum towards

the exterior of the body which is visible for at most three con-

secutive sections. This outgrowth, however, does not reach the

exterior, for it is plugged with cortical tissue. But one can

readily see in the sections, of which the drawings referred to

are copies, that a denser fibrous layer surrounding the uterus is

prolonged towards the exterior, in the same fashion as, but

further than, the uterine cavity. This peculiarity seems to me

to be explicable on the hypothesis that we have here either

a vestige of, or the beginning of, separate uterine pores such

as exist in the undoubtedly closely allied, if not identical, genus

Ophidotenia. I am, however, convinced that there are not in

the present species any actual pores.

The uterus in the ripest proglottids is very full of eggs which

have a narrower and thicker outer shell and a wider and thinner

inner shell. There are no external processes such as Schwarz has

figured in Ichthyotenia nattereri*. The eggs are not in any way

massed into balls such as occurs in the allied genus Acantho-

tenia y+; they lie, as it were, anyhow, but with some granular

material between them. This tends to aggregate them into a

continuous mass.

I believe this species to be different from any that have

been described. Its general size and the size of the scolex are

perhaps nearest to those of “ Tenia” racemosa, as described by

Schwarz. But the material belonging to this latter species, which

was examined by Schwarz, was not in a satisfactory state of

preservation. And, moreover, Jchthyotenia racemosa seems to

frequent South American snakes, while that which forms the

subject of the present communication is African in range. More-

over, the cirrus is unlike that of my species in not being coiled

and only pursuing an undulating course through the cirrus-sac.

The testes in J. gabonica do not appear to be so large as those of

I. racemosa. Nor are the diverticula of the uterus so well marked

as in J. racemosa.

Ophidotznia russelli, sp. n.

Of this species an example was obtained from a Russell’s Viper

(Vipera russelli) in June 1911. The general appearance of the

worm is that of a typical Ichthyotenia or Ophidotenia, which

genera do not differ to the naked eye unless it be ultimately

proved that a small scolex characterises Ophidotenia and a large,

or at any rate larger, scolex characterises Jchthyotenia. The

worm was very active when alive, and the specimen when

extended was a foot or so in length. The scolex, as already

mentioned, is very small and not more than one-half of the

width of that of /chthyotenia gabonica just described. It is of

* Loe. eit. Taf. iii. fig. 7.

+ Beddard, P. Z.S. 1913, text-figs. 6, 7, p. 20.

OPHIDIAN TAPEWORMS. 161

course unarmed, The neck is long. The posterior proglottids

are longer than broad and from 2-3 mm. wide. The generative

pores alternate, but there are often as many as four or so

consecutively on one side. The external anatomy of this species

indeed hardly differs from that of Ophidotenia naic, for which

I have recently founded the genus Ophidotenia*.

The internal structure, too, is very similar. In transverse

sections the calcareous bodies are by no means so plain as in

Ichthyotenia gabonica just described. But in pieces of the body

mounted entire in glycerine the calcareous corpuscles are quite

obvious, and appear to be restricted to the lateral regions of the

segments, being absent or very few in the median dorsal and

ventral regions. It appeared to me that the glandular sub-

cuticular layer of the present Ophidotenia does not consist of

such large cells as that of the other species of the genus. But the

arrangement of the longitudinal muscles was quite similar. It is

possible that the existence of the strong internal longitudinal

fibres in Ophidotenia in the sexual proglottids and their very

feeble development in Ichthyotenia gabonica, may prove to be

a generic distinction between these types.

T could find in this worm only a single water-vascular trunk on

each side of the body. In this the two species of Ophidotenia (af

there be two) agree; but there isa small difference to be observed

which helps to justify a separation.

In the present species the water-vascular tube lies further away

(towards the centre of the proglottid) from the vitelline strip

than in Ophidotenia naie. And this difference is even greater

than appears by a mere inspection ; for the transverse diameter of

the sections of Ophidotenia naiw was greater than that of those

of Ophidotenia russell. In the latter species I observed two and

a half to three ripe testes to lie between the water-vascular tube

and the vitelline strip, whereas there was only room for one or a

little more in Ophidotenia nae.

The reproductive organs also show some slight difference in the

present species from what I have observed in its congener. The

testes ave quite absent from the middle of the proglottids, and are »

laterally pressed up close to the strip of vitelline glands. They

seem to me to extend further towards the middle line in Ophido-

tenia naic. The cirrus-sac and the coil of the vas deferens

together reach to nearly the middle of the segment. The cirrus

is not by any means long and the coil within the cirrus-sac 1s

disposed in one or two loops only, thus contrasting with that of

Ichthyotenia gabonica described above. It appears to me that the

coiled region of the male duct lying within the cirrus-sac in

Ophidotenia naic was rather larger than in the present species.

There is no doubt that the cirrus-sac is larger in the first-named

species. In any case, the small number of the coils lying within

the cirrus-sac of the present species contrasts very markedly with

* P,Z.S, 1913, p. 25.

162 DR. F. E, BEDDARD ON

what I have observed in Jehthyotenia gabonica, and glycerine

preparations of the proglottids of the two species are very easy to

distinguish. There are no special comments to make concerning

the ovary and the female ducts in the neighbourhood of the

ovary ; they appear to agree entirely with those of Ophidotenia

naie. The oviduct widens to form the end of the vagina, and the

latter suddenly dilates at its external orifice to form a muscular

sac quite as large at the terminal section of the cirrus. It is

nearly always in front of the opening of the cirrus; I found it

posterior only in one case. I could not find a definite sphincter

muscle surrounding the terminal section of the vagina, such

as exists in Ophidotenia naiw. The preparations that I have

made of the present species which illustrate the structure of the

uterus, serve rather to increase our knowledge of this organ in

the genus Ophidotenia than to accentuate differences between the

two species of the genus.

The accompanying drawing (text-fig. 35) shows the uterus in

an incompletely mature proglottid, which is therefore not very

long in proportion to its breadth. The uterus seems to lie exactly

in the middle line and to extend from near the posterior to near

the anterior border of the proglottid. In this young proglottid

the lateral diverticula of the median stem of the uterus were only

just beginning as inconspicuous buds. At the anterior end the

uterus opens directly on to the exterior by a large and very con-

Spicuous pore, which can be easily seen by careful focussing to

have clear-cut outlines due to the cuticle. It is quite circular in

contour. It is a noteworthy fact to find one definite uterine

pore only. For the fact brings the peculiarities of this genus

Ophidotenia more into line with the Bothriocephalids and tends

to show that, as might be expected, the frequent pores of later

stages are a secondary state of affairs, and thus not inimical to

the main point of resemblance urged between this genus and the

lower tapeworms. But, although there is only one large definite

external uterine pore to be seen in this preparation, the sub-

sidiary pores, much smaller, are to some extent recognisable

prolongations of the uterus approaching to very near the surface,

if not actually opening on to it.

In the proglottid in front of and in that behind the one which

is figured in the annexed drawing and has just been described,

there is not a large anteriorly situated uterine pore. But a few

rather indistinct pores are visible, like the subsidiary ones noticed

in the case of the first proglottid to be examined. The in-

distinctness of these pores leads me to infer that they can

be temporarily closed and, perhaps, indeed they may become

permanently closed, thus approximating to the conditions that I

have described above in [chthyotenia gabonica. I have naturally

examined these pores in transverse sections. In such sections a

depression in the outer layer of the body which forms the external

part of the uterine pore is conspicuous and relatively large.

Nearer to the centre of such a depression the cuticle is seen to

OPHIDIAN TAPEWORMS. 163

cease rather rapidly, leaving an obvious discontinuity. As I

have pointed out in Ophidotenia naiw, there is in the present

species no question of an artificial rupture of the cuticle due to

Text-fig. 35.

_ut.

The upper figure represents an incompletely mature proglottid of

Ophidotenia russelli.

The lower figure is of the anterior uterine pore more highly magnified.

ov. Ovary. #. Testes. wt. Uterus, the external orifices of which are represented

black. ¢ & 9. Male and female terminal organs.

an accident in the processes of section cutting. The cuticle is

seen to become thinned toa point on both sides of such an orifice

164 DR. F, E. BEDDARD ON

viewed in transverse section. But the break in the tissues

beneath the cuticle was not in my sections coextensive with the

area of this pore.

In such sections the area of the body upon which the pores lie

does not appear to be raised above the general surface of the body.

But in the large terminal orifice of the uterus which I have

depicted in text-fig. 35, the uterine pore is clearly borne upon

the summit of an elevation. I have already pointed out that

this orifice is circular in outline, and it may be added that the

underlying soft tissues correspond to this, the discontinuity

corresponding with that of the cuticle I have just mentioned ; and

there is here an apparent contradiction—that in my transverse

sections there is no such correspondence between the areas of the

cuticular pores and the narrow canals leading thereto from

the uterus. This apparent contradiction will be reconciled by

a consideration of text-fig. 36, which represents an older pro-

glottid than that illustrated in text-fig. 35. It is considerably

longer in proportion to its breadth, and the coils of the vas

deferens gorged with sperm are plainly visible, which is not the

case in the shorter proglottid. Furthermore, the ovary has

gained greatly in bulk, as will be noticed in a comparison of the

two figures. Differences in the structure of the uterus I shall

refer to later.

It will be observed that the external uterine pores are quite

obvious on this superficial view, but that the orifices are not

always circular as has been described in the younger proglottid.

The preparation from which text-fig. 36 was drawn consisted

of three proglottids. In all of them there was an uterine pore

coinciding with the anterior termination of the uterus. The pore

was not circular but of an elongated oval form, and other slit-like

forms were observed behind this point. It follows, therefore,

that in transverse sections a given pore will occupy a considerable

number of individual sections of the series. As to the structure

of the uterus itself, it will be seen that the lateral diverticula

have grown in length as compared with the younger stage. They

are apt to be irregular in position, not being always symmetrically

paired ; indeed, the diverticula are sometimes lacking for a con-

siderable distance on one side of the median stem. The stalked

glandular cells covering the cveriena agree with those of

Ophidotenia naic.

Ichthyotenia sp. ?

Of this species a number of examples were obtained from

the Mocassin Water Viper (Ancistrodon piscivorus). ‘The living

worms reached 14 inches in length and they measured 8 to 10

inches when in spirit. The unarmed suckers were quite mobile

and independent of each other. The neck region can contract

and move like the whole body. The scolex is large and measures

in the contracted state 2 mm. in width or even rather more.

OPHIDIAN TAPEWORMS. 165

Text-fig., 36.

Ripe proglottid of Ophidotenia russelli.

P. Anterior uterine pore.

Towards posterior end of uterus three other uterine pores, also left white, are seen.

Lettering as in text-fig. 35.

Tam disposed, on account of the external characters of this worm,

to regard it as in all probability identical with Ichthyotenia

166 DR. F. E. BEDDARD ON

marenzelleri described by Barrois *, and later by Schwarz}, which

was obtained by Calmette from the same species of snaket. But,

inasmuch as there were only just indications of the reproductive

organs I am unable to write positively upon the matter, and thus

Text-fig. 37.

€

s ¢

Ss st

Transverse section through neck-region of Ichthyotenia sp.

7. Longitudinal muscles. w.v. Water-vascular tubes.

prefer not to give it a name. It is rather remarkable that in

proglottids situated 8 inches or so from the scolex, there were

merely traces of the reproductive organs—in fact, only the

* Bull. Soc. Sci. Agr. Lille, 1898.

+ “Die Ichthyotenien der Reptilien,” etc., Inaug.-Diss. Univ. Basel, 1898.

+ “ Trigonocephalus piscivorus (Erigonocephalus piscivorus).”

OPHIDIAN TAPEWORMS. 167

beginning of the formation of the ducts passing between the two

water-vascular tubes.

Text-fig. 38.

w.V

Ww. Vv,

Transverse sections through portions of proglottids of Ichthyotenia sp.

im. Medulla. w.v. Water-vascular tubes.

T have, however, a few remarks to add to Schwarz’s description

of this worm. This author does not mention the fact that the

neck region, at any rate just behind the scolex, is provided with a

very thick layer of longitudinal muscle fibres occupying the whole

of the space between the subeuticular layer and the medullary

168 ON OPHIDIAN TAPEWORMS.

parenchyma. The number of fibres in a single vertical row was

about 12; and they were not associated into bundles. The

individual fibres were, indeed, rather far apart. The section

illustrating the structure of this region of the body is represented

in text-fig. 37, and it also shows that the medullary region is

to a slight extent invaded by these longitudinal fibres. It will

be observed that we can detect in this species of Jchthyotenia an

arrangement of the muscular system like that of the Ichthyo-

teeniids of Varanus, which I follow v. Linstow in assigning to a

separate genus Acanthotenia*. The resemblance, however, is

not exact; for in dAcanthotenia the fibres are associated into

definite bundles, which is as definitely not the case in the present

species. Still there is a likeness which so far weakens the case

for the generic distinctness of Acanthotenia. And, furthermore,

I can find in the /chthyotenia under consideration no marked

layer of longitudinal fibres in the body generally, such as is

obvious, for example, in Ophidotenia russelli described above.

Indeed, Schwarz, remarks that “die innere Langsmuskulatur ist

schwach.” I do not assert that there may not be some delicate

fibres here; but there is nothing so conspicuous as is to be met

with in many other forms. The musculature therefore in this

region of the body agrees with that of Acanthotenia ; but there is

no spiny covering of the scolex. Schwarz has remarked upon the

presence in /. marenzellert, as well as in J. calmettet, of transverse

muscular fibres occupying the whole of the medullary layer.

I cannot interpret the appearance of the transverse sections of

the present species in that fashion.

The accompanying drawings (text-fig. 38, p. 167) show two

sections of this species, of which the upper one is from a region

of the body anterior to the lower one. It will be seen that in

both the medullary parenchyma differs from that of Ichthyotenta

gabonica figured abovet by the more strongly marked fibrous

looking network in the meshes of which the homogeneous ground

substance lies. Furthermore, it seemed to me that this network

was considerably more emphasized in the posterior region of the

body than in the more anterior segments. In the posterior

region, moreover, I did not always detect the water-vascular

tubes, which are quite easy to see more anteriorly as is to be

gathered from an inspection of the figure to which I have referred.

The difficulty of seeing these tubes is further evidence of the

greater thickness of the network in the more posteriorly situated

proglottids. Whether muscular fibres lie in this network I have

not been able to see; but I am of opinion that the network is

not a transverse muscular layer, but merely an exaggeration of

the network which is always visible in this situation in these and

other tapeworms, and in which may le muscular fibres inde-

pendent from it. I have already directed attention to the

presence of such muscular fibres in Jchthyotenia gabonica

described above.

* P. Z.S. 1913, p. 5. + Supra, p. 155, text-fig. 33.

WMS), Idk ASN

ID)

ECHINOCARDIUM CORDATUM.

ON THE STRUCTURE OF ECHINOIDS. 169

13. The Anterior Ambulacrum of Echinocardium cordatum

Penn.,and the Origin of Compound Plates in Echinoids.

By Herpert L. Hawkrys, M.Sc., F.G.S., Lecturer in

Geology, University College, Reading.”

[Received November 8, 1912: Read February 4, 1913. ]

(Plate XX VI.+ and Text-figures 39-41.)

INDEX.

Page

Structure of ambulacrum III. in Spatangide .................. 171

Pr of petals in Clypeaster................00.cecee vec eee eos 178

eB Ofsperionathicy indepen ee eee eee elee ears eee 178

Physiology : use of podia of amb. III. in Hehinocardium ... 176

Development of new coronal plates in Regular Echinoids

and Spatangide .............. Wan ied sa LO,

Variation in position of apical apictiann fl in Mie nioceide .. 174

I. Introduction.

During the course of research into the anatomy of the test: of

the primitive Irregular Echinoids, I had occasion to re-examine

the corresponding structures in living forms, and naturally

included among these the common British ‘“ Heart-Urchin,”

Kchinocardium cordatum Penn. It might have been thought

that, in the case of a species so long known and so frequently

examined, all the essential structural features had been deter-

mined. It was with considerable surprise, therefore, that I

found in it a character (from my point of view of first-rate

importance) that has not hitherto been certainly recognised, and

never adequately figured or described.

Lovén, with his customary fidelity, indicated some of the

complexity in ambulacrum III. of the species (Etudes sur les

Echinoidées, Pl. xii. fig. 107, and Pl. xxxix. fig. 222), but he

gave no comment on the structure, and his drawings of it are

incomplete. Beyond this casual recognition, I have ‘been unable

to find any account or figure of the very remarkable features

here described, and, indeed, most of the descriptions and drawings

of the area that I have seen are positively misleading. This is

undoubtedly due to the fact that a surface examination, however

careful, shows little or no trace of the elaborate structure present.

It was not until I accidentally discovered a new method of dis-

playing sutures that I had any suspicion of the remarkable

character of the area.

The examination of some 150 prepared specimens of various

sizes has led to considerations which have an important bearing

on the question of the origin of “ plate-crushing,” and a discussion

* Communicated by Dr. F. A. BatuEr, F.R.S., F.Z.S.

+ For explanation of the Plate see p. 181.

Proc. Zoou, Soc.—1913, No. XII. 12

170 MR. H. L. HAWKINS ON THE

of the various views held on that point follows the description of

the anterior ambulacrum of /. cordatum.

IL. Technique.

The test in ambulacrum IIT. of £. cordatum is exceedingly

thin, even in large examples being only as thick as ordinary

paper. Viewed fone the outside, ane area between the apical

system and the anterior border of the internal fasciole is covered

with an almost uniform granulation, and no trace of sutures can

be distinguished in the adapical part. From the inside a clue

can be obtained as to the existence of complexity in the structure

of the area, but it seems a general rule among Hchinoids that the

sutures shown on the inner surface of the test differ to some

degree in their disposition from those on the outside. After a

great number of experiments, I have perfected a method whereby,

as may be judged from a, SOCTIE fig. 2, the details of the

sutures, however minute, may be rendered clear.

The specimen should be well dried, and may have been stored

in a collection for many years without lessening the efficacy of the

process. It should not, however, be macerated, and the bleached

specimens of #. cordatum often found on sand-dunes rarely prove

suitable material.

When the specimen is quite dry and brittle, the test should be

split open along the ambitus, and all the sandy and organic con-

tents removed. Especial care is needed to remove all the dried-

up ampulle of the area to be examined. It is usually necessary

to moisten the specimen for this purpose. When all the radioles

and accessible organic matter have been brushed away from both

surfaces, the specimen should be thoroughly dried again. A

hurried drying at this stage (over a flame or near a fire) gives the

best results.

The part to be examined is then saturated with a staining

solution. Of all the fluids that I have used, none give clearer

results than the aniline colours supplied as cheap ink at a penny

per bottle. The red, blue, and green colours are all equally

effective, but I prefer to use the last-named, as being more

restful to the eyes, and admirably suited to the requirements of

photography. It is advisable to apply the stain with a fine

camel’s-hair brush to both the inner and outer surfaces, and

the test should be quite saturated.

When the area is again perfectly dry, the brush should ke

dipped into strong hydrochloric acid, and drawn lightly once or

twice over the outer surface. Care must, of course, be taken to

wash off all the acid (from both surfaces) very quickly. The

etching with acid should be continued until the sutures stand

out clearly. The process depends on the fact that a thin film

of organic matter exists between the plates, so that, when the

caleareous substance is partly etched away, the densely s stained

tissue in the sutures remains standing out in relief.

STRUCTURE OF ECHINOIDS. 171

The acid has no apparent effect on the green dye, and the

results show no serious sign of fading or other degeneration after

three years. (The specimen should not be kept for too long

exposed to bright sunlight.) Although the process sounds some-

what involved, it can be very quickly accomplished. I have

completely prepared fifty specimens within two hours of their

having been cleaned. The method will probably be found useful

for distinguishing sutures in other Hchinoids, but I have not

obtained such clear results when applying it to thick plates, such

as those of Hehinus. For the guidance of any who may adopt

the method I add the following particulars :—

It is inadvisable to reverse the order of procedure outlined

above-——staining should always precede etching.

The process destroys the finer surface characters of the plates,

and should not be used on rare specimens.

The etching should be continued until the plate surfaces are

almost colourless, but care must be taken that the plates are not

eaten through, or distortion of the sutures will result.

The clearness of the preparation may be increased by slightly

etching the inner surface.

The sutures should be viewed by reflected light, and will not

show in perfection unless the specimen is dry.

If the staining has not been sufficiently intense, but little

advantage is gained by applying more. ‘The specimen may often

be made useful, however, by saturation with glycerine and

examination by transmitted light.

III. Description of ambulacrum ITT, of Echinocardium

cordatum.,

The part of the ambulacrum here considered is enclosed by the

internal fasciole. In EH. cordatwm this region is depressed to

form a considerable groove passing into the anterior notch. The

surface of the area is covered with a uniformly fine granulation,

which tends to become more sparse along the perradial suture in

the anterior part. A few very small tubercles are sometimes de-

veloped on it, although some of the largest tubercles on the test

occur on the sides of the groove, which are built of interam-

bulacral plates. The shape of the groove (and also of the fasciole)

is rarely symmetrical, being normally expanded more on the right

side than on the left.

The character of the pores is strikingly different in various

parts of the area :—(i.) Near the apical system the pores are

minute, and usually uniserial in arrangement. (ii.) In the

middle section they are much larger, more or less pyriform or

transversely elongated in shape, and arranged in a very complex

order, biserially, triserially, or irregularly. They often form

broad “ poriferous zones” that occupy half the width of each half

of the ambulacrum. (The area in this section expands to its

greatest width.) It is here that the longest podia are situated.

In both the foregoing sections the pores are closely packed.

MR. H. L, HAWKINS ON THE

Text-fig. 39.

172.

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with considerable lateral expansion.

There are 67 ambulacrals in column a, and 69 in column 6.

In both columns 45 plates remain primaries.

Ambulacrum III. of Hehinocardium cordatum,

STRUCTURE OF ECHINOIDS.

Text-fig. 40.

Ambulacrum III. of 2. cordatwm, with slight lateral expansion.

There are 69 ambulacrals in each column.

In column a@ 29 plates, and in column 6 27 plates, remain primaries.

173

174 MR. H. L. HAWKINS ON THE

(iii.) Finally, the pores become almost circular in shape and well

spaced out anteriorly, progressively decreasing in size until the

fasciole is reached.

No sutures can be seen on the outer surface of the test in the

two adapical sections of the area, but some indication of them,

resulting from the inflation of the plates, can be seen from within.

By applying the method above described, they can be rendered

clearly visible from the outer view, and the following features

become manifest.

“ Plate-crushing ” is restricted to sections 1. and 11. of the area.

In those sections the plates are very low, especially in i. and the

anterior part of 11. In in. they rapidly increase in height.

In section 1. there is usually a large preponderance of primary

plates, and the type of “‘plate-crushing” found is that charac-

teristic of most Echinoids, namely, the production of demi-plates

(in contact with the adradial, but not reaching the perradial,

suture). ‘The demi-plates are developed quite irregularly.

In section ii., where the ambulacrum attains its greatest width,

the structure is far more complex. The plates in some parts

become very low, often being mere strips of calcite. Demi-plates,

occluded plates (in contact with the perradial, but not reaching

the adradial suture), and even “klasma”-plates (not reaching

either vertical suture) are developed, and a consequent displace-

ment of the pores occurs. In some rare instances the pores have

atrophied in occluded plates. The anterior part of this section

is built of relatively high plates, and the pores are of very large

size. The “ plate-crushing” here results in the formation of

occluded plates, which, however, almost reach the adradial suture.

In all of the specimens (over 150) examined, individual pecu-

liavities and irregularities, though frequent, are insufficient to

mask the general pattern. The differences met with seem,

moreover, to conform to two general conditions. These can be

roughly indicated as being (a) the relative width of the area, and

(6) the length of the area affected, measured in proportion to the

size of the specimen.

(a) There is a great variation in the degree of lateral expansion

attained by the ambulacrum in its middle section. This character

has no connection with different age, as specimens of practically

the same size may show the extremes of difference. It seems an

invariable rule that in the narrower areas the development of

occluded and ‘“‘klasma”-plates is greater than in the broader ones.

A comparison of text-figs. 39 and 40 will make this clear. The

development of these small plates can thus be ascribed directly to

lateral compression.

(6) Individuals of the same size have approximately the same

number of ambulacrals (about 69 in large specimens) between

ocular IIT. and the fasciole. The fasciole crosses the ambulacrum

from the 8th or 9th interambulacral plates on each side irre-

spective of the size of the specimen. The position of the apical

system is variable, being sometimes almost central, and at others

STRUCTURE OF ECHINOIDS. 175

considerably to the front of the centre. This results in a corre-

sponding length or shortness of the part of the ambulacrum

within the fasciole, although the number of plates present is the

same in both cases. In the shorter areas, the degree of crushing,

as indicated by the development of demi- and occluded plates, is

relatively increased. This result is obviously brought about by

the greater vertical compression in the area.

For the originals of text-figs. 39 and 40 I have selected speci-

mens of almost exactly the same size. There are the same

number of ambulacrals in both in the part of the area considered,

but their characters show the two extremes of difference. (Both

figures are drawn to the same scale directly from photographs

in an enlarging lantern.) Text fig. 39 represents a laterally-

expanded area, in which sections i. and i. occupy most of the

length up to the fasciole. Text-fig. 40 represents a comparatively

narrow area, in which section iii. occupies almost a quarter of the

region within the fasciole. It will be noticed that the degree of

plate-crushing is far greater in text-fig. 40 than in text-fig. 39,

more notably in the unexpanded adradial than in the perradial

tracts.

IV. The meaning of the Structure.

The only other Spatangoid in which I have seen a similar

structure to that just described is Heteraster oblongus from the

Lower Cretaceous. In that form the pores of ambulacrum III.

are more or less biserially arranged, and ‘“ plate-crushing,” never

approaching in intensity that of #. cordatum, is developed. I

have been able to satisfy myself that no disturbance of the serial

arrangment of the pores, and no trace of ‘“ plate-crushing,” occur

in Hehinocardium flavescens. The same remark applies to the

species of Wicraster and Hemiaster that I have seen, but there are

indications of some irregularity in the anterior ambulacrum of a

Schizaster from the London Clay in my collection.

Now in Heteraster, Schizaster, and Hchinocardium cordatum,

ambulacrum IIT. is situated in a fairly deep groove; while in

L. flavescens, Micraster, and Heniaster the area is more or less

flush with the surrounding test-surface in the adapical part. This

seems to indicate some connection between the presence of an

anterior sulcus and “ plate-crushing” in the anterior ambulacrum.

I recently (P. Z. 8. 1912, p. 464) argued that the sulcus below

the anus in many Jurassic Nucleolitidee might be ascribed to an

excessive growth of the plates there as a result of the interference

of the periproct. The same explanation seems to suffice in this

case. The number of interambulacral plates bordering on ambu-

lacrum IIT. in Z. cordatum remains constant during all the later

growth-stages, but the number of ambulacrals increases steadily

with the growing size of the individual. This indicates a propor-

tionate increase in the potential length and expanse of the

ambulacrum compared with that of the surrounding part of the

test. That would result in a sagging inwards of the over-

176 MR. H. L. HAWKINS ON THE

developed region. <A further support to this view is found in the

fact that in very young specimens of Z. cordatum (where “ plate-

crushing ” is very slightly developed), the anterior groove is

scarcely perceptible.

It appears, then, that after an early stage, the development of

new coronal plates from the oculars in Z. cordatum is arrested,

save in the case of ocular IIT., where the issuing ambulacrum is

continually growing by the addition of: new plates. The rest of

the test grows by the increase in size of the plates already

developed. Little or no resorption of coronal plates oceurs in

Spatangids, and ambulacrum IIT. of #. cordatum is no exception

to the rule. Indeed, the influence of the continuous growth of

the area seems confined by the anterior part of the internal

fasciole. In consequence, all the re-adjustment due to the con-

tinuous growth of new plates has to be carried on in the

‘“‘petaloid” region of the area. The results are: firstly, an

inward sagging of that part of the ambulacrum to form the

anterior sulcus, and secondly, mutual resorption and rearrange-

ment of the plates, causing “plate-crushing.” The additional

feature enters, as described above, that when the area is expanded

laterally the intensity of the “ plate-crushing ” is proportionately

reduced.

The physiological advantages of the character of the area in

EL. cordatum are obvious. <A far greater number of podial pores

can be compressed into the short space available than would be

possible by other means, and this results in a greater number of

prehensile podia being developed. Again, the absence of the

complexity in #. flavescens seems explicable on the grounds of

habitat. 2. flavescens does not occur between tide-marks, and

seems to be rarely, if ever, deeply buried in the sand, but

E. cordatum is often more than six inches from the surface, and

must therefore find greater difficulty in collecting food. Hence

the development of a larger number of podia in the latter

species.

The present paper is concerned more with the origin and

development of the structure than with its physiological meaning,

and I pass to a consideration of the views that have been expressed

to explain ‘ plate-crushing” in other groups of Echinoids.

V. Previous views on * Plate-crushing.”

Most of the work that has been done on ambulacral structures

has had reference to the Regular Hchinoids, among which the

various characters have been largely used for purposes of classi-

fication since the publication of Duncan’s paper in 1885 (Quart.

Journ, Geol. Soc. vol. xli. PP. 419-452). Duncan expressed the

view that ‘‘ growth-pressure” was the cause of “ plate-crushing,

and he included as a subsidiary cause of the pressure the grow th

of tubercles on the plates. Many of his figures, however, show

that the sutures separating the elements of a compound plate

STRUCTURE OF ECHINOIDS. 177

pass across the boss of a tubercle without appreciable deflection.

Loven (Etudes) expressed a similar view to that of Duncan.

A. Agassiz (Revision of the Echini) stated that compound

plates might be developed by the subdivision of original primaries,

but no other author seems to haye accepted this view.

Lambert, in 1892 (Bull. Soc. géol. France, vol. xx. pp. 38-100),

referred to plating-complexity, and ascribed it to the pressure of

new plates, not mentioning any influence of tubercle-growth.

But in 1900, the same author (Bull. Soc. Sci. Yonne, vol. Ji.

pt. 2, pp. 3-57) expressed the opinion that the chief cause of

“plate-crushing” was to be found in the expansion of the

tubercles, and that normal ‘“ growth-pressure ” exerted but a

secondary influence.

Both Duncan and Lambert considered, then, that ambulacral

** nlate-crushing” had a dual origin, the former holding that the

pressure of new plates was the main cause, the latter regarding

tubercle-growth as the chief agent.

More recently, Bather (1909, ‘ Triassic Echinoderms of Bakony’)

has suggested that the oncoming of “ plate-crushing” in the

Diademoid (Centrechinoid) ambulacrum is to be correlated with

the growth of the perignathic ambulacral processes, which restrain

the plates from passing freely to the peristome margin. ‘This

view is, in effect, a return to that of Duncan, that growth-pressure

of new plates is the main cause of the development of com-

pound ambulacrals. Jackson (1912, ‘ Phylogeny of the Echini’)

considers ‘‘ growth-pressure” alone in his discussion of ambulacral

structures.

In considering the corresponding features among the Holecty-

poida, I have lately taken the same view (Geol. Mag. 1910 &

1911), and, as the structures above described in Hehinocardium

cordatum seem to have a bearing on the problem, I propose to

give a discussion of the subject.

VI. The Origin of “ Plate-crushing.”

The ambulacra contain compound plates in all groups of Regular

Kchinoids except the Cidaroida. Excluding the highly specialised

and aberrant Perischoechinoida, the Regular Echinoids may be

separated into two sections, in one of which (the Cidaroida)

resorption of old plates at the peristome takes place freely and

continuously, so that the proportionate diameter of the peristome

remains constant throughout growth; and in the other (the

Centrechinoida) the resorption does not keep pace with the

growth of the corona, so that the peristome of an adult is pro-

portionately smaller than that of a young form. Im all the

Cidaroida, and almost all the Centrechinoida, the growth of new

plates at the edge of the oculars is continuous and regular. Thus

while the Cidaroid ambulacrals can pass freely down the full

length of their area, those of the Centrechinoid are to some degree

hindered. The result must obviously be a compression of the

earlier ambulacrals against the obstacle (e.g., Henvipedina, and,

178 MR. H. L. HAWKINS ON THE

for the Holectypoida, Pygaster). The fact that the compression

first shows itself near the peristome is significant; for, to my

mind, it strongly supports the view that it is to the trans-

ference of the chief perignathiec structures from the inter-

ambulacra to the ambulacra that the structural differences

between Cidaroids and Centrechinoids are to betraced. Gradually,

both in ontogeny and phylogeny, the region of compression

extends further from the point of its origin, until in many forms

(e. g., Hchinus) it reaches practically to the ocular margin.

From the opposite standpoint, it is true that in the Cidaroida

alone the ambulacra support mere granules, while every stage of

complexity among the Centrechinoida corresponds with a pro-

gressive development of large tubercles. And indeed, there is

no direct evidence for either side to be derived from a study of

Regular Kchinoids, so closely do “ plate-crushing ” and tubercle-

growth coincide in their appearance.

A study of the ambulacral structures of the Irregular Echinoids

has, however, convinced me that the “‘ srowth-pressure” of new

plates is, for them at least, the essential and original cause of

“ plate-crushing.”

Among the Holectypoida compound plates (on the Centr-

echinoid pattern) are developed to an ever-increasing degree as

the group is traced from the Lias to the Chalk. With few excep-

tions, the amount of resorption of old plates at the peristome seems

to decrease in corresponding order; and finally, the size, both

relative and actual, of the tubercles steadily diminishes. In

Pygaster, to take a typical genus, an interesting feature can be

noticed in this connection. On the adoral surface the tubercles

become large, and their areole often cross the sutures between

the ambulacral plates. The very slight degree in which the

development of the tubercles affects the arrangement of the

plates can be estimated by an examination of text-fig. 41. The

triple arrangement of the ambulacral plating of Pygaster, and

presumably of the other Holectypoida, must then owe its

development to some more fundamental cause than that of

tubercle-growth.

In many of the larger Clypeastroida (e. g., Clypeaster), the

petaloid parts of the ambulacra are built up of alternate

primaries and demi-plates. he tuberculation differs in no

respect from that of the extra-petaloid regions of the test. Here

the closing round of the end of the petal affords a new point of

resistance to the downward passage of plates, and, as a result,

a secondary type of ‘‘ plate-crushing ” is developed.

In the Nucleolitidee and “ Cassidulide,” although most of the

ambulacrum is built up of primaries, there is, near the peristome,

a phyllode or ‘‘hypophyllode,” composed of demi- or occluded

plates. There is no definite difference in the coarseness or

arrangement of the tubercles in this region, and the structure is,

as I have previously shown (Geol. Mag. 1911), cognate with that

of the Holectypoida or Centrechinoida,

STRUCTURE OF ECHINOIDS. 179

In all the groups mentioned above, little or

the growth of new coronal pl

the life of the individual.

no interruption of

ates from the oculars occurs during

Among the Spatangide, the most

Text-fig. 41,

Adoral part of ambulacrum of Pygaster semisulcatus, showing the slight

influence of tuberculation on the position of sutures.

highly specialised of moder

n Echinoids, the full complement of

plates is early attained, and,

for the most part, subsequent growth

180 ON THE STRUCTURE OF ECHINOIDS.

is the result of enlargement of the plates already developed,

rather than of the addition of new ones. In many genera a

considerable development of large tubercles occurs (though rarely

on the ambulacra), and no distortion of the sutures follows such

a secondary development. But in Yehinocardium cordatwm (and

probably in Heteraster oblongus), ambulacrum III. continues to

develop in the more primitive Echinoid way throughout lie.

And here, with an exceptional character in the manner of growth,

we find a corresponding exception in the structure. Practically

no tubercles, and no granules of considerable size, occur on the

area, but the ‘“‘ plate-crushing ” is carried to a degree of complexity

surpassing that of any other Hchinoid.

It is important to remember that in the Regularia, Holecty-

poida, and Nucleolitidee, the ‘“ plate-crushing,” when it exists, is

always most strongly developed towards the peristome. More-

over, in most members of these groups the arrangement of the

compound plates is founded on a regularly three-fold plan. In

the Clypeastroida and Spatangide, when “ plate-crushing ” occurs,

it is restricted to some part of the adapical surface, and there

seems to be no definite pattern in its arrangement.

It seems safe to conclude that the structures found in the

last-named sections have no phylogenetic connection with those

of the former groups, and a reason for their development must be

sought. From the discussion given above it seems clear that the

only feasible explanation lies in the assumption that the high

specialisation of the petaloid parts of Clypeastroid and Spatangid

ambulacra precludes the passage of plates from those regions to

lower positions. The ‘“ plate-crushing ” is confined to the petals,

and the remainder of the areas have, in large measure, lost their

distinctively ambulacral characters as regards the plating.

For the Irregular Echinoids, mechanical compression due to

the growth of new plates seems the only available cause of the

development of compound plates. To my mind it affords the most

reasonable explanation of similar structures among the Regularia.

The development of a tubercle is a superficial phenomenon,

having no visible influence on the texture of the plate-substance

in the deeper layers. It seems likely that the lateral expansion

of a tubercle may cause loca} distortion of sutures at the outer

surface (although in many cases it obviously has no such influence),

but, at the most, this effect would be local and secondary.

VIL. Summary.

A new method for exposing sutures in dried specimens of

Kehinoids is described. The method combines the processes

of staining and etching; depends on the presence of organic

tissue between the plates; and gives a fairly lasting result easily

photographed.

The complex structure of the anterior ambulacrum of Hehino-

cardium cordatum is described in detail for the first time, and a

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ON COPEPODA FROM CHRISTMAS ISLAND. 181

corresponding development in Heteraster oblongus and Schizaster

is indicated.

A general description of the growth of compound ambulacrals

among the great groups of HKchinoids is given; and, as a result

of the facts brought forward, it is argued that ‘‘ plate-crushing ”

is due solely to the mechanical ‘“‘ growth-pressure ” caused by the

development of new plates ; while the appearance of tuberculation

corresponding in arrangement with the ambulacral elements is

considered to be an after-effect of their fundamental characters,

and to modify very slightly, if at all, the structure of the areas.

EXPLANATION OF PLATE XXVI.

Fig. 1. Denuded test of Hchinocardium cordatum, showing the relative smoothness,

and the crowding of the pores, of ambulacrum IIT.

Fig. 2. The same specimen, after staining and etching, showing the plate sutures

to} : 1 : to} top) co} 5)

particularly in ambulacrum III.

14. Plankton from Christmas Island, Indian Ocean.—II.

On Copepoda of the Genera Oithona and Paroithona.

By G. P. Farran.*

[Received November 5, 1912: Read February 4, 1913.]

(Plates XX VII.-_XXXI.7)

INDEX.

GEOGRAPHICAL ZooOLoGy. Page

Oithona, spp. of, from Christmas I., Indian Ocean ...... 182

Paroithona, sp. of, <*) 5 SLA on cect 182

SYSTEMATIC.

Oithona vivida, sp. n., description and figs. ............... 183

Oithona decipiens, sp.n., 5, 5 Se nant ters OA

Oithona fallax, sp. n., 5 * 185

Oithona attenuata, sp.N., 5, TOM al fess hageocu core 187

Oithona simplex, sp. 0., Ms aie) hy eeoeuyea eae 187

Oithona oculata, sp. 0., BS PS BESS 2 Can aue al eel dots)

Paroithona pulla, sp. n., m ee LO,

Oithona pelagica, Farran, synonym of O. setigera eee 183

Oithona tropica, Wolfenden, synonym of O. setigera... 183

Oithona setigera, Dana, discussion of synonymy of...... 182

Oithona, diagnostic table of spp. of...................c0.00 0 191

Paroithona, diagnostic table of spp. of ..................055 192

In a previous paper published in the Proceedings of this

Society (1911, pp. 282-296), an account was given of the species

of Coryceus and Corycella found in a small collection of Plankton

made in 1908 by Sir John Murray, K.C.B., F.R.S., and Dr. C. W.

Andrews, F.R.S., at Christmas Island, in the Indian Ocean, and

presented to the British Museum by Sir J. Murray. The

following account deals with the species of Oithona and Par-

oithona found in the same collection.

* Communicated by Dr. W. T. Catman, F.Z.S.

+ Fcr explanation of the Plates see p. 193.

182 MR. G. P. FARRAN ON

Of the genus Oithona eleven species were represented. Five

of these,

O. plumifera, ; O. robusta,

O. setigera, O. nana,

O. linearis,

are already known. The remaining six appear to be unknown

and are described below under the following names :—

O. vivida, O. attenuata,

O. decipiens, O. simplex,

O. fallax, O. oculata.

The closely allied genus Paroithona, previously known from a

single species in the N.K. Atlantic, is represented by a new

species, P. pulla.

The fact that twelve species, out of the twenty contained

in the two genera, were present will serve to illustrate the

remarkable richness of the collection.

The number of gatherings in the collection was eight. Three

of these were taken with nets of larger mesh than the rest, and

contained only those species which exceeded 1 mm. in length.

The smaller species, including all the new forms, were found

exclusively in the other five.

Notes on the Species.

OITHONA SETIGERA Dana.

Found in five gatherings and was, next to O. plumifera, the

commonest species of Oithona.

This species was described by Dana (8) in 1849, from specimens

taken in the Pacific, and Guiesbrecht (13) gave a complete

diagnosis of it also from Pacific specimens. Most of the Christ-

mas Island specimens could be recognised by their large size

(16 to 1-7 mm., with a few as large as 1°9 mm.), and by the

possession of well-marked clavate sete, usually coloured red, on

the outer edges of the second basal joints of the swimming-feet.

In a few specimens, however, these sete were long and slender

and did not show signs of thickening. ‘These specimens agree

with O. pelagica which I described (11) from the west coast of

Treland, and also with O. tropica described by Dr. Wolfenden (17)

from a single specimen from the Maldives. Other specimens

showed a good deal of variation in the amount of the thickening,

and in some instances the sete on the fifth pair of feet were

thickened and coloured.

As regards O. tropica, Wolfenden states that “the imner

marginal bristles of the 2nd basals are not swollen as in O. seti-

gera.” In this sentence the word “inner” is evidently a clerical

error for outer. The proportional lengths of the joints of the

Ist antenne, as given for O. tropica, differ from those found in

O. setigera, but the difference can be explained by supposing the

3rd and 4th and the 6th and 7th joints, the segmentation

COPEPODA FROM CHRISTMAS ISLAND. 183

between which is imperfect, to be united and the 8th and 10th

joints each to have been divided into two.

It seems, then, that O. pelagica and O. tropica cannot be main-

tained as species distinct from O. setigera, but the fact is worth

noting that, in the N.E. Atlantic, O. setigera, whether the cause

be racial or environmental, is not found with thickened setze,

while in tropical regions these setze are almost always present.

In addition to the large form mentioned above there occurred

a very few specimens of a form measuring from 1:20 to 1:26 mm.

in length. These are the same size as, and, at first sight, look very

like O. plumifera, but examination of the appendages shows that

they agree with O. setigera. The sete of the basal joints of the

swimming-feet are not thickened. There did not appear to be

any specimens of intermediate size connecting the larger and

smaller forms.

Distribution. North Atlantic, Gulf of Guinea, Mediterranean,

Gulf of Suez, Indian Ocean, Pacifie Ocean.

OITHONA PLUMIFERA Baird.

Present in all the gatherings and very common in most

of them.

Almost all the specimens bore tufted plumose outer-edge setz

on the 2nd basal joints of the first three pair of swimming-feet,

and in the few cases where they were absent it seemed probable

that the plumose tip of the seta had been broken off. The

amount of feathering varied in different specimens. In most

cases it was opaque and of a bright red colour; in others it was

transparent and colourless and difficult to observe. In the 1st

maxilla there was no seta on the 2nd inner lobe and the seta

on the endopodite was very minute. The endopodite of the

mandible bore only three sete. The size of the specimens varied

from 1:20 to 1:37 mm.

Distribution. Mediterranean, Gulf of Guinea, Red Sea, Arabian

Sea, Indian Ocean, Pacific Ocean, Gulf of California, off Cape of

Good Hope.

OITHONA LINEARIS Giesbrecht.

This easily recognised species was present in six gatherings,

being almost as common as O. setigera.

Distribution. Red Sea, Indian Ocean, Pacific Ocean.

OITHONA VIVIDA, sp.n. (Pl. XXVII. figs. 1-8.)

Female (fig. 4).—Length -68—-74 mm.; cephalothorax °37 mm.;

abdomen *34 mm.; proportional lengths of abdominal segments

and furea 15:40: 22:18:17:15.

Rostrum (fig. 5) sharp-pointed and produced anteriorly, as in

O. plumifera and O. setigera.

Ist antenna (fig. 1) very slender, reaching to the genital

openings ; proportional lengths of antennal joints :

IL 3 4 6 6 F938

11 9 8+3+10 17 53 114+23 5 9 10°

184 MR. G. P, FARRAN ON

There are two imperfect jointings close together in the centre of

the third joint and another in the sixth; a row of minute

spinules runs along the lower edge of the fourth and fifth joints.

2nd antenna large, robust, of the usual form.

Mandible (fig. 2) with very slender 2nd basal; endopodite with

four small sete, the two central ones being the shortest ; exopodite

with four large feathered sete and one slender one.

Ist maxilla (fig. 3).—I1st inner lobe well developed with the

usual setz; 2nd inner lobe with one fine seta; 3rd inner lobe

with three spines; endopodite comparatively large with three

very small sete; exopodite with four sete.

2nd maxilla and maxillipede of the usual structure but com-

paratively short.

lst foot (fig. 6): exopodite with 1.1.3 outer edge spines and

1.1.4 inner edge setz, the seta on the inner edge of the 1st joint

being very minute, terminal spine long and slender, longer than

the exopodite ; endopodite with 0.1.4 inner edge and 0.0.1

outer edge setze.

2nd foot (fig. 7): exopodite with 1.1.3 outer edge spines and

1.1.4 inner edge sete, seta on inner edge of Ist, joint very

minute, terminal spine longer than the exopodite and broader

than in the Ist foot; endopodite with 1.2.4 inner edge and

0.0.1 outer edge sete.

3rd foot (fig. 8): exopodite with 1.1.2 outer edge spines, the

most distal very large, the rest very minute, and ‘1.1.4 inner

edge sete, terminal spine much longer than the exopodite;

endopodite with 1.2.4 inner edge and 0.0.1 outer edge sete.

4th foot rather small: exopodite appears to have 1.1.3 or

1.0.3 very minute slender outer edge spines and 1.1.4 inner

edge setee ; endopodite with 1.2.3 outer edge and 0.0.1 inner

edge setze.

5th foot of usual form.

The possession of a pointed rostrum and of a large number of

outer edge spines, though reduced in size, on the swimming-feet,

and also the presence of a large endopodite on the Ist maxilla,

places this species near to O. robusta, but the 1st antenna is of a

different type and seems to be peculiar in not having a spine on

the distal end of the 3rd (primitively 12th) joint. As O. robusta

is the least specialised of the rostrate Oithonas, so O. vivida seems

to have taken the first step towards the reduction of the spines

on the swimming-feet which is found in the other rostrate

species.

Four specimens were found in one gathering.

OrrHONA ROBUSTA Giesbrecht.

Found in four gatherings, a few specimens in each.

Distribution. Indian Ocean and Tropical Pacific.

OITHONA DECIPIENS, sp. n. (Pl. XXVIII. figs. 4-11.)

Female (fig. 7).— Length 56-62 mm.; cephalothorax *34 mm. ;

COPEPODA FROM ‘CHRISTMAS ISLAND. 185

abdomen (ineluding Th. 5) -28 mm., proportional lengths of

abdominal segments and furca 12:26:12:10:9:8. Rostrum

(fig. 4) pointed, not visible in dorsal view, similar to that of

O. similis but more hooked at the tip. Form of body slender.

lst antenna just reaches to the genital openings ; proportional

lengths of joints :

a 2 3 4, Ba Ge “Se oe

(16+16) (8+10+13412) (14414) (144+15+15+16) 12 28 10 12 14’

the first four joints being subdivided by imperfect jointing in the

proportions indicated ; there is a sinall spine on the end of the

2nd joint.

2nd antenna very small.

Mandible (fig. 5): endopodite small with two set, exopodite

with five sete. ;

1st maxilla (fig. 6) small, endopodite without sete.

2nd maxilla and maxillipede comparatively short and weal

but of usual form.

Ist foot (fig. 8): exopodite three-jointed with 1.0.2 outer

edge spines and 0.0.4 inner edge sete; endopodite two-jointed

with 1.5 inner edge setz and | outer edge seta.

2nd foot (fig. 9): exopodite with 1.1.2 outer edge spines and

0.1.5 inner edge sete; endopodite with 1.2.4 inner edge and

0.0.1 outer edge sete ; an inner edge seta was found on the Ist

basal of the lst and 2nd feet, but not of the 3rd and 4th, an outer

edge seta was present on the 2nd basals of all the swimming feet.

3rd foot (fig. 10): exopodite with 1.0.1 outer edge spines and

0.1.5 inner edge sete; endopodite with 1.2. 4 inner edge and

0.0.1 outer edge sete.

Ath foot (fig. 11): exopodite without outer. edge spines all

with 0.1.5 inner edge sete ; endopodite with 1. 2, .3 inner edge

and 0.0.1 outer edge Sece

5th foot of usual foot

This species resembles O. similis so closely that it is almost

impossible to distinguish it without examination of the spines on

the swimming-feet. It differs from that species in the absence

of-a spine on the 2nd joint of the exopodite of the Ist foot and

in the presence of a spine on the corresponding joint of the

2nd foot. It also differs in not having an outer edge spine on

the 3rd joint of the exopodite of the 4th foot, unless that spine

may have been accidentally knocked off in all the specimens

examined, as sometimes happens in the case of O. similis, and in

having a well-developed spine on the centre of the outer margin

of the 3rd joint of the exopodite of the 2nd foot, ‘which in

O. similis is only: represented by a small tooth.

About thirty-five specimens were found in one gathering.

OITHONA FALLAX, sp.n. (Pl. XXVIL. figs. 9-12; Pl. ORV ALL.

figs. 1-3.) |

toy™

Female (Pl. XXVIII. fig. 3).—Length -88- 94 mm.; cephalo-

Proc. Zoot, Soc. IGS, Wao. LONE. I3

186 MR. G. P. FARRAN ON

thorax 48 mm.; abdomen (including Th. 5) -42 mm. puma

lengths of abdominal segments and furca 9: 20:9: 9:8:6.

Rostran (etl, XXVIII. fig. 1) pointed, not visible in dorsal

view, similar to that of O. similis and O. decipiens.

Ist antenna reaches to the beginning of the genital segment ;

proportional lengths of joints :

1 2 3 4 5 6 (ee). Oy.

(24428) (4412) (16416) 40 (20+20+20+4+22) (16438) 14 16 20°

the first three and the 5th and 6th joints being subdivided by

imperfect jointing in the proportions indicated.

2nd antenna slightly larger comparatively than in O. decipiens.

Mandible (Pl. XX VIII. fig. 2): endopodite with four sete.

Ist maxilla: exopodite with three moderately strong terminal

setee, endopodite with one small seta.

2nd maxilla and maxillipede of the usual type.

Ist foot (PI. XX VII. fig. 9): exopodite with 1.1.2 outer edge

spines and 0.1.4 inner edge sete, terminal spine shorter than

the exopodite ; endopodite two-jointed with 1 .5 inner edge and

1 outer edge sete.

2nd foot (Pl. XX VII. fig. 10): exopodite with 1.0.2 outer

edge spines and 1.1.5 inner edge sete ; the inner edge seta on

the Ist joint 1s small; terminal spine approximately equal to the

exopodite ; there is a small tooth on the outer edge of the 3rd

joint of the exopodite proximal to the first spine; endopodite ~

with 1.2.4 inner edge and 1 outer edge sete.

3rd foot (Pl. XX VII. fig. 11): exopodite with 1.0.1 outer

edge spines and 1.1.5 inner edge seta, inner edge seta on Ist

joint small; there is a small tooth in the middle of the outer

edge of the 3rd joint; terminal spine slightly longer than the

exopodite, —

Ath foot (Pl X XVII. fig. 12): exopodite with 0.0.0 outer

edge spines and 1.1.5 inner edge sete, inner edge seta on

1st joint small; terminal spine slightly longer than the exopodite ;

endopodite with 1.2.3 inner edge and 1 outer edge sete.

There is a well-developed outer edge seta on the 2nd basals of

all the swimming-feet and an inner edge seta on the Ist basals.

5th feet of usual form.

This species, like O. decipiens, can with difficulty be dis-

tinguished from O. similis without examination of the swimming-

feet. The shorter terminal spines and the presence of two outer

edge spines on the 3rd joint of the exopodite of the 2nd foot and

of a small inner edge seta on the Ist joints of the 2nd to 4th

feet are the distinguishing characters which can be most easily

observed.

Four specimens were found in three gatherings.

OITHONA NANA Giesbrecht.

Occurred in five gatherings, but in two only was it numerous.

Distribution. N. Atlantic, Mediterranean, Red Sea, Arabian .

Sea, Indian Ocean, Gulf of Califor nia, aes: of Good Hope.

COPEPODA FROM CHRISTMAS ISLAND. 187

OITHONA ATTENUATA, sp. n. (Pl. XXX. figs. 3-7.)

Female (fig. 5).—Length *88 mm.; cephalothorax -4 mm. ;

abdomen (including Th. 5) ‘48 mm., proportional lengths of

abdominal segments and furea 16:8:8:6°5:8; furcal rami long

and narrow, arrangement of sete as in O. nana. Front of

cephalon (fig. 3) slightly produced but not pointed.

lst antenna (fig. 4) reaches to the beginning of the genital

segment ; proportional lengths of joints :

lO 4616 7 BOO me

16 23) 9) 19 31 1010 8 8) 199

Small spine on the end of the 3rd joint.

2nd antenna comparatively large, of the usual structure.

Mandible: 2nd basal with one stout terminal seta; endopodite

large, rounded, with four sete ; exopodite with five sete.

lst maxilla: structure not made out.

2nd maxilla and maxillipede as in O. nana.

Swimming-feet (figs. 6, 7) with three-jointed exopodites and

endopodites ; exopodites of Ist to 3rd feet with 1.1.3 outer edge

spines; exopodite of 4th foot with 1.1.2 outer edge spines ;

terminal spines of 2nd and 3rd feet shorter than the exopodites ;

short strong seta on the inner edge of the Ist joint of the ex-

opodites of the 2nd and drd feet, not seen on the Ist foot.

5th feet of the usual form.

This species is closely allied to O. nana and differs mainly in

the long drawn-out abdomen and furea and in the longer Ist

antenna.

A single specimen was found in the collection.

OITHONA SIMPLEX, sp. n. (Pl. XXIX. figs, 10-14; Pl. XXX,

figs. 1, 2.

Female (Pi. X XIX. fig. 11).—Length -38-:42 mm. ; cephalo-

thorax *24--26 mm., abdomen (including Th, 5) :14-16 mm,

(Pl. XXIX. fig. 12), proportional lengths of abdominal segments

and furea 7:21:9:7:3:7. Rostrum absent, front of cephalon

(Pl. X XIX. fig. 10) rounded as in O. nana.

lst antenna (Pl. XXX. fig. 1) reaches to the beginning of the

2nd thoracic segment; proportional lengths of joints:

BS By BS BO. NO Ab

Q'S OTS °S ALS TL BSS

2nd antenna rather short and stout.

Mandible, 1st maxilla, 2nd maxilla, and maxillipede as in

O. nana.,

Ist foot (Pl. XXX. fig. 2): exopodite three-jointed with 1.1.3

outer edge spines and 0.1.4 inner edge sete, terminal spine longer

than the exopodite ; endopodite two-jointed with 0.5 inner edge

and 0.1 outer edge setz ; process of the 2nd basal between the

exopodite and endopodite rather large and acute.

13*

188 ro. MR. -G.P, FARRAN ON

Ond foot: éxopodite three-jointed with 1.1.3 outer edge spines

and 1.1.5 inner edge sete, inner edge seta on Ist joint very

small, terminal spine shorter than the exopodite; endopodite

three-jointed with 1.2.4 inner edge and 0.0.1 outer edge setze ;

outer edge seta present on 2nd basal, not seen on 3rd and 4th

feet.

: Brd foot (Pl. XXIX. fig. 13) similar to 2nd except for absence

of outer edge seta on 2nd basal. a

4th foot (Pl. X XIX. fig. 14): exopodite with 1.1.3 outer edge

spines and 1.2.5 inner edge sete; the most distal outer edge

spine is well developed, the others being small and slender;

inner edge seta on first joint very small; terminal spine shorter

than the exopodite; endopodite with 1.2.3 inner edge and

0.0.1 outer edge sete.

5th feet very small but appear to be of the usual form with

one terminal and one basal seta.

Occurred in two gatherings, about twenty specimens.

- Inthese gatherings it could be recognised by its small size and

the shortness of the anal segment, less than the furca. .

. OIFTHONA OCULATA, sp. n. (PI. XXX. figs. 8,9; Pl. XXXT.

figs. 2-9.)

Female (Pl. XX XT. figs. 2, 3).—Length -9 mm.; cephalothorax

‘54 mm.; abdomen (including Th. 5) -36 mm.; proportional lengths

of abdominal segments and furca 7:18:10:9:8:83 fureal rami

about two and a-half times as long as wide, with an outer edge

seta situated in the middle of the outer margin. Rostruin absent,

cephalon slightly produced into a rostral prominence in front of

the antenne. The cephalic ganglia form a large bilobed mass, in

front of which are situated, in the rostral prominence, a pair of

small clear refractive areas.

lst antenna (Pl. XX XI. fig. 4) reaches to the 2nd thoracic

segment; proportional lengths of joints:

1 2 is Aa OTe Seog O EELS

LIEGE Gan Ae ele 7 AeTo Menno IG)

3rd joint with an imperfect jointing near its proxiinal end,

Ath joint. with a small spine distally on its upper margin.

2nd antenna of the usual form.

Mandible (Pl. XXXI. fig. 5): 2nd. basal with two terminal

(inner edge) spines and a small seta on its lower (inner) margin

opposite the endopodite ; endopodite sloped outwards, with three

terminal and two inner edge sete: exopodite with two terminal

and three inner edge sete. oyh os

Ist maxilla (Pl. XX XI. fig. 6): 1st inner lobe well developed,

of the usual form, 2nd inner lobe with one strong seta, 3rd inner

lobe with three terminal and one lateral sete ; endopodite mode-

rately large, bent outwards, with two lateral and two terminal

sete; exopodite large, with four terminal sete, the outermost

being slightly the largest. 73a OS Byte eae

COPEPODA FROM CHRISTMAS ISLAND. 189

2nd maxilla and maxillipede (Pl.. XXX. fig. 8) of. the usial

form but rather short.

Ist foot (Pl. XX XI. fig. 7): exopodite with 1.1.3 outer edge

spines and ]1.1.4 inner edge setw, the inner edge seta on the

Ist joint being very small; termina] spine shorter than the

exopodite ; endopodite with 1.1.4 inner edge and 0.0.1 outer

edge sete; there is a distinct tooth between the terminal and

the outer edge seta; inner edge seta on the 2nd basal longer

than the first two joints of the endopodite.

2nd and 3rd feet (Pl. XXXI. figs. 8,9) with 1.1.3 outer

edge spines and 1.1.5 inner edge sete, the inner edge seta on

the 1st joint being very small; terminal spine of 2nd foot a little

longer than the exopodite and of 3rd foot about equal to it; in

both cases it is considerably longer than the endopodite ; inner

edge seta on Ist basal of 3rd foot and outer edge seta on

2nd basal, both these setee being absent from the 2nd foot.

Ath foot (Pl. XXX. fig. 9): exopedite with 1.1.2 outer edge

spines and 1.1.5 inner edge sete, the inner edge seta on the

1st joint being very small, terminal spine slightly longer than

the exopodite ; endopodite with 1.2.3 inner edge and 0.0.1

outer edge sete ; Ist basal with inner edge seta, 2nd basal with

outer edge seta.

5th foot small, with two terminal sete and one at its base.

This species is closely allied to O. rigida, differing mainly from

it in the greater length of the terminal spines of the swimming-

feet. In O. rigida, as figured by Giesbrecht (14) and also by

Cleve (7), these spines are shorter than the terminal joints of

the exopodites. Another difference is the comparatively large

size of the inner edge sete on the Ist joints of the exopodites in

O. rigida. In O. oculata they are very small. In size and appear-

ance and in the proportional lengths of the abdominal segments

and antennal joints the two species agree very closely.

Common in one gathering, a single specimen in another.

Genus PaRoITHONA.

This genus was described in 1908 (10) from specimens taken

in deep water off the west coast of Ireland, only one species

having been met with until the Christmas Island collections

yielded a second representative. The genus is closely allied to

Oithona, and is distinguished mainly by having a two-jointed

endopodite on all the swimming-feet and by the form of the

mandible palp. It is questionable whether the differences which

separate it from some species of the genus Oithorna are airy

greater than those which separate O. nana from such species as

O. plumifera and O. setigera, but the occurrence of a second

species, closely resembling that first described, is in favour of the

retention of the genus.

190 MR. G. P. FARRAN ON

PAROITHONA PULLA, Sp.n. (PI. X XIX. figs. 1-9; Pl. XXXT.

figs)

- Female (Pl. XXIX. fig. 1)—Length -40—-43 mm.; cephalo-

thorax ‘23 mm.; abdomen (Pl. X XIX. fig. 3) (including Th. 5)

"19 mm., proportional lengths of abdominal segments and

furea 8:20:9:11:6:6. Rostrum absent, front of cephalon

(Pl. XXXI. fig. 1) rounded as in O. hebes. Furca with the

two longest sete superposed on the inner half of the terminal

margin.

1st antenna (Pl. X XIX. fig. 2) reaches to the end of the 2nd

thoracic segment; proportional lengths of joints :

LC LN Gee) NO UT

GOs 7f Da 7 ae 276:

The 3rd joint is divided into two by a partial jointing, and there

appears to be a small spine on the end of the 4th joint.

2nd antenna (Pl. X XIX. fig. 5) rather small and slender, two-

jointed.

Mandible (Pl. X XIX. fig. 4): second basal with a single large

terminal spine on the inner lobe and a much smaller spine some

distance away on either side of it; endopodite bent outwards,

with two terminal and two much smaller lateral sete ; exopodite

with four subequal sete.

lst maxilla: structure not properly made out.

2nd maxilla and maxillipede as in P. parvula, of the usual

Oithona structure but rather small.

1st foot (Pl. X XIX. fig. 6) with two-jointed exopodite oncil

endopodite, the division han ween the 2nd and 3rd joints of the

exopodite being only faintly indicated ; exopodite with 1.1.2

outer edge spines and 0.1.4 inner edge sete, the terminal spine

considerably longer than the exopodite; endopodite with 1. 4

inner edge and 0.1 outer edge sete; Ist basal with an inner

edge and 2nd basal with an outer edge seta.

2nd to 4th feet with three-jointed exopodites and two-jointed

endopodites.

2nd foot (Pl. X XIX. fig. 7): exopodite with 1.1.2 outer edge

spines and 0.1.5 inner edge setz, terminal spine ilar than

the exopodite ; endopodite with two ‘inner edge, one terminal and

one outer edge setz on the distal end of the 2nd joint; no sete

seen on the basal joints.

3rd foot (Pl. X XIX. fig. 8): exopodite with 1.0.1 outer edge

spines and 0.1.5 inner edge sete; there is a small tooth on the

middle of the outer edge of the 3rd joint; terminal spine very

slender, longer than the excpodite ; endopodite as in 2nd foot;

2nd basal with an outer edge seta, no seta on Ist, basal. °

4th foot (Pl. X XIX. fig. 9): exopodite with 0.0.1 outer edge

spines and 0.0.5 inner edge set, terminal spine longer than

the exopodite ; endopodite as in 2nd and 3rd feet except for a

stout feathered seta on the inner edge of the Ist joint; 2nd

basal with an outer edge seta, no seta on Ist basal.

COPEPODA FROM CHRISTMAS ISLAND. 191

5th foot very small with a short terminal and a basal seta.

Four specimens were found in one gathering.

This species seems to agree in appearance with Oithona hebes,

but differs in the jointing and spinulation of the swimming-feet.

In structure it is separated from Parotthona parvula, which it

closely resembles, by the reduction in number of the sete on the

endopodites of the swimming-feet. Not much reliance can be

put on the presence or absence of sete on the basal joints of

the swimming-feet, as they may have been overlooked in the

descriptions of both species.

The following is an attempt to draw up a diagnostic table of

the species of Onion and Paroithona of which a recognisable

description exists, which will assist in the identification of the

species described above. The number following each name refers

to the paper in which it was first described.

1. Rostrum present.

A. Rostrum antero-ventrally directed, visible in dorsal

view.

a. Exop. of 1st foot with 1.1.2 outer edge spines.

aa. 2ud basals of swimming- -feet with plumose sete ;

length 10-15 mm. . O. plumifera (1).

bb. 2nd basals with smooth sete ; Tength 1 0-1'16mm. O. atlantica (11).

b. Exop. of 1st foot with 1.1.3 outer edge spines.

aa. Exop. of 4th foot with 0.0.1 outer edge spines ;

Temeriln, NAHAS SWI, cco scecas esaconeacaas000c08000500 O. frigida (15).

bb. Exop. of 4th foot with 1.1.3 outer edge spines ;

length ‘68-74 mm. . O. vivida.

ce. Exop. of 4th foot with 0.0.2 outer edge spines ; ;

length 1°2-1'9 mm. a . O. setigera (8).

. Rostrum ventrally directed! nee Rianied in ies cell view.

a. Exop. of Ist foot with 1.0.2 outer edge spines;

length ‘56-62 mm. ........ . O. decipiens.

b. Exop. of 1st foot with 1.1. 2 onliae dee spines.

aa. Exop. of 2nd foot with 1.0.1 onter edge

spines; length °73="8 mm...........-.--2.-0-0-002-- O. similis (6).

bb. Exop. of 2nd foot with 1.0.2 outer edge

spines; length “88-94 mm. ................000602. O. fallax.

c. Exop. of 1st foot with 1.1.3 outer edge spines.

aa. Exop. of 4th foot with 1.1.2 outer edge spines.

a. Rostrum long, slender, straight; length

TNE Sg ST ae a ae tera ee cr oA O. robusta (18).

8. Rostrum short, curved; length *7 mm. ...... O. brevicornis (18).

bb. Exop. of 4th foot ee 0.0.1 outer edge

spines; length 1:07-1:12 mm. ..................... O. linearis (12).

2. Rostrum absent.

A. Endop. of 2nd-4th feet three-jointed.

a. 5th foot with one terminal seta.

aa. 4th foot with 1.1.3 outer edge spines; length

*388—42 mm. ......... O. simplex.

bb. 4th foot with 1.1. 2 gmiet 650 55 spines.

a. Cephalothorax much longer than abdomen ;

length *55 mm. ..... O. hebes (18).

. Cephalothorax slightly longer than abdomen ;

length ‘50-53 mm. .... O. nana (18).

y. Cephalothorax shorter than abdomen ; 2 ‘length

ie) TUNING Men dod nano bde DoMBoTGCOIDOUG AOE HOB paBBBUGHS TOE MANAHG DOK: RAGS

192 ... -MR.-G. P. FARRAN ON’

_ 6. 5th foot with two terminal sete.

aa. Mandible with large Spine on 2nd baal

length ‘64mm. ... ..... O.minuta (16).

bb. Mandible with small spines on 2nd basal.

a. Terminal spines of exopodites of swimming-

feet very long; length 9 mm. ...... .. O. oculata.

8. Terminal spines of exopodites of swimming-

feet shorter than the 3rd joint; length 15 -

1313) MING soredsoe .. O.rigida (14).

B. Endop. of all swimming- oeneh “in sonic. “(Gea

Paroithona).

a. Exop. of 4th foot with 1. 1.1 outer edge spines ;

length ‘46mm. ...... .. P. parvula (10).

‘6. Exop. of 4th foot with 0.0.1 outer ‘edge spines ;

length -40--43 mm. ........ beeen ee publa.

I have omitted from the nis: 0. apes (3) (4), O. spini-

Srons (2), O. spinirostris (5), O. helgolandica (5), and O. pygmea

(2). The identity and probable synonymy of these species have

been fully discussed by various writers and it is unnecessary to

consider them further.

LITERATURE REFERRED TO.

(1) Barrp, W.—“ Notes on British Entomostraca.” Zoologist,

vol. i. 1843.

(2) Boncx, A.—Vid. Selsk. Forhandl. Christiania, 1864.

(3) Brapy, G. S.—‘ Challenger’ Reports: Copepoda, pt. Xxiil.

1883.

(4) Brapy, G. 8.—Die Marinen Copepoden der Deutschen Siid-

polar-Expedition, 1910.

(5) Cuaus, C.—Die freilebenden Copepoden, 1863.

(6) Craus, C.—“ Die Copepoden Fauna von Nizza.” Schr. Ges.

Naturw. Marburg, 1866.

(7) Curve, P. 'T.—‘ Plankton from the Indian Ocean and Malay

Archipelago.” Konigl. Sv. Vet.-Akad. Handl., Bd. 35,

WOO. LION:

(8) Dana, J. D.—Amer. Journ. Sci. (2) vol. viii. 1849,

(9) Dana, J. D.—U.S. Explor. Exped., vol. xiii. Crustacea, 1852.

(10) Farran, G. P.—Second Report on the Copepoda of the

Irish Atlantic Slope. Fisheries, Ireland, Sci. Invest.,

1906, 11. [1908].

(11) Farran, G. P.—‘ Note on the Copepod Genus CHUB

Ann. Mag. Nat. Hist. (8) 11. 1908.

(12) Gresprecut, W.—Atti Accad. Lincei Roma, vol. vii. 1891.

(13) Ginsprecu1t, W.—Fauna u. Flora des Golfes von Neapel;

vol. xix. Copepoda, 1893.

(14) GresBrecut, W.—“‘ Ueber pelagische Copepoden des Rothen

Meeres.” Zool. Jahrb., Syst. Bd. ix. 1897.

(15) GinsBRECHT, W.— -Resultats du Voyage du S8.Y. Belgica.

Copepoden, 1902.

(16) Scorr, T.—‘ Report on Entomostraca from the Gulf of

Guinea.” Trans. Linn. Soe. London (2) vol. vi. 1893.

(17) Woxrexven, R. N.—Fauna and Geography of the Maldive

and Laccadive Archipelagoes, vol. 11. Suppl. 1, Copepoda,

1905.

COPEPODA FROM CHRISTMAS ISLAND. 193

EXPLANATION OF THE PLATES,

All the figures have been drawn with the aid of a camera lucida.

Pratt XXVII.

Fig.

. Oithona vivida 2, Ist antenna.

mandible palp.

1st maxilla.

dorsal view.

cephalon, lateral view.

1st foot.

2ud foot.

3rd foot.

| Oithona ‘fallas 2, Ist foot.

2nd foot.

8rd foot.

4th foot.

Pratt XXVIII.

Fig. e Oithona fallax °, cephalon, lateral view.

Fig.

be)

mandible palp.

dorsal view.

Oithona decipiens 2, cephalon, lateral view.

FOODS TB ow po

mandible palp.

Ist maxilla.

dorsal view.

Ast foot.

2nd foot.

3rd foot.

4th foot.

PratTE XXIX.

Paroithona pulla §, dorsal view.

Ist-antenna.

abdomen.

mandible palp.

_2nd antenna.

1st foot.

2nd foot.

3rd foot.

4th foot.

10. Oithona simplex 2, cephalon, lateral view.

Ai Ag

TIS senile

aes

dorsal view.

abdomen.

8rd foot.

4th foot.

Prater XXX.

be)

. Oithona simplex 9, 1st antenna.

Ist foot.

. Oithona attenuata 2, cephalon, lateral view.

1st antenna.

dorsal view.

8rd foot.

Ist foot.

| Oithona oculata 2, maxillipede.

5. ”

oO 39

oF)

4th foot.

Prarr XXXI.

Paroithona pulla 2, cephalon, lateral view.

Oithona oculata 2, dorsal view.

GOGI ENED BOE ES WS)

lateral view.

1st antenna.

mandible palp.

Ist maxilla.

1st foot.

2nd foot.

3rd foot.

194 MR. H. B. PRESYON ON AGNATHOUS

15. Diagnoses of New Species and Varieties of Agnathous

Mollusca from Equatorial Africa. By H. B. Presron,

ld Bas

[Received November 8, 1912: Read February 18, 1913.]

(Plates XX XIT.-XXXV.*)

INDEX. Page

Geocraplicall See ened ne ae see ec esanteese aetna eens Oms

Systematic :

Streptaxis: 5 new species .......................... 194-195

Ennea: 53 new species; 11 new varieties ...... 196-216

Ennea syngenes, nom. nov. tor E. consobrina... 216

The species of which descriptions are given in the present

paper were chiefly collected in Hast and East Central Africa by

Messrs. Robin Kemp, A. Blayney Percival, and C. W. Woodhouse

in their recent travels in those parts of the African continent,

and, further, go to show the enormous field for conchological

research awaiting the student of this very fruitful region, in

many parts of which it would appear that, probably owing to

variation in rainfall, altitude, etc., each range of hills has toa

certain extent its own special molluscan fauna, often characterised

by certain local and peculiar phases not only common to the

species but also to the genera occurring in that particular

locality.

SrREPTAXIS PERCIVALI, sp.n. (Pl. XXXII. fig. 4.)

Shell ovate, somewhat inflated, thin, semitransparent, yel-

lowish white; whorls 5, the earlier whorls regularly increasing,

the later proportionately large, the last two distorted, sculptured

with fine, very oblique, and closely set costule which become

obsolete on the base of the shell; suture impressed, narrowly

margined below ; umbilicus moderately narrow, deep ; columella

outwardly expanded, descending in a strong curve, diffused above

into a thin, ill-defined, projecting callus which reaches the upper

margin of the labrum; labrum white, narrowly expanded and

reflexed ; aperture somewhat oblique, roundly ovate.

Alt. 9°25, diam. maj. 7, diam. min. 6 mm.

Aperture: alt, 4, diam. 3°5 mm.

Hab. Northern slopes of Mt. Marsabit, British East Africa

(A. Blayney Perciwal).

Allied to S. kirkii Dohrn T, but differing chiefly in having one

whorl less.

STREPTAXIS WOODHOUSEI, sp. n. (PI. XXXII. fig. 5.)

Shell differing from S. percivali, to which it is nearly allied,

in its more rectangular and less inflated form, more obtuse apex,

* For explanation of the Plates see p. 217.

+ Proc. Zool. Soc. Lond. 1865, p. 232.

TE, Ae SOUS) JEN, SOOMUL.

WS, 1724

AH. Searle del, et Lith, Huth imp,

NEW STREPTAXIS AND ENNEA FROM EQUATORIAL, AFRICA.

P.Z.5.19138.PlLXXXII1.

18. ava, 14a

AH. Searle del et lith,

NEW ENNEA FROM EQUATORIAL AFRICA.

Huth imp.

P.Z.S. 1913. Pl. XXXIV.

A.H.Searle, del et lith.

NEW ENNEA FROM EQUATORIAL AFRICA.

PZ:S. 1013), Pl OOK;

14.

AH.Searle, del et lith.

NEW ENNEA FROM EQUATORIAL AFRICA.

MOLLUSCA FROM EQUATORIAL AFRICA. 195

and rather coarser transverse sculpture; the umbilicus is nar-

rower and the columella descends almost vertically in a very

slight curve and extends above into a thickish, well-defined,

par rietal callus; the aperture is also much more rectangular than

in that species.

Alt. 9°5, diam. maj. 7, diam. min. 5°75 mm.

Aperture: alt. 3°75, diam. 3 mm.

Hab. Mt. Elgon, Uganda (C. W. Woodhouse).

STREPTAXIS MARSABITENSIS, Sp. n. (Pl. XXXIT. figs. 2, 2a.)

Shell very depressedly turbinate, cream-coloured; whorls 44,

regularly increasing, the earlier whorls smooth, polished, shining,

the later whorls sculptured with moderately fine, closely set,

arcuate riblets which become obsolete on the base of the shell;

suture impressed; umbilicus wide, deep; columella descending in

an oblique curve, white, slightly expanded, outwardly extending

above into a somewhat thick, well-defined, polished, parietal

callus which reaches the upper margin of the labrum ; labrum

white, considerably expanded-and reflexed below, slightly so

above, where it is gibbously bent somewhat downwards in front,

thus forming, with the parietal wall, a broad, but very shallow,

sinus ; aperture obliquely and broadly sublunate.

Alt. 7, diam. maj. 13°75, diam. min. 11°25 mm.

Aperture: alt. 5, diam. 4°5 mm.

fab. Northern slopes of Mt. Marsabit, British East Africa, at

an altitude of 4600 feet (A. Blayney Percival).

STREPTAXIS URGUESSENSIS, sp.n. (PI. XXXII. figs. 3, 3 a.)

Shell allied to S. marsabitensis, but differing from that species

in its larger size, broader, but proportionately no higher, form,

and rather coarser transverse sculpture above, though the basal

sculpture, while being much more marked, is considerably

finer; the umbilicus is proportionately shghtly narrower and the

columella more obliquely descending; the labrum has converging

margins and is more sinuous and gibbously bent downwards over

the aperture, which is subrectangular and much broader in shape

than is the case with S. marsabitensis.

Alt. 9°5, diam. maj. 16, diam. min. 13 mm.

Aperture: alt. 7, diam. 5°5 mm.

Hab. Urguess, British Hast Africa (A. Blayney Percival).

STREPTAXIS DESIDERATA, sp.n. (Pl. XXXII. figs. 1, 1a.)

Shell suborbicular, very depressedly turbinate, polished, shining,

cream-coloured ; shore 5}, regularly increasing, the. last sims.

what rapidly br oadening towards the aperture, marked only with

indistinct, transverse growth-ridges; suture impressed, rather

coarsely crenellate; umbilicus open, broad, deep; columella

descending in an oblique curve above, very oblique below, reflexed,

whitish, diffused into a thin, fairly well-defined, and broadly,

outwardly extending, parietal callus which reaches the upper

196 - MR. H. B, PRESTON ON AGNATHOUS.

margin of the labrum; labrum thickened, expanded, reflexed and

considerably receding below, then scarcely reflexed, projecting

and somewhat inwardly bent over the aperture and finally again

receding above; aperture very broadly and compressedly, obliquely

sublunate.

Alt. 11, diam. maj. 20, diam. min. 16 mm.

Aperture: alt. 8°5, diam. 8°25 mm.

Hab. Urguess, British Kast Africa (A. Blayney Percival).

ENNEA BUCCINA, sp. n. (PI. XXXII. fig. 11.)

Shell rather large, edentulate, somewhat ovately trigonal,

white; whorls 63, the first five rather rapidly, but regularly

increasing, the last very large, rapidly ascending in front, slightly

diverging from the axis of the remainder, indistinctly, transversely,

arcuately plicate, the plicee becoming obsolete on the last whorl

which is nearly smooth; suture lightly impressed; umbilicus

appearing as a very narrow, shallow, curved, elongate depression ;

columella descending in a slightly oblique curve, diffused above

into a thin, opaque, rather ill-defined, parietal callus which

terminates just in front of the upper margin of the labrum;

labrum white, porcellanous, outwardly expanded and very nar-

rowly reflexed except near its upper margin where, though

thickened, this is scarcely the case, projecting in front, receding

below and slightly above to form a shallow curved sinus; aperture

very oblique, wleeceing ula, with rounded base.

Alt. 20°25, diam. maj. 15-5, diam. min. 12°75 mm.

Aperture: alt. (nearly) 9°5, * diam. 5°75 mm.

Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude

of 6000 feet (Hobin Kemp).

ENNEA MARGARITA, sp.n. (PI. XXXIV. fig. 8.)

Shell ovate, polished, shining, whitish cream-colour; whorls 6,

rapidly increasing, the penultimate gibbous, the last slightly

oblique, the earlier whorls sculptured with transverse riblets

which become obsolete on the later whorls except at the suture,

the last whorl marked with very fine, almost imperceptible,

scratch-like, spiral striew ; suture impressed, margined below, the

margin rather coarsely sculptured by the terminations of the

transverse riblets ; umbilicus (in the adult) quite closed, the um-

bilical area being represented by a shallow depression ; columella

slightly excavated above, obliquely descending below; labrum

pure white, expanded and narrowly reflexed, the margins joined

by a thin, well-defined, pariet tal callus; aperture subquadrate,

edentulate. ,

Alt. 12°5, diam. maj. 7°5 mm.

Aperture: alt. 5, diam. 3°25 mm.

Hab. Kigezi, extreme 8.W. Uganda, at an altitude of 6000 feet

(Robin Kemp).

Young specimens are somewhat narrowly, but profoundly,

umbilicate. __

MOLLUSCA FROM. EQUATORIAL AFRICA. uegi7e

- EINNEA KIVUENSIS, sp. n. (Pl. XXXIV. fig. 3.)

Shell differing from H#. margarita in its smaller size, rather

more cylindrical form, and generally thinner texture; the trans-

verse sculpture is more marked, the penultimate whorl is pro-

portionately less gibbous, and the last whorl is comparatively

ee oader than is the case in that species.

Alt. 10°5, diam. maj. 6°25 mm.

Aperture : alt. 4°25, diam. 3 mm.

Hab. Near Lake Kivu, Belgian Congo (Lobin Kemp).

ENNEA INNOCENS, sp. n. (Pl. X XXIII. fig. 11.)

Shell scarcely rimate, rather large, cylindrically ovate, cream-

coloured, smooth, shining; whorls 6, rather flat, the first three

regularly, the remainder rapidly, increasing in size, the last very

slightly ascending in front ; suture impressed ; columella vertically

descending, slightly bulging in the middle, “narrowly outwardly

expanded ; labrum white, very narrowly reflexed and dilated

below; aperture irregularly subquadrate, edentulate.

Alt. 13°75, diam. maj. oer

Aperture: alt. 5°25, diam. 3°25 mm.

Hab. Jombene Hills, British East Africa, at an altitude of

6000 feet (Lobin Kemp).

ENNEA ELGONENSIS, sp.n. (PI. XX XIII. fig. 3.)

Shell allied to #. innocens, but much smaller and with more

obtuse apex, though having the same number of whorls; the

umbilical depression is proportionately broader, the columella

descends in a curve, the labrum is not dilated below, and the

aperture is ovate instead of being subquadrate as in #. innocens.

_ Alt..10°75, diam. maj. 5°75 mm.

Aperture: alt. 4, diam. 2°75 mm.

Hab. Mt. Elgon, Uganda (C. W. Woodhouse),

. ENNEA InGLoRIA, sp. n.. (Pl. XX XIII. fig. 10.)

Shell small, shortly cylindrical, thin, whitish; whorls 6,

the first four regularly increasing, the fifth and sixth rather

rapidly increasing in length, but not in breadth, smooth but for

coarse growth-markings, the last two whorls convex; suture

impressed, strongly crenellate below; perforation very narrow;

columella slightly oblique and outwardly expanded ; labrum nar-

rowly dilated below and reflexed ; aperture edentulate, subovate.

Alt. 4:5, diam. maj. 2°25 mm.

Aperture: alt. 1, diam. °5 mm.

Hab. Mt. Nyiro, to the south of Lake Rudolph, at an altitude

a 8300 feet (A. Blayney Percival).

ENNEA RECTANGULARIS, sp. n. (Pl. XXXV. fig. 10.)

Shell rectangularly cylindrical, with planulate apex, thin, trans-

parent, polished, shining, vitreous, pale greenish ; whorls 6, the

first two flattened above, the third increasing in length and

198 MR. H. B. PRESTON ON AGNATHOUS

breadth, the fourth also increasing in length, but slightly dimin-

ishing in breadth, the fifth increasing in length and again in

breadth to equal that of the third, the sixth increasing in length

but not in breadth ; suture impressed, narrowly margined below ; '

columella rather obliquely descending, white, porcellanous ;

labrum white, narrowly, outwardly expanded and reflexed except

near the upper margin, where it becomes almost acute; aperture

subquadrate, edentulate.

Alt. 6°75, diam. maj. 3 mm.

Aperture: alt. 1:5, diam. 1 mm.

Hab. Urguess, British East Africa (A. Blayney Percival).

ENNEA BACCATA, Sp.n. (Pl. XXXII. figs. 10, 10.)

Shell cylindrically ovate, cream-coloured; whorls 8, the first

very small, the second large in proportion, the third, fourth, fifth,

and sixth regularly increasing in length and breadth, the seventh

increasing in length, but not in breadth, the eighth increasing in

length, but diminishing in breadth, sculptured, especially on the

upper whorls, with fine, slightly oblique, closely set, transverse

strie; suture lightly impressed; umbilical area presenting a

narrow depression; columella not very obliquely descending,

spreading above into a very thin, almost imperceptible, but well-

defined, parietal callus which reaches to the upper margin of the

labrum; labrum sinuous, outwardly expanded, especially at the

base, reflexed ; aperture armed with an erect, oblique, parietal

lamella situated very near the upper margin of the labrum, a

rather coarse lamella-like projection on the outer lip, a small

nodulous basal denticle at the foot of the columella, above which,

and rather more exteriorly situate, occurs another and stronger

nodulous denticle, and above this again a rather broad, very

interiorly situate, columellar projection.

Ait. 6, diam. maj. 3 mm.

Aperture: alt. 1:25, diam. °75 mm.

Hab. Urguess, British Kast Africa, at an altitude of 6000 feet

(A. Blayney Percival).

ENNEA ARTHURI, sp.n. (Pl. XXXII. figs. 9, 9a.)

Allied to #. baccata, but much smaller, with more obtuse apex;

it has one whorl less, the first four increasing regularly, the fifth

proportionately in length, but not in breadth, while the sixth and

seventh increase also in length, but diminish in breadth; the

surface is quite smooth, polished and shining, without any trace

of the transverse stri# which are characteristic of H. baccata;

the suture is narrowly margined above, the labrum is not sinuous,

the columella is much more oblique and lacks the internal pro-

jection on the upper part which is so well developed in that

species.

Alt. 7:25, diam. maj. 3°75 mm.

Aperture: alt. 3, diam. 1°5 mm.

Hab. Jombene Hills, British Fast Africa, at an altitude of

4000 feet (A. Blayney Percival).

MOLLUSCA FROM EQUATORIAL AFRICA. 199

ENNEA ADELPHA, sp.n. (PI. XXXII. figs. 7,7a; Pl. XXXIV.

figs. 1, 1 a.)

Shell allied to H. vicina Smith* from Chiradzulu, British

Central Africa, but differing from that species chiefly in its

coarsely crenellate suture and coarse, transverse, plicate sculpture ;

the small parietal denticle of H. vicina is also much more developed

and the broad, tridentate projection on the inner margin of the

outer lip less so.

Alt. 6°5, diam. maj. 3°25 mm.

Aperture: alt. 2°25, diam. 1-75 mm.

Hab. Between Masaka and Entebbe, extreme 8.W. Uganda

(Robin Kemp).

The very distant locality of #. vicina, which is situated to the

south of Lake Nyassa, has to a certain extent influenced me in

separating the present species.

EINNEA PERTURBATA, Sp. n. (Pl. XXXYV. figs. 7, 7 a.)

Shell allied to #. adelpha and ina measure to /. vicina Smith,

but shorter and with less convex whorls than either, besides

lacking the two internal basal denticies which seem to be a

character common to both those species.

Alt. 6, diam. maj. 3 mm.

Aperture: alt. 1°75, diam. 1 mm.

Hab. Mt. Elgon, Uganda (C. W. Woodhouse).

ENNEA COENI, sp.n. (Pl. XXXII. figs. 15, 15 a.)

Shell differing from 1. perturbata and its allies H. vicina Smith

and #. adelpha in lacking the group of three closely situated and

projecting lamelle on the upper part of the columella, which, in

the present species, are replaced by a single, small, and slightly

curved denticie.

Alt. 5°75, diam. maj. 2°75 mm.

Aperture: alt. 1°5, diam. °75 mm.

Hab. Mt. Elgon, Uganda (C. W. Woodhouse).

ENNEA WOODHOUSEI, sp. n. (PI. XXXV. figs. 15, 15a.)

Shell allied to #. perturbata, but considerably smaller, with

rather coarser and more distant transverse plicee and more basally

constricted aperture ; the tridentate projection on the inner side

of the outer lip, which is common, not only to the above species,

but also to #. vicina Smith, and to #. adelpha, in the present

case appears as a very broad, bidentate, shelf-like protuberance,

almost choking the aperture.

Alt. 4°75, diam. 2°5 mm.

Aperture: alt. 1, diam. °75 mm.

Hab. Mt. Elgon, Uganda (C. W. Woodhouse).

ENNEA NYIKAENSIS, sp. n.

Shell differing from /. fortidentata Smith? in its larger and

* Proc. Zool. Soc. Lond. 1899, pp. 580-581, pl. xxxiii. figs. 1, 2.

7 Ann. & Mag. Nat. Hist. 1890, vi. p. 162, pl. vi. fig. 6.

200 _oMR. H. B. PRESTON ON AGNATHOUS -

much broader. form, the. crenulation atthe suture is also con-

siderably finer, and it also lacks the small parietal denticle of that

species.

Alt. 9:25, diam. maj. 6°25 mm.

Aperture: alt. 4, diam. 3°25 mm.

Hab. Nyika Range, Nyassaland.

Ennea fortidentata was described by Mr. E. A. Smith, in the

year 1890, from specimens collected by Mr. Last at Mamboia,

German Hast Africa, and in 1897 a number of specimens of an

Ennea were received at the British Museum from Sir H. H.

Johnston which had been collected far to the south on the Nyika

Range in Nyassaland*. As the dentition appeared to be nearly

identical these specimens were considered to be a variety of

E. fortidentata and have been regarded as such until the present

time. Through the courtesy of Mr. Smith I have now been able

to examine both the type specimens of /. fortidentata and also

the series of shells from Nyassaland, besides, through the kindness

of Mr. A. Blayney Percival, a vast number of fairly typical

specimens of that species from Mt. Elgon in Uganda, with the

result that Iam forced to the conclusion that the Nyassaland

shells are quite distinct from those collected at Mamboia and more

recently on Mt. Elgon.

The type of H. nyikaensis is in the British Museum.

ENNEA CAREA, sp.n. (Pl. XXXII. figs. 13, 13 a.)

Shell small, cylindrical, white, smooth, polished, shining ;

whorls 6, very flat, the last broad; sutures incised, narrowly

margined above; columella arched above; aperture irregularly

subquadrate, armed with a nodulous, parietal lamella, two diver-

gent lamelle on the columella and a bifurcate lamella on the

upper margin of the outer lip; labrum whitish, thickened,

scarcely reflexed.

Alt. 3:25, diam. maj. °75 mm.

Hab. Gazi, British East Africa (Robin Kemp).

ENNEA CopPiosa, sp.n. (Pl. XXXIT. figs. 16, 16a.)

Shell cylindrically ovate, with very obtuse apex, thin, trans-

parent, polished, shining, vitreous, whitish ; whorls 63, the first

four and a half regularly increasing in length and breadth, the

penultimate increasing proportionately in length but not much in

breadth, the last whorl increasing in length but considerably

diminishing in breadth, marked only on the upper whorls with

oblique, transverse striz, the latter half of the last whorl and the

base of the shell being finely costulate, especially round the

umbilical area; suture impressed, narrowly margined below and

showing occasional traces of nodulous crenellation ; umbilical

depression ovate, moderately open and deep; columella very

* Proc. Zool. Soc. London, 1899, p. 581.

MOLLUSCA FROM EQUATORIAL AFRICA. 201

slightly, obliquely descending, broadly, outwardly expanded,

vitreous, both this and the labrum, which is also rather broadly,

outwardly expanded and narrowly reflexed at the edge, showing

the white laminiferous, somewhat irregular growth-periods ;

aperture irregularly subquadrate, armed with an erect sickle-

shaped parietal lamella, which starts from the upper margin of

the labrum and proceeds downwards in an oblique direction for

some distance, when it terminates in a broad, shallow, cup-like

projection ; two glassy, lamella-like teeth on the inner margin of

the labrum, of which the lower is the larger and is much more

quadrate than the upper; a small, interiorly situate, basal denticle ;

a minute, sharp, thorn-like denticle at the base of the columella,

and a triangular, trifid, projecting denticle or lamella on the

upper portion of the columella.

Alt. 6, diam. maj. 3°25 mm,

Aperture: alt, 1:5, diam. 1 mm.

Hab. Urguess, British Hast Africa (A, Blayney Percival).

Var. ROBUSTA, Var. n,

Shell differing from the type in its larger and more eylindrical

form and in the stronger and more fully developed apertural

armature, though the system remains the same,

Alt. 7°75, diam, maj. 3°75 mm.

Aperture: alt. 1:75, diam, 1°25 mm.

Hab, Laregi Hills, British East Africa, at an altitude of 6000

to 7000 feet (A. Blayney Percival),

ENNEA CURYVICOLUMELLA, sp.n, (PI. XXXII. figs, 17,174.) .

Shell small, perforate, shortly cylindrical, smooth, shining,

vitreous, creamy white ; whorls 6, flattish, the first slime rather

rapidly increasing, the fourth and fifth gradually so, the sixth

rather broad in proportion to the remainder; suture impressed,

margined below ; umbilicus moderately wide and deep; colu-

mella descending in an angular curve and excavated at the

angle, where, on the inner margin, is developed a coarse, lamelli-

form protuberance projecting into the interior of the shell;

labrum continuous, somewhat expanded and reflexed, bearing two

lamelliform plaits (of which the lower is the larger) on the inner

margin of the upper portion, and a coarse, slightly curved, erect,

parietal lamella above; aperture rather elongately subquadrate.

Alt, 2°75, diam. maj. 1°25 min,

Hab. Between the Jombene Hills and Nyeri, British East

Africa (Robin Kemp).

ENNEA DECUSSATULA, sp.n, (PI. XXXJIT. figs. 1, 1 a.)

Shell cylindrically ovate, thin, cream-coloured ; whorls 7, the

first five regularly increasing, the sixth long in proportion to

those above, the seventh also long, but diminishing in breadth

and bistrangulate just behind the labrum, sculptured with rather

Proc. Zoou, Soc.—1913, No. XIV. 14

202 MR. H. B. PRESTON ON AGNATHOUS

coarse, oblique, transverse costule, and minutely, spirally malleated

so as to present a slightly decussate appearance ; suture incised,

strongly crenellated by the terminations of the transverse

costule; umbilical area appearing as a narrow and shallow

depression ; columella slightly oblique; labrum white, narrowly

expanded and reflexed, the margins distant; aperture sub-

rhomboidal, armed with a single, erect, nearly vertically de-

scending, parietal lamella, a small denticle and a somewhat weak

lamella on the inner side of the outer lip, a well-developed basal

denticle at the base, and a broad squarish shelf-like projection on

the columella bearing a central depression.

Alt. 11°75, diam. maj. 5°5 mm.

Aperture: alt. 4, diam. 2°5 mm.

Hab. Kigezi, extreme S.W. Uganda, at an altitude of 7000 feet

(Robin Kemp).

ENNEA DISSEMINATA, sp.n. (Pl. X XXIII. figs. 2, 2 a.)

Shell small, rather shortly cylindrical, whitish, slightly polished ;

whorls 7, the third, fourth, and fifth somewhat convex, the

sixth and seventh flattish. sculptured, especially on the later

whorls, with indistinct, oblique, transverse costule; suture im-

pressed, rather distantly crenellated by the terminations of the

transverse costule ; umbilicus reduced toa mere fissure; columella

curved, extending above into a narrow, but thickish, parietal

eallus ; labrum white, narrowly reflexed ; aperture subquadrate,

armed with a short, erect, parietal lamella, a broad, subbifid,

tooth-like projection on the inner margin of the outer lip, a small

basal denticle at the foot of the columella, and above this a broad,

rounded projection, very interiorly situate.

Alt. 4°25, diam. maj. 2 mm.

Aperture: alt. °75, diam. °5 mm.

Hab. Near Lake Kivu (type specimen); also between Lake

Mutanda and the region between that lake and Lake Kivu

(Robin Kemp).

ENNEA EUSSOENSIS, sp.n. (Pl. XX XIII. figs. 4, 4 a.)

Shell ovately cylindrical with obtuse apex, scarcely rimate,

moderately solid, shining, dark cream-colour ; whorls 6, the first

three rapidly, the remainder regularly, increasing, slightly con-

vex, smooth but for lines of growth ; suture impressed ; columella

white, rather broadly, outwardly expanded; labrum white,

expanded, slightly reflexed; aperture rather broad and short,

armed with a coarse, oblique, parietal lamella situated on the

right-hand side, a short, tooth-like lamella on the inner margin

of the outer lip and a small lamella situated on the median region

of the columella.

Alt. 5:5, diam. maj. 2°5 mm.

Aperture : alt. 1-5, diam. 1:25 mm.

Hab. Eusso Nyiro, British East Africa (Robin Kemp).

MOLLUSCA FROM EQUATORIAL AFRICA. 203

ENNEA FOVEOLATA, sp.n. (PI. XX XIII. figs. 5, 5a.)

Shell small, cylindrical, cream-coloured, polished; whorls 6,

rather flat, plicate at the suture, otherwise smooth with the

exception of the latter portion of the last whorl which is plicate

throughout, last whorl bearing three hollows or dents just behind

the peristome; suture impressed; columella produced internally

into a very broad, squarish, shelf-like projection ; aperture some-

what square, armed with a small, rounded denticle immediately

below the shelf-like columelJa, a broad projection inside the outer

lip, a small denticle above this and a curved, erect, parietal

lamella ; labrum white, expanded, reflexed.

Alt. 4°25, diam. maj. 1°5 mm.

Aperture: alt. ‘75, diam. -75 (nearly) mm.

Hab. Gazi, British Kast Africa (Robin Kemp).

ENNEA FUNEREA, sp.n. (Pl. XX XIII. figs. 6, 6a.)

Shell cylindrical, rather solid, somewhat shining, covered with

a dark blackish-brown periostracum ; whorls 7, the first rapidly,

the last three moderately increasing, flattish, sculptured with

regular, very oblique, transverse costule ; suture impressed ;

columella vertically descending, outwardly broadened and ex-

panded, greyish white; labrum slightly granular, also greyish

white and outwardly expanded with reflexed margin; aperture

subquadrate, bearing a coarse, erect, parietal lamella, two slightly

distant lamellae on the inner margin of the outer lip, of which

the lower is the larger, a moderately sized, basal, tooth-like

lamella, and a very coarse and blunt, almost bifurcate lamella on

the columella.

Alt. 6, diam. maj. 2°75 (nearly) mm.

Aperture: alt. 1°75, diam. °75 mm.

Hab. Chanler Falls, Eusso Nyiro River, British Kast Africa

(Robin Kemp).

Var. LAEVIS, var. n.

Shell differing from the typical form in being almost destitute

of the transverse sculpture.

Alt. (of type specimen) 7, diam. maj. 2°75 mm.

Aperture: alt. 2, diam. ‘75 mm.

Hab. Between the Eusso Nyiro River and Mount Marsabit,

British Hast Africa (A. Blayney Percival).

Though the series of the typical form collected by Mr. Kemp

at the Chanler Falls scarcely varies in size, the present variety

shows enormous differences, as below :—

Largest specimen : alt. 8°5, diam. maj. 3°25; aperture: alt. 2,

diam. °75 mm.

Smallest specimen: alt. 4°75, diam. maj. 25; aperture:

alt. 1:5, diam. -75 mm.

ENNEA HECTOR, sp.n. (PI. XXXITI. fig. 7.)

Shell large, ovately cylindrical, rather thin, smooth but for

14*

204 MR. H. B. PRESTON ON AGNATHOUS

lines of growth, somewhat shining, creamy white; whorls 73, the

first five regularly, the remainder rapidly, increasing, the last

contracted below; suture impressed; umbilicus moderately

narrow; columella obliquely descending, white, reflexed, diffused

above into a lightish callus which joins the upper margin of the

labrum; labrum white, sinuous, expanded and reflexed, especially

in the basal region; aperture irregularly subquadrate, armed

with an erect, somewhat bent, oblique, parietal lamella slightly

broadening below, a group of three denticles on the inner margin

of the labrum, of which the middle one is the largest and the

lowest the smallest, a small, triangular, basal denticle and a strong

curved lamelliform projection about halfway up the columella.

Alt. 18°25, diam. maj. 9 mm.

Aperture: alt. 6, diam. 4°25 mm.

Hab. Between the Jombene Hills and Nyeri, British Hast

Africa (type) (Robin Kemp).

A very handsome species which it would be difficult to con-

found with any other as yet recorded from the region in which it

oceurs; specimens were also sparingly found by Mr. Kemp on

the Jombene Hills at an altitude of 6000 feet and at Mweru.

‘ ENNEA INGEZIENSIS, sp.n. (Pl. XXXIII. figs. 9, 9a.)

Shell small, ovately cylindrical, cream-coloured, somewhat

shining; whorls 7, the first five regularly increasing, the last

two longer but narrower than the fifth, the last deeply strangu-

late just behind the labrum, marked with obsolete, oblique,

transverse riblets; suture well impressed, strongly crenellated by

the terminations of the obsolete, transverse riblets ; umbilical

area presenting a rather deep depression ; columella descending

in a somewhat angular curve and developed into a broad, shelf-

like projection well within; labrum white, expanded and

narrowly reflexed, the margins joined by a well-defined, parietal

callus; aperture subrectangular, armed with a coarse, erect,

somewhat curved, parietal lamella, a large, squarish, tooth-like

projection on the labrum corresponding to, and formed by, the

deep strangulation on the last whorl, a small, triangular denticle

at the base of the columella, above which and very interiorly

situate is a broad, shelf-like projection.

Alt. 4:5, diam. maj. 2 mm.

Aperture: alt. 1:25, diam. 1 mm.

Hab. Ingezi, Mfumburu Mountains, extreme 8.W. Uganda ;

also from Kigezi in the same region at an altitude of 6000 feet

(Robin Kemp).

Var. PUSILLA, var. n.

Shell much smaller than the typical form, with one whor] less,

which gives to it a more cylindrical appearance; the columella

also descends more vertically than is the case with the type.

Alt. 3:25 (nearly), diam. maj. 1:5 (nearly) mm.

Hab. Between Mbarara and Kigezi, extreme 8.W. Uganda

(Robin Kemp).

MOLLUSCA FROM EQUATORIAL AFRICA. 205

Var. MBARARAENSIS, var. n.

Shell smaller than the typical form and proportionately shorter,

with one whorl less, the denticle at the base of the columella is

also removed to a position considerably higher up and almost

overhanging the internal, shelf-like projection.

Alt. 3°5 (nearly), diam. maj. 1°75 mm.

Aperture: alt. 1 (nearly), diam. ‘75 mm.

Hab. Between Mbarara and Masaka, extreme 8.W. Uganda

(Robin Kenp).

ENNEA INSULSA, sp. n. (Pl. XX XIII. figs. 13, 13 a.)

Shell elongately ovate, very minute, cream-coloured ; whorls 43,

very convex, sculptured with oblique, transverse riblets, the last

ascending in front; suture deeply impressed; aperture sub-

quadrate, armed with a parietal lamella bearing a small, internal,

erect denticle towards the base of the outer lip and an internal

lamella on the columella; labrum white, outwardly expanded,

continuous.

Alt. 1°75, diam. maj. °75 mm.

Hab. Gazi, British East Africa (Robin Kemp).

EINNEA INTRADENTATA, sp.n. (Pl. XXXITI. figs. 14, 14 a.)

Shell small, cylindrical, yellowish white; whorls 64, the first

two and a half rapidly, the remainder scarcely, increasing, rather

flat, smooth ; suture impressed ; umbilical area forming a shallow

depression ; columella descending obliquely, notched above,

bulging inwardly below; labrum white, sinuous, narrowly re=

flexed; aperture irregularly, somewhat elongately subquadrate,

armed with an almost vertical, parietal lamella, an upwardly

directed, tooth-like lamella on the outer lip, below which and very

interiorly situate occurs a nodulous lamella, two converging,

deeply situate, broad, basal denticles, and two broad lamellee also

very interiorly situate on the columella, of which the uppermost

is the stronger.

Alt. 3°5, diam. maj. 1°25 mm.

Hab. Jombene Hills, British East Africa, at an altitude of

6000 feet; a single specimen was also taken on the route between

the Jombene Hills and Nyeri (Lobin Kemp).

ENNEA IRIDESCENS, sp.n. (Pl. XXXIII. figs. 12, 12 a.)

Shell rather elongately cylindrical, thin, semitransparent,

whitish, polished, shining, slightly iridescent; whorls 7, the first

four regularly increasing, the remainder increasing in length but

shightly diminishing in breadth, marked with coarse, oblique

growth-lines; suture impressed; umbilical area presenting the

appearance of a rather broad and shallow depression ; columella

almost vertically descending; labrum rather broadly expanded

and reflexed; aperture subquadrilateral, armed with a small,

rather interiorly situate denticle and a coarse, erect lamella on

the parietal wall, a projection on the inner margin of the outer

lip, and immediately below this and in the interior of the shell

206 MR. H. B. PRESTON ON AGNATHOUS

a stout, squarish denticle, a broad, well-developed, basal denticle

well within the aperture, and a bifid lamella on the columella, the

upper arm of which is much the coarser.

Alt. 4, diam. maj. 1°5 mm.

Aperture: alt. °75, diam. -5 mm.

Hab. Near Kigezi, extreme 8.W. Uganda (Robin Kemp).

Var. CURTA, var. n.-

Shell shorter and proportionately a little broader, with one

whorl less; the large, parietal lamella is less strongly developed

and the basal denticle more strongly so than in the typical

form.

Alt. 3°5 (nearly), diam. maj. 1:5 (nearly) mm.

Aperture: alt. °75, diam.-5 mm.

Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude

of 6000 feet (obin Kemp).

ENNEA BURUNGAENSIS, sp. n. (PI. XXXII. figs. 12, 12a.)

Shell differing from #. iridescens in its shorter, broader, and

rather less cylindrical form, in being obsoletely plicate on the

upper whorls, and in having the suture strongly crenellate; the

aperture is ovate and lacks the small, interiorly situate, parietal

denticle of that species, the projection on the inner margin of the

outer lip is much mere square in shape, stronger and more

marked, and the denticle below is, in the present form, developed

into a long, curved, erect lamella; moreover there are two large

basal denticles, the lower being interiorly situate, instead of the

single basal denticle of H. iridescens, and the bifid lamella on

the columella of that species is replaced by a broad, shelf-like

projection.

Alt. 4°25, diam. maj. 1°75 mm.

Aperture: alt. °75, diam. °5 mm.

Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude

of 6000 feet (Robin Kemp).

ENNEA KENIANA Preston, var. PARVULA, var. n.

Smaller than the typical form * from Mount Kenia.

Alt. 2°5, diam. maj. 1:5 mm.

Hab. Between the Jombene Hills and Nyeri, British East

Africa (Robin Kemp).

Var. INTERMEDIA, var. n.

Intermediate between the typical form and var. parvula.

Alt. 3 (about), diam. maj. 1:75 mm.

Hab. Jombene Hills, British East Africa, at an altitude of

6000 feet (Zeobin Kemp).

Both the above varieties appear to be well-marked local

races, the species seeming to become more stunted in the lower

country.

* Ann. Mag. Nat. Hist. vol. vit. 1911, p. 464, pl. xi. fig. 3.

MOLLUSCA FROM EQUATORIAL AFRICA, 207

ENNEA ABERDARENSIS, sp.n. (PI. XXXII. figs. 6, 6 a.)

Shell differing from H. keniana Preston* in its more ovate

form and much smaller size, in having but 6 whorls, a more

curved columella, more regularly rectangular aperture, and pro-

portionately stronger armature, though the arrangement of the

apertural Jamellz is quite similar.

Alt. 2°75, diam. maj. 2°75 (nearly) mm.

Hab. Mt. Kenangop, Aberdare Range, British East Africa

(Robin Kemp).

ENNEA KIGEZIENSIS, sp.n. (Pl. XXXIV. figs. 2, 2a.)

Shell small, shortly cylindrical, whitish, somewhat shining ;

whorls 63, the first four and a half regularly increasing, the last

two regularly increasing in length, but very slightly diminishing

in breadth, marked with growth-lines, otherwise smooth, except

in the subsutural region, where the shell is coarsely plicate ;

suture well impressed; umbilicus laterally compressed into a

fissure ; columella somewhat curved, outwardly expanded ; labrum

white, expanded outwards and neliexed, the margins converging ;

aperture subquadrilateral, armed with a single, rather coarse,

erect and vertical, parietal lamella, a coarse, nodulous denticle on

the inner margin of the outer lip, below which and very interiorly

situate occurs a broadly triangular lamella, and a broad, 1

teriorly situate projection on the columella ending in three

distant, pointed denticles.

Alt. 3 (about), diam. maj. 1-5 mm.

Hab, Kigezi, extreme 8.W. Uganda (Robin Kemp).

ENNEA LAQUEUS, sp. n. (Pi. XXXIV. figs. 5, 5a.)

Shell rather small, ovately cylindrical, solid, whitish ; whorls

63, the first two and a half regularly increasing, the third much

larger, the antepenultimate proportionately so, the penultimate

increasing in length but scarcely in breadth, and the last also

increasing in length but diminishing in breadth, the apical

whorls faintly and finely, decussately sculptured with transverse

and spiral striz, the remaining whorls sculptured with moderately

fine, closely-set, oblique costule, which become finer and still

more closely set in the strangulated area Just behind the labrum,

and where also traces of microscopic, spiral striz are visible ;

suture well impressed ; umbilical area represented by an obliquely

elongate depression ; columella descending in a curve and bearing;

very internally, a broad, sloping, shelf- like projection ; labrum

white, porcellanous, continuous s, outwardly expanded and reflexed,

callously flattened against the parietal wall; aperture irregularly

ovate, bearing an erect, parietal lamella, and a projection on the

upper portion of the outer lip which bears two small, fine lamelle

above and a coarser one below, the parietal lamellee fitting

between the two former; at the base of the aperture, somewhat

* Ann. Mag. Nat. Hist. vol. vii. 1911, p. 464, pl. xi. fig. 3.

208 MR. H. B. PRESTON ON AGNATHOUS

interiorly situate and a little to one side, appears a moderately

small denticle.

Alt. 4 (nearly), diam. maj. 2 mm.

Aperture: alt. °75, diam. °5 mm.

Hab. Jombene Hills, British East Africa, at an altitude of

4000 feet (A. Blayney Percival).

A very remarkable species, which, apart from the decussate

sculpture of the apical whorls, visible only by the aid of a micro-

scope, 1s easily separable from hitherto-described species of the

group by the interlocking of the parietal lamella with those on

the projection situated just within the aperture on the outer lip.

Var. HERCULES, var. n.

Differing from the typical form in its much larger size, coarser

sculpture, and rather more obtuse apex.

Alt. 6°25, diam. maj. 3 mm.

Aperture: alt. 1°75, diam. 1:25 mm.

Hab. Urguess, British East Africa, at an altitude of 6000 feet

(A. Blayney Percival).

ENNEA CLAUSTRUM, sp.n. (Pl. XXXII. figs. 14, 144.)

Shell allied to H. laqueus, but larger, more cylindrical and less

ovate, it has half a whorl more and is more strongly, transversely

costulate on all but the apical whorls; the columella descends in

a much stronger curve, and instead of the broad, internal, shelf-

like projection, which in the present species is reduced to a very

slight protuberance, there is, very internally situated, a strong,

curved lamella on the outer wall.

Alt. 4°75, diam. maj. 2 mm.

Aperture: alt. ‘75, diam. -75 mm.

Hab. Between the Laikipia Plateau and the Eusso Nyiro River,

British East Africa (A. Blayney Percival).

ENNEA LIMA, sp.n. (Pl. XXXIV. figs. 6-6 0.)

Shell ovately cylindrical, white, somewhat shining; whorls 7,

the first very small, somewhat flattened, the second rather large

in proportion, the third, fourth, and fifth regularly inereasing

in length and breadth, the sixth and seventh increasing in

length, but diminishing slightly in breadth, the apical whorls

smooth, polished, the remainder rather curiously, coarsely,

eancellately sculptured with spiral sulci, the interstices being

crossed by short, closely-set ridges, which develop on the latter

half of the last whorl and on the base of the shell into fine

transverse costule ; suture impressed, narrowly margined below

with white; umbilical depression wide and moderately deep, finely,

transversely costulate; columella white, porcellanous, broadly

and somewhat erectly, outwardly expanded, slightly obliquely

descending, diffused into a white, rather coarsely costulate, well-

defined, parietal callus which reaches to the upper margin of the

MOLLUSCA FROM EQUATORIAL AFRICA. 209

labrum ; labrum white, outwardly expanded and reflexed below ;

aperture subquadrate, armed with a coarse, oblique, parietal

lamella, two coarse, broad, projecting denticles on the inner

margin of the labrum, of which the lower is the larger, a small,

slightly interiorly situate, basal denticle, a minute denticle at the

base of the columella, and a broad, bifurcate, projecting lamella

on its median part.

Alt. 6°25, diam. maj. 3 mm.

Aperture: alt. 2, diam. 1°25 mm.

Hab. Urguess, British Hast Africa (A. Blayney Percival).

ENNEA MALASANGIENSIS, sp.n. (Pl. XXXIV. figs. 7, 7 a.)

Shell cylindrical, yellowish white, somewhat polished ; whorls

6, the first very small, the second and third proportionately large,

regularly increasing, the remainder increasing in length but not

in breadth, moderately convex, having a somewhat weathered

appearance ; suture deeply impressed; umbilicus rather wide,

shallow; columella vertically descending, broadly expanded ;

labrum broadly expanded and reflexed below, continued along

the parietal wall; aperture subrectangular, armed with a small,

interiorly situate denticle, and a strong, curved, parietal lamella,

a broad, squarish projection on the inner margin of the outer

lip, an interiorly situate, basal denticle, and a broad, bifurcate

projection on the columella, of which the upper bifureation is the

stronger.

Alt. 3°25 (nearly), diam, maj. 1:25 mm.

Aperture: alt. °75, diam. *5 mm.

Hab. Malasangi, S.W. Uganda (Robin Kemp).

ENNEA MASAKAENSIS, 8p. n. (Pl. XXXIV. figs. 9, 9a.)

Shell small, acuminately ovate, in dead condition yellowish ;

whorls 6, the first five regularly increasing, the sixth increasing

in length but slightly decreasing in breadth, sculptured with

somewhat distant, vertical, transverse costule; suture im-

pressed ; umbilicus reduced to a narrow and shallow depression ;

columella obliquely descending; labrum continuous, rather broadly

expanded, reflexed ; aperture subquadrate, armed with a parietal

lamella and a broad denticle on the inner margin of the outer

lip.

ae 4, diam. maj. 1°75 mm.

Aperture: alt. °75, diam. ‘75 mm.

Hab. Between Masaka and Entebbe, extreme 8.W. Uganda

(Robin Kemp).

ENNEA MIKENOENSIS, sp.n. (Pl. XXXIV. figs. 10, 10a.)

Shell cylindrically ovate, dark yellowish white; whorls 8, the

first six regularly increasing, the last two increasing in length

but very slightly decreasing in breadth, sculptured with rather

coarse, oblique costule; suture impressed; umbilicus narrow,

deep, rapidly widening below; columella somewhat vertically

210 MR. H. B. PRESTON ON AGNATHOUS

descending; labrum continuous, expanded and slightly reflexed,

notched on its outer margin; aperture subrectangular, armed

with one very small, somewhat interiorly situate, parietal denticle,

and one coarse, curved above and straightly descending, parietal

lamella, a coarse, bifurecate denticle on the inner margin of the

labrum, the upper bifurcation being much smaller than the

lower, a very interiorly situate basal tooth and a coarse, squarish

lamella-like tooth on the columella.

Alt. 4, diam. maj. 2°25 mm.

Aperture: alt. 1, diam. 75 mm.

Hab. Burunga, Mount Mikeno, Belgian Congo, at an altitude

of 6000 feet (Robin Kemp).

ENNEA MONTICOLA, sp.n. (Pl. XXXIV. figs. 12, 124.)

Shell allied to H. commoda Smith * from the Mau Escarpment,

E. Uganda, but smaller, of a more cylindrical shape, with finer

transverse sculpture and having one whorl more; the lamella

on the outer lip is also much more strongly bifurcate than in

E. commoda, and there is, in the present species, no internal basal

denticle immediately below it.

Alt, 3°75, diam. maj. 2 mm.

Aperture: alt. °75, diam. °5 mm.

Hab. Lavogi Hills, British East Africa, at an altitude of 6000-

7000 feet (A. Blayney Percival).

ENNEA MWERUENSIS, sp.n. (PI. XXXIV. figs. 14, 14 a.)

Shell rimate, ovately cylindrical, shining, whitish ; whorls 7

the first four regularly, the remainder rather rapidly increasing,

flattish, the last ascending in front, sculptured with indistinct,

oblique, transverse striae, which are more apparent in the sub-

sutural region and become very marked on the base of the shell ;

suture impressed, scarcely crenellate ; umbilicus reduced to a

narrow fissure; columella obliquely descending, bulging inwardly

and slightly, erectly, outwardly reflexed; labrum reflexed,

expanded, especially on the side opposite to the columella ;

aperture almost irregularly triangular with rounded base, armed

with an erect, oblique, parietal lamella, a projecting, lamelliform

denticle on the inner margin of the outer lip and two denticles,

one situate at the base of, and the other about halfway up, the

columella.

Alt. 7, diam. maj. 3°5 mm.

Aperture: alt. 2°5, diam. 2 mm.

Hab. Mweru, British East Africa (type); also found on the

Jombene Hills at an altitude of 6000 feet (Jtobin Kemp).

It may be well here to point out that the village of Mweru

above mentioned should not be confounded with the well-known

Lake of the same name situated far to the south on the borders

of the Congo State and Northern Rhodesia.

* J. Conch., Leeds, x. p. 315, pl. iv. fig. 9.

MOLLUSCA FROM EQUATORIAL AFRICA, 211

ENNEA NAIVASHAENSIS Preston*, var. ELGONENSIS, var. n.

Shell even more cylindrical than the typical form, with more

closely set and more pronounced transverse riblets, the aperture

also, through not being contracted at the base, has a less triangular

appearance.

Alt. 1°75, diam. maj. 1 mm.

Hab. Mt. Elgon, Uganda (C. W. Woodhouse).

ENNEA MUTANDAENSIS, sp. n. (Pl. XXXIV. figs. 13, 13a.)

Shell differing from H. naivashaensis Preston, from Naivasha,

British East Africa, in its darker colour, more cylindrical form,

less convex and rather longer whorls, and in the broader and

more basally rounded aperture.

Alt. 2, diam. maj. 1 mm.

Hab. Between Lake Mutanda and Lake Kivu (Robin Kemp).

ENNEA NYIROENSIS, sp.n. (Pl. XX XV. figs. 1, 1 a.)

Shell small, moderately solid, ovately cylindrical, yellowish ;

whorls 6, the first four convex, regularly increasing, the fifth less

convex and only slightly increasing in breadth, the sixth some-

what decreasing in breadth, sculptured with fine, very closely-set,

rather oblique, transverse riblets; suture impressed ; umbilical

area presenting the appearance of an elongate, strangulate

depression; columella white, very obliquely descending, out-

wardly expanded, extending above into a well-defined, parietal

callus which reaches the upper margin of the labrum; labrum

white, narrowly expanded and narrowly reflexed, except above,

where it is somewhat contracted ; aperture very obliquely sub-

quadrate, armed with a strong, curved, parietal lamella, a broad,

bifid, tooth-like projection rather interiorly situate on the inner

margin of the outer lip, and a very interiorly situated, lamella-

like, basal denticle.

Alt. 4, diam. maj. 2 mm.

Aperture: alt. 1, diam. -5 mm.

Hab. Mt. Nyiro, to the south of Lake Rudolph, at an altitude

of 8300 feet (4. Blayney Percival).

EwneA oprata Preston, var. oBESA, var. n. (Pl. XXXYV.

figs. 3, 3 a.)

Shell much broader and more ovate than the typical form

and with half a whorl less; moreover, the lower denticle below

the lamella on the inner margin of the labrum is almost obsolete.

Alt. 12, diam. maj. 6°5 mm.

Aperture: alt. 3°5, diam. 2°25 mm.

Hab. Mt. Kenangop, Aberdare Range, British East Africa

(ltobin Kemp).

* Rev. Zool. Africaine, Brussels, 1911, vol. i. p. 219, pl. xi. fig. 3.

+ Ann. Mag. Nat. Hist. 1911, vil. pp. 464-465, pl. xi. fig. 4.

912 MR. H. B. PRESTON ON AGNATHOUS

Var. MAJUSCULA, var.n. (Pl. XXXV. figs. 2, 2a.)

Shell much larger than either the typical form or the var.

obesa ; it is also more cylindrical than the latter, though the

dentition agrees perfectly with that variety.

Alt. 13°75, diam. maj. 6°5 mm.

Aperture: alt. 3°75, diam. 3 mm.

Hab. Mt. Kenangop, Aberdare Range, British East Africa

(ftobin Kemp).

The typical series from Mt. Kenia was collected in a dead

condition and appeared, though polished and shining, to be pure

white; both the present varieties were collected alive and are of

an olive-greenish hue, which is also doubtless the normal colour

of the typical form.

EINNEA PAPYRACEA, sp.n. (Pl. XXXV. figs. 4, 4a.)

Shell ovately cylindrical, white, very thin; whorls 8, not very

convex, the first seven regularly increasing, the eighth narrower,

sculptured with rather coarse, very oblique, transverse riblets,

and, on the last whorl, with fine, rather indistinct, spiral strie ;

suture very lightly impressed, strongly crenellated by the

termination of the transverse riblets; umbilical area presenting

the appearance of a very shallow, elongate fissure; columella

somewhat vertically descending, extending above into a thin,

parietal callus; labrum white, narrowly reflexed; aperture sub-

quadrate, armed with a parietal lamella, a small denticle, and

below this a large erectly squarish denticle just within the outer

lip, a small basal denticle and a broad projection on the columella

terminating in three denticles.

Alt. 11:75, diam. maj. 6 mm.

Aperture: alt. 3°5, diam. 2°25 mm.

Hab. Near Lake Kivu, Belgian Congo (Robin Kemp).

EINNEA PERGRATA, sp.n. (Pl. XXXV. figs. 6, 6 a.)

Shell rimate, rather large, elongately cylindrical, semitrans-

parent, whitish, shining; whorls 8, not very convex, the first

four regularly increasing, the remainder slowly increasing in

length but not in breadth, sculptured with irregular, very

oblique, transverse plicee, which are more noticeable in the

sutural region; suture impressed, coarsely crenellate below by

the terminations of the transverse plice ; umbilicus moderately

narrow; labrum white, shining, reflexed and outwardly ex-

panded, especially below; columella vertically descending, widely

reflexed above, narrowing below, bearing a coarse, projecting

denticle in the median region; aperture subquadrate, armed

with a coarse, erect, vertical, parietal plait, two denticles on the

inner surface of the outer lip, of which the uppermost is the

smaller, and a broad denticle at the base.

Alt. 12°5, diam. maj. 4°25 mm.

Aperture: alt. 3°5, diam. 1‘5 mm.

MOLLUSCA FROM EQUATORIAL AFRICA. 213

Hab. Mt. Elgon, Uganda (C. W. Woodhouse).

An extraordinarily cylindrical form which does not appear,

so far as is at present known, to have any close ally in Hast or

East Central Africa.

ENNEA PERVITREA, sp.n. (Pl. XXXV. figs. 8, 8a.)

Shell cylindrical, polished, shining, glassy, transparent whitish ;

whorls 74, the first four and a half regularly increasing in

breadth and length, the last three increasing in length but not

in breadth, the last rapidly ascending in front, quite smooth but

for occasional growth-markings ; suture very slightly impressed,

narrowly margined below; umbilicus very narrow, deep; colu-

mella obliquely descending, spreading above into a thin, somewhat

extended, sinuously margined, parietal callus; labrum white,

outwardly expanded and reflexed, especially below; aperture

semiovate, armed with a strong, curved, parietal lamella, a

moderately stout denticle on the inner margin of the outer lip,

and a broad, but less pronounced, denticle on the columella.

Alt. 6°75, diam. maj. 3°25 mm.

Aperture: alt. 1°75, diam. 1:25 mm.

Hab. Forests to the north of Mt. Kenia, British East Africa

(A. Blayney Percival).

ENNEA JOMBENEENSIS, sp.n. (Pl. XX XIII. figs. 8, 8a.)

Shell allied to #. pilula* Preston from Mt. Kenia, but of a

thinner texture, lighter colour, more cylindrical form, and having

a less outwardly expanded columella and less strongly developed

labrum ; nevertheless the subparietal lamella is more strongly

marked, and beside it occurs, on the parietal wall, a small, vertical

denticle, which is quite lacking in /. pilula; in addition to this

the present species possesses an extra basal lamella.

Alt. 12°25, diam. maj. 6°5 mm.

Aperture: alt. 2°75, diam. 1°5 mm.

Hab. Jombene Hills, British East Africa, at an altitude of

6000 feet (type); also taken at Mweru (Robin Kemp).

ENNEA POLLONERA, sp.n. (Pl. XXXV. figs. 9, 9 a.)

Shell cylindrically rectangular, whitish, somewhat solid ; whorls

64, the first two flattened above, the remainder flattened laterally,

the upper whorls regularly increasing, the last two decreasing in

breadth, sculptured with fine, oblique, transverse costule ; suture

almost linear, crenellated by the terminations of the transverse

costule ; umbilical area marked by a shallow, elongate depression ;

columella descending in a curve, extending above into a well-

defined, white, parietal callus which reaches the upper margin of

the labrum; labrum white, slightly polished, expanded and

veflexed; aperture subcircular, armed with a coarse, sinuously

curved, parietal lamella extending far into the imterior of the

* Ann. Mag. Nat. Hist. 1911, vii. p. 465, pl. xi. fig, 5.

214 MR. H. B. PRESTON ON AGNATHOUS

shell, two elongate lamelle on the outer wall, the upper of which

iherana hmeayyessf in a coarse, nodulous denticle on fe inner margin of

the upper portion of the labrum and a broad, bidenticulate

projection on the columella.

Alt. 12, diam. maj. 6°75 mm.

Aperture: alt. 2:5, diam. 2 mm.

Hab. Lake Kivu, Belgian Congo (Robin Kemp).

ENNEA PRETIOSA Preston, var. NYIROENSIS, var. n.

Shell differing from the typical form * from Mt. Kenia and the

surrounding region in its larger size and more rounded shape; it

also has one whorl more, a shallower suture, and more circular

aperture ; the last whorl is proportionately narrower towards the

base and the transverse costule in the large series before me are

less pronounced.

Alt. 5°75 (nearly), diam. maj. 3°25 mm.

Aperture: alt. 1:25, diam. 1:25 mm.

Hab. Mt. Nyiro, to the south of Lake Rudolph, at an altitude

of 8300 feet (A. Blayney Percival).

ENNEA ADJACENS, sp.n. (Pl. XXXII. figs. 8, 8 a.)

Shell differing from ZH. pretiosa Preston from Mt. Kenia in

its broader and more solid form, rather more circular aperture,

and in the transverse sculpture, which, in the present species, is

much finer, rather more oblique, and very much more closely set

except on the latter half of the last whorl; the labrum, which is

continuous, is also more broadly outwardly expanded than in

i. pretiosa.

Alt. 5, diam. maj. 3 mm.

Aperture: alt. °75, diam. *5 mm.

Hab. Between the Jombene Hills and Nyeri, British Hast

Africa (obi Kemp).

ENNEA SPATIUM, sp.n. (Pl. XXXYV. figs. 12, 12a.)

Shell allied to H. pretiosa, but differing from that species in its

much more distant, transverse costule, which are also much less

numerous and placed at about twice the distance apart from

one another than is the case with that species.

Alt. 4°25, diam. maj. 2°75 mm.

Aperture: alt. °75, diam. °75 mm.

Hab. Larogi Hills, British East Africa, at an altitude of

6000-7000 feet (A. Blayney Percival).

ENNEA MIRIFICA, sp.n. (PI. XXXIV. figs. 11, 11 a.)

Shell allied to #. pretiosa, but considerably broader with

whorls angled above, the transverse costule are more erect

except in the subsutural region and are exaggerated into knife-

* Ann. Mag. Nat. Hist. 1911, vii. pp. 465-466, pl. xi. fig. 6.

MOLLUSCA FROM EQUATORIAL AFRICA. 215

like projections at the angle of the whorls; the spaces between

the costule are quite smooth and glassy; the aperture is obliquely

elliptical, and the parietal lamella is larger, broader, and more

projecting, especially above; the basal denticle is almost obsolete

and the columella is devoid of armature.

Alt. 4°25, diam. maj. 3 mm.

Aperture: alt. 1, diam. °5 mm.

Hab. Mt. Nyiro, to the south of Lake Rudolph, at an altitude

of 8300 feet (4. Blayney Percival).

ENNNEA RENIFORMIS, sp.n. (Pl. XXXYV. fig. 11.)

Shell perforate, ovately cylindrical, yellowish white; whorls 6,

the first four regularly increasing, the next two increasing in

length but not in breadth, the last ascending considerably in

front, sculptured with coarse, oblique, transverse plicee which are

more noticeable in the subsutural region; suture impressed,

coarsely serrated by the terminations of the transverse plice and

considerably overhung in the latter half of the shell by the whorl

above; umbilicus rather wide, deep; columella obliquely de-

scending above, sharply curved below, extending into a thick,

somewhat well-defined, parietal callus, which reaches the lip

above and gives the appearance of a continuous labrum; labrum

white, sinuous, narrowly expanded and reflexed; aperture reni-

form, bearing a single short, oblique, parietal plait and a denticle

on the inner surface of the outer lip in the median part.

Alt. 6°5, diam. maj. (including the aperture) 4°5 mm.

Aperture: alt. 2°25, diam. 1 mm.

Hab. Mt. Elgon, Uganda (CO. W. Woodhouse).

A curious species, easily recognizable by the rather large

umbilicus and ascending last whorl, but especially by the kidney-

shaped aperture.

ENNEA SUAVISSIMA, sp.n. (PI. XXXYV. fig. 138.)

Shell imperforate, cylindrical. thin, transparent, cream-coloured,

polished, shining ; whorls 8, smooth, the first five regularly in-

creasing, the sixth increasing in length but little in breadth, the

last two flattish, also increasing in length but decreasing in

breadth; suture very lightly impressed, narrowly margined

below ; columella white, rather obliquely descending, somewhat

twisted, bearing a plait towards the median region and extending

above into a thin, well-defined, parietal callus, which reaches the

upper margin of the labrum and bears a short, curved, parietal

lamella; labrum white, thickened, very slightly expanded but not

reflexed, bent slightly inwards and thickened towards the middle

of the outer margin; aperture subrhomboidal.

Alt. 8, diam. maj. 3°75 mm.

Aperture: alt. 2°5, diam. 1:25 mm.

Hab. Mt. Nyiro, to the south of Lake Rudolph, at an altitude

of 8300 feet (A. Blayney Percival).

216 MR. H. B. PRESTON ON AGNATHOUS

ENNEA PERCIVALI, sp.n. (Pl. XXXV. figs. 5, 5a.)

Shell allied to #. suavissima, but differing in its much smaller

size, proportionately more elongately cylindrical form, more

convex lower whorls, and in possessing half a whorl more ; it is

narrowly perforate, and the columella, which is not twisted, is

somewhat broadly expanded above; moreover, the aperture is less

rhomboidal in shape than in /. swavissima.

Alt. 5°75, diam. maj. 2°5 mm.

Aperture: alt. 1°75, diam. °75 mm.

Hab. Mt. Nyiro, to the south of Lake Rudolph, at an altitude

of 8300 feet (A. Blayney Percival).

ENNEA VIATORIS, sp.n. (Pl. XXXV. figs. 14, 144.)

Shell large, cylindrical, pale yellowish brown; whorls 8, the

first five regularly increasing, the sixth increasing very little in

breadth, the last two slightly decreasing in breadth, sculptured

with closely-set, rather fine, oblique, transverse costule ; suture

lightly impressed ; perforation narrow, partly concealed by the

outward expansion of the columella; columella vertically de-

scending, outwardly expanded ; labrum somewhat broadly dilated

and narrowly reflexed except near the upper margin where it is

rather contracted ; aperture subrhomboidal, armed with a coarse,

oblique, erect, parietal lamella, two rather weak lamelle on the

inner margin of the outer lip, of which the lower is the coarser, a

‘basal denticle situate well within, and a rather broad, projecting,

tooth-like lamella on the columella.

Alt. 12°25, diam. maj. 5°75 mm.

Aperture: alt. 3°5, diam. 2 mm.

Hab. Mt Nyiro, to the south of Lake Rudolph, at an altitude

of 8300 feet (A. Blayney Percival).

ENNEA SYNGENES, nom. nov. for Hnnea consobrina Preston,

non Ancey.

Mr. J. R. le B. Tomlin having kindly pointed out to me that

the above name used by me in the ‘Annals and Magazine of

Natural History,’ vol. vii. 1911, p. 463, is already preoccupied *,

I now venture to propose the name “ syngenes” as a substitute.

ENNEA (PTYCHOTREMA) KOMBAENSIS, sp. n. (Pl. XXXIV.

figs. 4, 4a.)

Shell cylindrical, moderately solid, yellowish white; whorls 5,

the first three regularly increasing, the fourth proportionately

much longer, the fifth, though increasing in length, slightly

decreasing in breadth, sculptured with somewhat coarse, oblique,

transverse plications; suture impressed, crenellated by the

terminations of the transverse plice ; umbilical area represented

by a narrow, elongate, shallow depression ; labrum. continuous,

white, porcellanous, outwardly expanded, and, except along the

* “British Naturalist,’ 1892, p. 125.

MOLLUSCA FROM EQUATORIAL AFRICA.

parietal wall, reflexed ; aperture ovate, armed with an elongate,

curved, interiorly entering lamella on the parietal wall, a small

denticle and below this a very sinuous, internally entering,

elongate lamella on the inner margin of the outer lip, a basal,

interiorly situate lamella and an oblique, bifid projection on the

columella.

Alt. 7, diam. maj. 3:25 mm.

Aperture: alt. 2°25, diam. 1-25 mm.

Hab. Komba, 8.W. Uganda (Robin Kemp).

EXPLANATION OF THE PLATES.

PuatE XXXII.

Streptaxis desiderata, sp. n.

Figs. 1-la,

2-2 a.

3-3 a.

Figs.

Proc. Zoou. Soc.—1913, No. XV.

marsabitensis, sp. n.

UrGFUESSENSIS, SP. Ni.

percivali, sp. n.

woodhousei, sp. n.

Ennea aberdarensis, sp. n.

adelpha, sp. n. (type).

adjacens, sp. n.

arthuei, sp. n.

baccata, sp. n.

buccina, sp. n.

burungaensis, sp. n.

cared, sp. Nn.

claustrum, sp. I.

coeni, Sp. n.

copiosa, sp. i.

curvicolumella, sp. n.

XXXIIT.

Hunea decussatula, sp. n.

disseminate, sp. n.

elgonewsis, sp. 1.

eUSSOEHSES, SP. N.

Soveolata, sp. n.

Sunerea, sp. n.

hector, sp. 1.

Jombeneénsis, sp. Ne

tngeziensis, sp. n.

ingloria, sp. n.

WMnocens, Sp. N.

tridescens, sp. 0.

insulsa, sp. 1.

intradentata, sp. n.

Pirate XXXIV.

Hunea adelpha, var.

Bb)

kigeziensis, sp. n.

kivuensis, sp. n.

(Ptychotrema) kombaensis, sp. n.

laqueus, sp. 1.

lima, sp. n.

malasangiensis, sp. n.

margarita, sp. n.

masakaensis, sp. n.

mikenoensis, sp. ni.

mirifica, sp. 0.

monticola, sp. n.

mutandaensis, sp. n.

MWELUENSIS, SP. N.

218 MR. W. A. LAMBORN ON THE

PLATE XXXV.

Figs. 1-la. Ennead nyiroensis, sp. n.

2-2 a. » optata Preston, var. majuscula, var. nu.

3-3 a. FA Ap 5 » obesa, var. 1.

4-4 a. >» papyraced, sp. 1.

5-5 a. . percivali, sp. 0.

6-6 a. » pergrata, sp. i.

7-7 a. » perturbata, sp. n.

8-8 a. » pervitrea, sp. 1.

9-9 a. » -pollonere, sp. N.

10. » rectangularis, sp. i.

11. » vreniformis, sp. 1.

12-12 a. » spatium, sp. n.

13. 5 suavissima, Sp. i.

14-144 » viatoris, sp. i.

15-lia. ,, woodhousei, sp. n.

16. Notes on the Habits of Certain Reptiles in the Lagos

District. By W. A. Lamborn, M.R.C.S8.*

[ Received November 25, 1912: Read February 18, 1913.]

INDEX.

; Page

YAGTHING, GOUODOPUIMD roc 50g s50539 oncano ceo sodebs doanBeeccondssose! Calls}

Abricane Crocodiles ewer en secreteeee esate aetec ieee eee

I. On the Habits of the Lizard Agama colonorum, Daudin, and

on Native Superstition in regard to the Chameleon.

For some two years I have devoted spare time in Nigeria

to the pursuit of Entomology, and one of the questions in which

I have interested myself has been as to the palatability of

Lepidoptera.

To test this various animals have been employed, and one which

has been of great service to me is the common West African

lizard Agama colonorum. This is by far the most abundant

lizard in the district—a camp called Oni on the bank of a fresh-

water lagoon 70 miles east of Lagos and 10 miles from the sea,

and so I have had ample opportunities of making observations as

to its habits.

I must express my grateful thanks to Mr. G. A. Boulenger,

F.R.S., for advice and for his kind help in identifying the species

of the lizards by means of the specimens in the Natural History

Museum.

Agamas are found on high ground only, such as is always

above flood-level, and in open spaces, and they group together in

communities consisting of six or seven adult females headed by a

male. Jam strongly of opinion that these females look to one

lord and master only, and that every such family is to be found

day after day in one particular locality. This point might, of

* Communicated py Prof. E. 3. Pouttron, ¥.R.S., F.Z.S.

HABITS OF CERTAIN REPTILES. 219

course, have been settled by marking the females, but the difh-

culty has been that they are particularly wary and hard to catch.

I have satisfied myself, however, that any particular male can

always be found within the limits of a small area day by day, and

that the females keep within the same neighbourhood is indicated

by both sexes habitually seeking the same resting place night after

night.

During several months of the year 1912 a male lizard

invariably slept under the eaves of my bungalow, gaining access

thereto by climbing one particular pillar. On one occasion when

it sought the usual resting place I drove it down, but it returned

a few minutes later. I frightened it off twice again and even

then it came back, and finally I allowed it to climb up.

An old female, which I knew well because it had a bulbous

stump in the place of its tail, always entered a particular hole in

the wall of the bungalow at night-time. The young lizards

wander to a far greater extent, not attaching themselves to any

particular group, and I do not think that they seek nightly any

regular sleeping place,

The male lizard is a strict disciplinarian, and if one of his harem

happens to offend him he chases her and endeavours to inflict a

bite. ‘The females have in consequence a due respect for him, so

much so, that if some one drops a number of butterflies in the

vicinity the females, as a rule, wait for the male to take his fill

before attempting to satisfy their appetites, and if one of the

females does presume to attempt to share the meal, the male

becomes so filled with the desire to inflict vengeance that he will

desert the prey and chase the offender with the utmost activity,

often leaving the other females to test the meal.

The remarkable subservience of the females appears to be

eonnected with the great disparity in the numbers of the sexes.

The male’s responsibilites seem to be in excess of his capacities, so

that the females are forced to resort to various artifices to secure

their share of his attentions, This solicitation usually takes the

form of running up to the male, elevating the tail in front of

him and turning it to one side, but even then he frequently

responds by biting the importunate female. If she runs away,

however, he will sometimes pursue, and unless she allows herself

to be caught too readily, her end may be attained. I am sure

that this little manceuvre does act as an incentive to the male.

Previous to the act of pairing, the male mounts the female from

one side and supports himself by gripping the neck in his jaws,

her tail being raised and deflected to the opposite side. When

coupling has been initiated, the male releases the neck of the

female and takes up a position more or less ata right angle to her

while it is in progress. If disturbed, both of course endeavour

to rush away, and the male will often drag the female for a distance

of some feet, sometimes up a tree, before uncoupling results,

The process usually lasts two or three minutes.

The eggs, in a cluster of three or four, are deposited in the

15*

220 MR, W. A. LAMBORN ON THE

ground excavated by the female and are then covered with earth.

The choice of a situation to the liking of the female seems to be

a difficult one, for she will frequently make several burrows

without ovipositing in any. These are left open, with the little

heap of earth at one side.

As is to be expected, the males are exceedingly combative, and

the entry of one into the preserves of another usualiy leads to a

battle. The males eye one another from a distance, raising and

lowering the forepart of the body repeatedly at a slow rate. One

then approaches a few feet nearer to the other and rests again,

watching its adversary, and then after a long wait rushes up from

a distance of two or three feet to within a few inches of him.

The tail is the offensive weapon, and to bring it into action the

males take up a position parallel to each other but head to tail.

Each seeks to overcome the other not by a number of strokes but

by a single well-directed blow, and there is much manceuvring

with a view to obtaining a favourable position. Many of the

blows are ineffective, for the lizards dodge each other very skilfully,

but when a blow does get home it may be a powerful one, judging

by the resounding smack which follows. After a few such blows

one turns tail and the victor pursues him, with the result that a

further round may be fought out, though often the loser is simply

chased away. The males never are killed in their combats, and I

have never seen one male attempt to bite another.

There is more tendency for the females to bite each other, and

one frequently sees a large female running open-mouthed at a

smaller one, especially if the smaller one happens to have pro-

truding from its mouth a morsel too large to be readily swallowed.

Both males and females will bite when cornered if an attempt

be made to handle them.

The lizards are usually insectivorous, but when insects are

scarce, as in the dry season, they will not uncommonly eat vege-

table matter such as lettuce, tomatoes, and various indigenous fruits.

Their voracity is remarkable. On February 12th, 1912, I

dropped no fewer than eighteen of the Pierine butterfly, Phris-

sura sylvia, before a male lizard, and all were consumed within

ten minutes.

I have left behind me in Nigeria a snake 10 inches long which

had been seized by a lizard of this species. I did not personally

see the incident, but on returning to camp after a short absence

on June 29th, I was shown the bodies of the snake and lizard, and

was told by my colleagues that they had seen the lizard walking

backwards dragging the snake by its upper jaw. Still proceeding

backwards it began to climb a palm tree, the snake actively

wriggling all the time, and it reached a height of six feet. Un-

fortunately, at this point some barbarian instinct prompted one

of the men to throw a stone which injured the lizard so that

the snake was released and fell to the ground, where it was

despatched.

HABITS OF CERTAIN REPTILES. 221

Incidentally I may say that it is easy to hit a lizard with a

stone. They seem unable to judge its flight and make little

attempt to dodge it.

If one endeavours to approach them they run off at a rapid rate

towards the nearest wall or tree, as they are able to scale a rough

perpendicular surface readily.

When on the look-out for prey they mount a stone or other

peint of vantage whence a more ready view of the neighbourhood

can be obtained.

It is of course the common belief among the West Coast natives

as it is among country people even in England, that the lizard is

a venomous reptile. In connection with this point I have been

asked to take this opportunity to communicate with my own facts

an observation made by Mr. J. A. de Gaye, F.L.S., Science Master

at King’s School, Lagos, on native superstition in regard to

chameleons. He thus describes an incident which he noticed in

July 1912, at Onitsha, a town about 200 miles from the mouth

of the Niger :—

“ While walking in the long grass not far from the Mission I

came across a couple of chameleons. The natives are very much

afraid of that animal. It is true that when disturbed it opens its

mouth wide as if to bite, but it is quite harmless. There is a

curious superstition about chameleons in Nigeria. When I showed

my chameleons to my boy, he rushed back shivering with terror

and shouted, ‘ Massa, massa, this be plenty bad. If he go bite

you, he bite you all time. When thunder he dey come, then he

go fear, he go.’

“‘T could not help laughing at the stupidity of the superstition,

and did my best to explain how foolish it was to believe in such

nonsense. It was of no avail, the only reply was, ‘he be true, all

dem people dey go tell you so.’”

Anyone familiar with the mind of the West African native

will know that such incidents can be paralleled wherever the

chameleon occurs. I remember how when I first went to the

- coast I brought a chameleon home from the bush, and happening

to look into the kitchen to give an order as I was passing, I was

astonished to see the cook depart via the window, as I came in by

the door.

He was an Accra man, and as far as I could make out on

questioning him, his dread of the beast was based less on its

supposed poisonous powers than on its possession of powers

associated with the evil eye. This belief in the occult powers

of the chameleon is illustrated by another incident in my

experience.

I had a chameleon in captivity, and a Benin boy who acted as

my gardener came to beg it of me, giving as his reason that

he wanted it for the purpose of making juju, but with what

particular idea I was unable to ascertain.

MR, W. A. LAMBORN ON THE

Il. On the Habits of African Crocodiles.

My second note relates to the eggs of a crocodile found at the

camp already mentioned.

On about April 15th, 1912, my attention was drawn by a

native to croaking noises, much like the sounds made by our

English frogs at the breeding-season, emanating from below the

path at a spot about 6 feet from my bungalow, which is about

50 feet above water-level and about 100 yards from the lagoon.

The native told me that crocodiles’ eggs must have been

deposited there, and that when the young ones are about to

hatch out they invariably make these sounds so that the mother

may learn that they are ready and come to dig up the eggs.

There were no surface indications of any disturbance of the

ground, and as the path had been made of laterite crushed and

rolled hard to a cement-like consistence three years before, and as

there had been a daily stream of passers by on it all this time, I

was not inclined to believe his story. However, on digging with

a cutlass we found 13 eggs at a depth of about 18 inches. The

eggs were elliptical, measuring 67 mm. in length and 42°5 mm.

in maximum diameter. The croaking sounds did come from

these, for some of the young crocodiles croaked as I held the

eggs in my hand, and when these ceased others seemed to begin

croaking in turn. The sounds were to be heard at regular

intervals of five seconds.

All the eggs, except a bad one, were starred by fracture at

a particular spot, at the side towards one pole, though the shell

was still held in shape by the unbroken subjacent membrane, and

all the young crocodiles hatched within half an hour of being

dug up. ;

In four cases the head only came out, and as the young seemed

too feeble to complete the exodus we pulled them out, after waiting

a reasonable time.

The young crocodiles attempted to run off in the long grass

as soon as they hatched and showed active resentment at

any molestation, viciously snapping at the hand if one touched

them. They were placed in a bath, in shallow water changed daily,

and for about two weeks after hatching they croaked at night,

especially when rain was in the air, but after that they made no

sounds unless one approached after dark with a light, when an

oceasional croak might be heard.

They remained healthy and active, though without food, for

two months, and then a small fish was administered to each, and

when I came away in July they were feeding of their own accord

on small fish and large freshwater shrimps.

I have been informed by natives that the young crocodiles,

immediately after hatching, attach themselves to the dorsal fringe

on the tail of the mother and are thus conveyed by her to the

HABITS OF CERTAIN REPTILES. Los

water, but I have not been able to obtain really reliable evidence

on this point.

in connection with the whole subject I find a most interesting

reference in Livingstone’s account of his first expedition to

Africa.

In December 1853 he encamped on the banks of the river

Leeba, a tributary of the Zambesi, and his statement runs as

follows :—

““ We saw 60 eggs taken out of a single nest. They are about

the same size as those of a goose, but perfectly round. ‘The shell

is partially elastic from having a strong internal membrane and

but little lime in its composition.

“The spot was about 10 feet above the water, and the broad

path leading down to the river-side furnished evidence of its

having been used for a similar purpose in former years. The

dam after depositmg her eggs covers them up, and returns to

assist the young out of their place of confinement. Assistance

seems necessary, for besides the tough membrane of the sheli,

they have four inches of earth upon them.

‘“* They do not, however, require immediate food, because they

retain a portion of the yolk equal to that of a hen’s egg in a

membrane in the abdomen as a stock of nutriment. When this

is expended, the dam leads them to the water’s edge and lets them

catch fish for themselves.”

It will be noticed that Dr. Livingstone described the eggs as

being ‘‘ perfectly round.” The eggs which I saw were distinctly

elliptical, and the measurements recorded were taken with

callipers. I have seen still larger crocodile eggs which were also

elliptical. They were 80 mm. in length and 51 mm. in diameter,

and were found in a shallow nest covered with dead leaves and

other débris at the foot of a large tree near the water.

Prof. Poulton has kindly furnished me with a translation by

Mr. E. A. Elliott, F.Z.S., of a paper read by Dr. A. Voeltzkow in

1899 before the Berlin Academy of Sciences, “‘ On the development

and habits of Crocodilus madagascariensis on the East Coast.”

The translation of the section dealing with points that I have

mentioned runs as follows :—

‘“‘In the workroom (study) of my house I kept crocodile eggs

in some boxes filled with sand, which I had always under obser-

vation, in order to observe the emergence of the young reptiles.

One day I heard sounds issuing from one of these boxes, and

supposed that possibly a young one had emerged and had emitted

these sounds while being suffocated in the sand—which would

not have surprised me, as I had long known that the young are

capable of emitting sounds.

“On digging down the astonishing fact was revealed that the

sounds proceeded from unbroken shells.

“The sounds were so loud that if the eggs were exposed they

could be heard distinctly in the adjoining room. If, as in nature,

224 ON THE HABITS OF CERTAIN REPTILES.

the eggs are covered by 1—2 metres of sand, the sounds are duller

but still distinct at the length of a room. ‘This crying of the

young can be induced by walking heavily past the receptacle

containing the eggs, by knocking against it, by taking an egg in

the hand and turning it; in fact, any shock causes the young one

to lift up its voice in the egg.

“* As the female visits the nest almost daily in order to convince

herself of its orderly condition, her passage from the water to

the nest and back shakes the ground and induces the production

of sound by those young ones which are sufficiently developed.

Thereupon the old one scrapes the sand out of the pit and

presently the young emerge. When eggs from my boxes were

dug up and kept exposed immediately after the first sounds were

heard, the young emerged in about three days.

‘““The fact that the young give out sounds was unknown to

the natives, and no one believed me until convinced by actually

listening to (the sound in) the egg. 1 demonstrated this striking

discovery to the English Consul and to the Freneh officials at

Majunga.

“The sounds are produced with the mouth closed, apparently

with strong contraction of the ventral muscles sometimes, as we

produce hiccough sounds, and the tone is similar.”

In this paper there is also a reference to an observation by

Humboldt who, in the case of Crocodilus acutus, notes that when

incubation is complete the female returns and calls the young,

which answer her voice.

Dr. Voeltzkow states that there is nothing to show whether

this statement was founded on Humboldt’s own observations or

was simply repeated from native observation.

There isa little doubt as to the identity of the species observed

by me, but Mr. G. A. Boulenger, F.R.S., in a letter to Prof.

Poulton, states that he regards the common Lagos crocodile as

identical with C. niloticus. My species is the common crocodile

of the Lagos district, the largest I have seen measuring 9 feet

from snout to tip of tail, and I have no doubt that it is the

western form of niloticus; but as the habits here deseribed do

not appear to have been recorded from the West Coast and as

some of the facts may be new, I have felt it desirable to put my

own account on record.

Edwin ‘Wilson, Cambridge.

R.Broom, del.

MP. Parker, lith.

SCYLACOPS CAPENSIS.

Je, Zs. Se NOMS) NEAL OOS Wiil.

R Broom, del. Edwin Wilson, Cambridge.

M_P. Parker, lith.

SCYMNOGNATHUS TIGRICEPS.

ON MAMMAL-LIKE REPTILES, 225

17. On the Gorgonopsia, a Suborder of the Mammal-like

Reptiles. By R. Broom, M.D., D.Sc., C.M.Z.S.

[Received December 31, 1912: Read February 18, 1913. ]

(Plates XXXVI. & XXXVII.*)

InDEX. Page

Gorgonopsia, Affinities of the .............:0.00cce eens 229

SEQUGEODS GCHDGHSAD cos c00 asa 000 cecgocaganndnoconandzaseeneno PAS)

Scymnognathus tigriceps .........ceceec cee eeeeee reeves 227

When Gorgonops torvus was described by Owen in 1876 he

regarded it as the type of a distinct group, partly on account

of the apparent peculiarity of the nose, and also because the

temporal region was believed to be roofed. Lydekker in 1890

and Seeley in 1895 both agreed that the temporal region was

roofed, and Seeley proposed the name Gorgonopsia as a distinct

order for the reception of Gorgonops. A re-examination of the

type in 1909 convinced me that though the parietal region is

wide, Gorgonops has temporal openings like the other Therapsida,

and the discovery by the Rev. J. H. Whaits of a nearly perfect

skull of Gorgonops torvus shows that my observation was correct.

Still one has long had the feeling that Gorgonops diftered

considerably from the typical Therocephalians. For a time I

inclined to place it with Z%tanosuchus and others in the Dino-

cephalia. Later on | put it back among the Therocephalians.

Now the discovery of one or two perfect skulls shows that the

difference from the Therocephalians is sutlicient to warrant

the re-establishment of Seeley’s group—the Gorgonopsia—as a

distinct suborder of the Therapsida. Iam well aware that the

formation of so many new orders and suborders of reptiles as

have been made in recent years has been criticized ; but it seems

to me wiser to separate into distinct groups forms that are

manifestly distinct than to group into one order animals that

are markedly different; and one can now say without any fear

of contradiction that Gorgonops differs from the typical Thero-

cephalians more than does a Carnivore from a Marsupial. As

will be seen from the present paper, the skull in many ways

approaches more nearly to that of the Anomodont.

SKULL.

Though nearly perfect skulls are known of Gorgonops torvus

and Scymnognathus whaitsi, in neither species can the sutures be

very clearly made out. The discovery by Mr. 8S. H. Haughton

of two fine skulls from the Cistecephalus zone, and of a good skull

obtained by the Rev. J. H. Whaits from the Pareiasawrus zone,

gives us material which reveals practically every point in the

structure of the skull. |

* Foy explanation of the Plates see p. 230.

226 DR. BR. BROOM ON

Scylacops capensis (Pl. XXX VI.).

Broom, Annals 8. African Mus. 1913.

Scylacops capensis is pretty closely allied to Gorgonops torvus.

The figures given show most of the cranial characters. The most

noteworthy features seen in the figure of the side view (fig. 1) are

the large septo-maxilla, the large prefrontal which meets the post-

frontal and shuts out the frontal from the orbital margin, and

the great forward extension of the zygomatic portion of the

squamosal,

The upper view of the skull (fig. 2) shows some much more re-

markable characters. The large size of the prefrontal is again seen.

The frontals, though fairly large, do not reach the orbital margins.

Behind the frontal is a large postfrontal bounded by the pre-

frontal, the frontal, the parietal, and the postorbital. Between

the back parts of the frontals and in front of the parietals is

a, well-developed median preparietal. Though a preparietal has

long been known in most Anomodonts, it has not hitherto been

detected in any other group. It is certainly absent in typical

Therocephalians such as Scylacosawrus and Lycosuchus, and is

apparently not present in Dinocephalians, while no trace of it

exists in Cynodonts, nor has it been detected in Pelycosaurs. A

careful re-examination of the skull of Galepus jouberti seems to

show evidence of a small but distinct preparietal round the front

of the large pineal foramen. We may thus regard the presence

of a preparietal as a character of the Anomodontia. the Gorgo-

nopsia, and the Dromasauria. In Anomodonts and in Galepus

the preparietal always forms at least the anterior wall of the

foramen. In the Gorgonopsia the foramen is between the

parietals, and the preparietal is some distance in front. The

parietals are large, and each has a process which extends far

back and forms an important part of the bony wall which

separates the temporal fossa from the occiput. The end of the

posterior process is clasped by the squamosal. Posteriorly

the parietals are mainly bounded by the large interparietal. The

postorbital is very large. It forms the greater part of the post-

orbital arch and the whole of the upper margin of the temporal

fossa.

Whatever may be the case with the temporal fossa in the

Plesiosaurs, the condition of the bones in this skull will, I think,

convince everyone that the fossa in the Therapsida is the homo-

logue of the inferior fossa of. the Rhynchocephalia, as I have for

a number of years maintained.

The occipital view (fig. 4) of the skull shows the great develop-

ment of the squamosals and the large interparietal. The suture

between the basioccipital and the exoccipitals cannot be made

out, nor is there any evidence of a distinct supra-occipital. The

condyle is single.

The palatal view (fig. 3) shows many features of great interest.

The front part of the palate cannot be seen, and is restored from

other specimens. Though the skull of Scylacops capensis shows

MAMMAL-LIKE REPTILES. ONT

only the general appearance of the palatal elements when viewed

from below, our knowledge of the morphological structure has

been gained mainly by the skulls of Scymnognathus tigriceps and

Seylacognathus parvus. The settling beyond question of the

nature of the palate, and especially of the vomer, was a matter

of such importance that I have not hesitated to break up the

skulls of both of these types. As museum specimens they are

none the worse for having been broken across in various direc-

tions and internally dissected, and, as a result, every detail of the

internal anatomy is now known.

For many years I have known that the type of Gorgonops

torvus showed an apparently undivided median vomer, but whether

it was a true vomer or a pair of prevomers fused as we get in

Ornithorhynchus it was impossible to tell. In typical Thero-

cephalians there are two prevomers as in Lizards and Rhyncho-

cephalians. In Anomodonts and Cynodonts there is a single

unpaired vomer as in Mammals. A good many years ago (1895)

I endeavoured to maintain that the mammalian vomer was not

homologous with the reptilian paired vomers, but that these

latter were the homologues of the dumb-bell bone of Ornitho-

rhynchus. The question has been discussed by Gaupp, Versluys,

Fuchs, Osborn, Williston, and others, but may be said to be still

unsettled. As the Gorgonopsia stand between the Thero-

cephalians on the one hand and the Anomodontia and Cyno-

dontia on the other, it is in this suborder that we must look for

a solution of the vomer problem. Unfortunately, the skulls

examined still fail to give us the solution, as both Scymnognathus

and Seylacognathus agree with the Anomodonts, Cynodonts, and

Mammals, and differ from the Therocephalians in having a

median impaired vomer and, so far as can be seen, no trace of

paired vomers.

Scymnognathus tigriceps (Pl. XX XVIT.).

Broom, Annals S. African Mus. 1913.

In Scymnognathus tigriceps the basisphenoid is very large. It

has two greatly developed thick descending processes, which pass

down a considerable distance below the level of the condyle. In

front there is a deep but thin median keel, which passes forwards

and meets the median keel formed by the united pterygoids.

Above, the sphenoid passes between the pterygoids, and forms

a large median plate that extends forwards and upwards as far

as the plane of the front of the orbit.

The pterygoid is a huge bone which more closely resembles the

pterygoid of the Pelycosaurs than that of the.other South-African

Therapsid suborders. The descending pterygoid process is very

large, but the most remarkable feature is the enormous develop-

ment of the ascending plates. These pass upwards and clasp the

front of the median sphenoid. In front of the sphenoid they

become ankylosed, and form a median plate which extends

forward to meet the vomer. Whether this large, thin, median

228 ; DR. R. BROOM ON

plate is entirely made up of the fused pterygoids, or whether

there is a median basi-cranial element as well, cannot be made

out in the specimen. Certainly it is largely composed of the

pterygoid plates. In Scylacognathus the pterygoid plates are

separate from one another, and a median cartilaginous element

might have been between them. Further forward the median

plate in Scymnognathus is clasped by the ascending plates of the

palatines. In front of the median plate we meet with the vomer.

In Seymnognathus it clasps the front of the median plate, but in

Scylacognathus it is seen clasping a median element which is not

the fused pterygoids but may be regarded as ethmoid, a fact

which suggests that not improbably there is in Scymnognathus

an ethmoid element between the pterygoid plates, though no

sutures can be detected. ‘There can, I think, be no question that

the vomerine bone is the true mammalian vomer. I can find no

evidence of paired prevomers. The relations of the palatine

bones, as viewed from underneath, can best be understood from

the figure of the condition in Seylacops (Pl. XXXVI. fig. 3).

The mandible is beautifully preserved in Scymnognathus, and

the condition is thus better known than in any of the Therapsidan

suborders. The outer aspect, of course, and something of the

internal relations of the bones, are known in all; but there

are gaps in our knowledge even of the Anomodont mandible,

and only of the Gorgonopsian jaw is our knowledge practically

complete. Full detailed figures are thus given for later com-

parison with the other types (figs. 6 & 7).

The dentary is large and powerful. The front of the jaw

is deep and formed by the dentary, except the base, which

is splenial. About two-thirds of the outer aspect of the jaw

are made up of the dentary, and there is a well-developed

coronoid process. Only a little of the dentary shows on the

inner aspects, as in front the inner side of the jaw is mainly

splenial, and further back most of the dentary is hidden by the

angular, gonial, and coronoid. The relations of these elements

to the dentary will best be understood by the section and figures.

The splenial extends backwards to opposite the point where the

angular begins to replace the dentary on the outer side. The

angular is the second largest bone in the jaw, and extends nearly

the whole length. In front it is fitted in between the dentary

and splenial. Posteriorly it forms the main part of the jaw. As

will be seen in the drawing, there is a curious doubling of the

back part, forming a deep groove for some structure. A very

similar condition is seen in Pelycosaurs, though, so far as known,

not in other Therapsida. There is a distinct coronoid bone, as

shown in the figures, lying on the inside of the back part of the

dentary. The surangular is of fair size and, with the angular,

forms most of the back third of the jaw. ‘The articular is large

but short. It forms the articulation. It is clasped by the

angular and the gonial. It has a peculiar posterior process

which curves downwards and forwards. The gonial clasps the

MAMMAL-LIKE REPTILES. 229

inner side of the articular in front, and extends forwards between

the coronoid and the angular to a little beyond the posterior end

of the splenial.

POSTCRANIAL SKELETON.

The upper cervical vertebra are very similar to those in other

Therapsidans. The proatlas is large, and the upper portion

of the atlas is very similar to that in Anomodonts and Dino-

cephalians; and the condition further resembles that of these

suborders in the two halves being free and not forming a single

arch as in Cynodonts.

The shoulder-girdle has a large scapula without any distinct

acromion process. The coracoid and precoracoid resemble those

of the Therocephalians. There is a distinct cleithrum (at least

in Scylacops), a large clavicle and interclavicle, and an ossified

sternum.

The carpus has a large radiale and ulnare and a small-inter-

medium. There are two centralia, of which the outer is the

smaller. There are four distal carpalia, but the fourth is very

broad and, I believe, formed of the ankylosed fourth and fifth.

The digital formula is 2, 3, 4,5, 3; the third toe having a

small phalanx and the fourth two small phalanges as in the

Therocephalia.

Very little is known of the posterior half of the body.

AFFINITIES OF THE GORGONOPSIA.

Till we know more of the structure of the Therocephalia it is

impossible to say how far the Gorgonopsia differ. The differences

in the skull may be tabulated as follows :—

Gorgonopsia. Therocephalia.

1. Parietal region broad. Parietal region narrow.

2. A preparietal bone. No preparietal bone.

3. A large postfrontal. Postfrontal small or absent.

4, Postorbital large, forming the Postorbital small, not extend-

upper temporal margin and ing far back and not meet-

meeting squamosal above. ing squamosal above.

5. A single median true vomer. A pair of prevomers.

6. Transpalatine closely united ‘Transpalatine separated from

to: pterygoid. pterygoid by large foramen.

7. Mandibles united by powerful Jaws loosely articulated.

symphysis.

8. Angular with deep groove. Angular perforated.

Most of the characters in which the Gorgonopsia differ from

the Therocephalia are characters in which they agree with the

Anomodontia. The Therocephalia are unquestionably the more

primitive group, but there are some early characters in the

Gorgonopsia and also in the Anomodontia. Of course we only

know well one or two of the later Gorgonopsians, and we have

good reason to believe that the group is very early—earlier in

230 ON MAMMAL-LIKE REPTILES.

South Africa perhaps than the Therocephalia, and pretty certainly

earlier than the Anomodonts.

As so many of our types are founded on imperfect snouts, it

will be some time, probably long, before it will be possible to

divide the Gorgonopsians from the Therocephalians. All we can

do at present is to give a list of the known Gorgonopsians and of

those that are probably members of that group.

The known Gorgonopsians are Gorgonops torvus Owen, Scymno-

guithus whaitst Broom, Scymnognathus tigriceps Broom &

Haughton, Scylacops capensis Broom, and Scylacognathus parvus

Broom.

The following are probably Gorgonopsians :—TZtanosuchus

ferox Owen, Titanosuchus cloetei Broom, Scapanodon duplessisi

Broom, Archeosuchus cairncrossit Broom, '

Besides these there are quite a number of supposed Thero-

cephalian genera which, on further investigation, may prove to

be Gorgonopsians, such as Jnostransewia, Hriphostoma, Lyco-

saurus, Cynosuchus, Arctognathus, Ictidosaurus, and a number

of others.

Archeosuchus occurs in beds which are Lower Permian. Scyla-

cognathus and a number of other probable Gorgonopsians occur

in the Middle Permian or Pareiasaurus zone. Gorgonops and

Scymnognathus whaitst are from the Hndothiodon zone; while

Scymnognathus tigriceps and Scylacops capensis are from the

Cistecephalus zone or top of the Permian, From the very top of

the Cistecephalus beds a small undescribed Gorgonopsian has just

been discovered, so that we can confidently affirm that the Gor-

gonopsia extend throughout the Middle and Upper Permian beds,

and that they probably extend from Lower Permian to Lower

Trias.

EXPLANATION OF THE PLATES.

Ang. Angular. Art. Articular. B.o. Basioccipital. B.s. Basisphenoid.

Co. Coronoid, D. Dentare. H.o. Exoccipital. Fr. Frontal. J.P. interparietal.

Ju. Jugal. D. Lachrymal. Mx. Maxilla. Na. Nasal. P.A. Prearticular or

Gonial. Pa. Parietal. Pal. Palatine. Pm. Premaxilla. Po.F. Postfrontal.

Po.O. Postorbital. P.P. Preparietal, Pr,F. Prefrontal. Pt. Pterygoid.

Q. Quadrate, S.Ang. Surangular. Sma. Septomaxilla. Sq. Squamosal. S¢. Stapes.

V. Vomer.

PratE XXXVI.

Scylacops capensis.

VWie.1. Side view of skull. About % nat. size.

2. Top view of skull. A little over 3 nat. size.

8. Palatal view of skull. About $ nat. size.

4. Occiput. About } nat. size,

Prate XXXVI.

Scymnognathus tigriceps.

Fig. 5. Side view of skull. Nearly 4 nat. size.

6. Inner aspect of mandible. Nearly { nat. size.

7. Sections across jaw at the places indicated in fig, 6.

West, Newman Eth

DEIAPOMUS 2 Ur Sings

ee eS

PZ.S.1913. Pl. XXX

eee

aes

< ae rreminconiseeen ee oe

= ie Se es

Z OO Oe aa

cee oerer Nea e ye

errs ae

ss OT orn

SNS ecw nema

(de) em, FS =

ea en RESIST INN

West,Newman lith.

ARUNELEA SUBSeATe a

EAS VOUS. Palas

West, Newman lith.

JMB INI IVA SIE) SAIN SUA Si.

ON NEW BRITISH ENTOMOSTRACA. 231

18. On Two British Entomostraca belonging to he Orders

Copepoda and Ostracoda. By G. Srewarpson Brapy,

MEIDss IID, JDSoay Melee, Cail /Zasy

_ [Received December 20, 1912: Read February 18, 1913.]

(Plates XX XVIII.-XL.*)

InDEXx. Page

Diaptomus pusillus, Sp. MN. .....c.cce eee 23

The two species here described are the result of a recent re-

examination of material collected many years ago. ‘The first

named species occurred abundantly in a gathering made by the

Rev. Canon Norman, F.R.S., in Loch Ness, Scotland. The other

species is, unfortunately, represented by only one specimen taken

by myself near Arundei in Sussex.

Order COPEPODA.

DIAPTOMUS PUSILLUS, sp.n. (Pl. XXXVIII.)

Female.—Length 1mm. Bodyslender; seen dorsally the anterior

portion (cephalothorax) is quite thrice as long as broad, its anterior

extremity well rounded, posterior broadly truncate with acutely

mucronate lateral angles (fig. 3). Abdomen two-segmented (fig. 3),

the first (genital) segment twice as long as the following segment,

wide and bulbously dilated at the base, which is produced laterally

and bears two aculeiform sete; caudal rami not much longer

than broad, terminal sete very feebly plumose. Anterior antennz

slender and almost entirely destitate of marginal hairs, extending

when reflexed as far as the apices of the caudal setz (fig. 1).

Basal joint of the last pair of legs (fig. 4) somewhat dilated, its

outer margin prominent in the middle and bearing a short spine-

like seta; inner branch short, simple, inarticulate, truncated at

the apex, which bears two or three very minute setx ; outer branch

about three times as long as the inner branch, its last joint forming

a strong dagger-shaped claw to the dilated base of which is

attached a short spine. Ovisacs very small, spherical, containing

not more than one or two ova.

Male somewhat smaller than the female, length 0-98 mm.

Right anterior antenna slender, very slightiy dilated in the

middle (fig. 2), the 13th joint bearing a single strong spine, the

10th and 11th each a very small needle-like hair, the other joints

almost or entirely hairless; antepenultimate and penultimate

joints simple, elongated, last jomt small, bearing a few apical

filaments (fig. 6). Last pair of legs strongly developed (fig. 5),

the right much the larger and reaching beyond the extremities

* Wor explanation of the Plates see pp. 233-4.

939 DR. G. 8S. BRADY ON

of the caudal rami; no inner branch; distal joint ending ina

long faleate claw and bearing a hook-like spine on its external

margin ; foot of the left side reaching scarcely as far as the two

proximal joints of the right foot, imperfectly three-jointed (the

basal joint being imperfectly divided), second and third joints

small, tapering to a sharp point.

This interesting little species was found abundantly in a

gathering made by my friend the Rev. Canon Norman, F.R.S.,

many years ago (1885) and kindly given to me by him. It had

remained unnoticed, partly on account of its small size and partly

owing to its being mixed up with numberless other Copepoda and

Cladocera, until discovered on a recent re-examination of the

collection. My attention was drawn to it chiefly by the very

conspicuous dark coloration of the spermatophores, which were

attached in unusually great numbers to the females—frequently

in bunches of three or four together; the small spherical ovisacs,

also very deeply pigmented, formed another mark of recognition.

The gathering was from Loch Ness “off Drumnadrochit in the

open lake.” In several other gatherings made by Dr. Norman

during the same tour I have found no trace of this species.

Order OSTRACODA.,

ARUNELLA, gen. Nov.

Animal like Candona except that there are three pairs of

ambulatory legs in addition to the flexile fourth pair, which is

contained within the shell; also a pair of setiferous appendages

arising from the ventral surface of the body—probably near the

base of the first pair of legs.

ARUNELLA SUBSALSA, Sp. n. (Pls. XX XIX. & XL.)

Male.—Antennules six-jointed, rather sparingly setiferous

(Pl. XX XIX. fig. 1); antenne stout, six-jointed, destitute of

natatory set, no apical claw (fig. 2), but bearing a few stout sete,

antepenultimate joint with a pair of rod-like sensory appendages

(fig. 3); mandibles like those of typical Candone, mandibular palp

four-jointed, with a small 7-setose branchial plate (fig. 4). First

pair of maxille composed of a body with four digitate segments

to which is attached an elongated setiferous branchial plate as in

the normal Cypridide (fig. 5); second pair strongly prehensile,

each bearing a small setiferous appendage (Pl. XL. figs. 5, 6).

First and second pairs of feet four-jointed, rather densely hispid

with small adpressed hairs (figs. 1, 2), last joint with two

strong apical claws, penultimate joint with a fascicle of about

six spine-like sete ; the third pair of feet (fig. 3) destitute of

hairs but armed at the extremity with a long falcate claw;

fourth pair flexuous and similar to those of the normal Cyprids

(fig. 4). Arising from the ventral side of the abdomen, pro-

bably near the bases of the first pair of legs, is an appendage

NEW BRITISH ENTOMOSTRACA. 233

consisting of two slender, hispid, linear branches—unjointed, but

bearing long apical and lateral sete—with a nipple-shaped

prominence separating their bases (Pl. XX XIX. fig. 6). Caudal

rami slender, bearing three terminal sete, and one near the

middle of the posterior margin (Pl. XL. fig. 7); copulatory

organs of the usual complex type (fig. 8); ejaculatory duct encased

in a very dense and massive capsule (Pl. XX XIX. fig. 7) ; whorls

of the duct very indistinctly visible through the fibrous covering.

Female unknown.

Of this remarkable species I have seen only one example, a

male. It was found unexpectedly on a re-examination of some

material collected many years ago in ditches by the side of the

River Arun, near Arundel, the water being, no doubt, slightly

brackish. The shell was so much encumbered with muddy

débris that it could not be distinctly seen, my attention being

drawn to it by the remarkably strong projection of the male

organs below the margins of the valves. ‘The animal presents

characters intermediate between the Cypridide and Cytheride,

the three pairs of ambulatory legs corresponding to those of the

Cytheride, while the following pair and the post-abdominal

rami are like the similar structures in Cypridide. But there is

in addition to these limbs a remarkable biramose appendage

which was detached in the process of dissection, but was probably

derived from the body of the animal close to the anterior legs.

This structure I take'to be possibly homologous with one described

by G. O. Sars * under the name “ processus piliferus,” and stated

by that author to be found between the feet of the first pair in the

male of Bairdia subdeltoidea. Other noteworthy characters are

the presence of a pair of rodlike sensory appendages on the ante-

penultimate joint of the antenne, and the hispid clothing of the

anterior legs and “ processus piliferus.”

I regret very much that no further specimens of Arwnella are

to be found in my eollected material, but must hope that some

future collector may be fortunate enough to find it, and thus be

able to place the species on a more secure basis. It must

evidently be hunted on the muddy bottoms of the ditches, its

swimming powers being to all appearance non-existent.

EXPLANATION OF THE PLATES.

Pruate XXXVIII.

Diaptomus pusillus.

Fig. 1. Female seen laterally, X 84.

2. Male seen from below, X 84.

3. Abdomen of female, with attached spermatophores, x 140.

4. Foot of fifth pair, female, X 300.

5. D : male, X 140.

6. Distal joints of right anterior antenna of male, X 240.

* ‘Nye Bidrag til Kundskaben om Middelhavets Invertebratfauna, p. 128,

pl. xviii. fig. 12.

Proc. Zoou. Soc.—1913, No. X VI. 16

234 MR. R. LYDEKKER ON

Pratn XXXIX.

Arunella subsalsa, 3g.

. Antennule, X 140.

. Antenna, X 140.

Sensory seta of antenna, X 300.

. Mandible and palp, X 140.

. Maxilla of first pair and labinm, X 84.

. Processus piliferns, X 84.

. Portion of ejaculatory duct and sac, X 84.

WTO OE Ot

PuatTe Xb.

Arunella subsalsa, g cont.

Fig. 1. Foot of first pair q

A second pair ¢ X 84.

5 third pair

4 fourth pair, X 140.

Posterior maxilla of right side, < 140.

Ky a left side, X 140.

. Caudal ramus, X 84.

Copulatory organ, X 84.

aOR NO

19. The Dwarf Buffalo of Southern Nigeria; with a

Revision of the Dwarf Buffaloes of Western Africa *.

By R. Lypexker, F.R.S., F.Z.8.

[Received November 14, 1912: Read February 18, 1913. ]

(Text-figures 42-44.)

INDEX. Page

Geographical: Africa, new Dwarf Buffaloes ............... 234

Systematic: Bos (Bubalus) caffer hunti........0.....0-.. 240

Bos (Bubalus) caffer beddingtoni ......... 240

In 1910 I gave an account of certain Dwarf Buffaloes seen

and shot by Lieut. H. J. L. Thompson in the Yala country of

Southern Nigeria, between the Aloda towns of Echimoga and

Iveku-Okaku, in an open grassy country watered by tributaries

of the Yaké+. The adult bulls were described as almost wholly

black, and the cows as dun or khaki-coloured ; immature bulls

being also dun. As I could not obtain specimens of these

Buffaloes for the British Museum, I refrained from giving them

a name. Recently, however, Lieut. A. W. Hunt, R.N., has

brought home from Southern Nigeria heads of Buffaloes doubt-

less belonging to the same race, and as he has presented one of

these, a cow, to the Museum, the time has come for naming the

South Nigerian Buffalo.

* [The complete account of the two new races described in this paper appears

here, but since the names and preliminary diagnoses were published in the ‘ Abstract,’

they are distinguished by being underlined.—Ep1ror. |

+ See my book ‘The Ox and Its Kindred,’ p. 242, 1912; in line 10 from bottom,

cow should be bx17.

DWARF BUFFALOES. 235

Before proceeding further, it may be remarked that the study

of the Dwarf Buffaloes of Western Africa is beset with great

diticulties owing to the fact that the localities of the type

specimens of Bos nanus, B. planiceros, and B. centralis (all of

which are in the Museum) are unknown.

Taking first the case of B. planiceros, which was named from

a frontlet and horns of a very old bull, it appears that these are

racially inseparable from the younger pair of horns on which

B. centralis was based, and as there is some clue to the locality

where the latter were obtained, there is a presumption that the

former came from the same region. Now the type of B. centralis

was collected by a Mr. Dalton about the year 1854, and as

Mr. Thomas* has shown that at least some of Dalton’s specimens

came from the Gambia, there is a presumption that this may

have been the case with the Buffalo-horns. This presumption is

strengthened by the fact that these horns are indistinguishable

from those of Buffaloes killed in Gambia by Messrs. G. Fenwick

Owen and G. Russell Roberts, two specimens of which—a mounted

head and a skull, with horns—were presented by the former

gentleman to the Museum. This renders it practically certain

that the Gambian Buffalo is identical with B. centralis, which

appears to be inseparable from B. planiceros ; and I accordingly

apply the name B, caffer planiceros to the Gambian, or Sene-

gambian, Buffalo, with the proviso that if the type of planiceros

should be proved to be racially distinct from that of centralis—

which I believe to be an impossibility——the latter name would be °

available for the Gambian animal. The head from the Gambia,

presented by Mr. Owen to the Museum, is remarkable for the

bright orange tawny colour of the throat and upper part of

the chest, which, together with a patch of the same colour below

each ear, contrasts strongly from the slaty black of the rest of

the head. A similar bright orange tawny band characterises the

whole of the under surface of a cow from Sierra Leone (adjacent

to Southern Senegambia), which was living in the Zoological

Gardens at Antwerp in 1875; the colour of the upper-parts

being yellow—or rather, perhaps, khaki. This cow, which is

described and figured by Sir Victor Brooke in the Society’s

‘ Proceedings’ for 1875, p. 455, pl. liv., I accordingly take to

represent the female of B. c. planiceros*.

Passing on to Gray’s Bubalus brachyceros, or Bos caffer brachy-

ceros, as I prefer to call it, a difficulty has arisen with regard to .

the sex of the type skulls brought by Denham and Clapperton

from the Lake Chad district. Hitherto I have regarded the

bigger of the two as representing an adult bull and the smaller

a full-grown cow, but the acquisition by the Museum of a larger

series of heads of female West African Buffaloes leaves no doubt

* Ann. Mag. Nat. Hist. ser. 8, vol. viii. p. 121, 1911.

+ Also figured in Brehm’s ‘Tierleben,’ ed. 3, Saugetiere, vol. iii. p. 320, 1891; the

figure being reproduced in the ‘ Royal Natural History,’ vol. ii. p. 201. Brooke

incorrectly stated that it came from Senegal. a

236 MR. R. LYDEKKER ON

in my mind that both are referable to full-grown cows, as was

considered to be the case by Sir V. Brooke. The male of the

Lake Chad race I take to be represented by a frontlet and horns

(the latter in a worm-eaten condition) brought by Captain H.

Cock from Northern Nigeria, some distance to the east of

Kontago, or Kontagora, and presented by him to the Museum

in 1904. The Dwarf Buffaloes met with during the Alexander-

Gosling Expedition* from Nigeria to the Sudan in the Shari

Valley and on an island in the upper Ubangui (a skull of one of

which from the locality last named is in the Museum) likewise

appear to be referable to this race. Two cows are described as

being respectively rich and dull tawny, with black legs, black

fringes to the ears, and black tail-tips; while the one bull was

dull tawny with similar black points. Dwarf Buffaloes shot by

Gen. P. 8. Wilkinson in Northern Nigeria on the Benue River,

near Abiusi, have been noticed by Mr. Pocock}. Of these an old

bull was jet-black, while an apparently younger animal of the

same sex 1s described as red, which is no doubt the equivalent of

the rich tawny in the account given in Boyd Alexander’s book

of the male buffalo from the Ubangui, which may likewise have

been comparatively young. The cows of the Benue herd appear

to have been all red or bright khaki-coloured.

Here it may be mentioned that the head of a Buffalo from the

interior of French Congo, described and figured by myself on

page 996 of the Society’s ‘Proceedings’ for 1910 as that of an

immature male (as it was stated to be by the donor), is really that

of an adult cow, as is indicated by the condition of the bases of

the horns, which, like those of the type specimens of B. c. brachy-

ceros, are fully formed and incapable of further growth.

From the cow of the Gambian race this head (text-fig. 42)

differs by the absence of an orange tawny area on the throat and

the more rufous colour of the hair, as well as by the much more

upward direction and greater incurving of the horns. In the

latter respect it is more like the cow, figured on page 321 of the

above-mentioned volume of Brehm’s ‘Tierleben,’ and said to be

from the Congo, but differs by the much larger amount of black

on the ear-fringes. The associated body-skin is wholly rufous,

with the exception of a black dorsal stripe; the shanks being

also dark. Unfortunately it is not known from what part of the

interior of French Congo this Buffalo was obtained; but if it

came from a place well to the east, there seems no reason why

it should not be within the distributional area of B. c. cottoni,

to which race it was provisionally assigned in the original

description. It is possible, however, that it may belong to

B. c. nanus.

As regards B. c. nanus, I take this race, which is typified by

the well-known frontlet and horns in the Museum, to be also

* See Boyd Alexander, ‘ From the Niger to the Nile,’ vol. ii. p. 394, 1907.

+ ‘The Field,’ vol. cxix. April 5, 1912.

DWARF BUFFALOES. 237

Text-fig. 42.

x Oe

AOA

Head of Cow of Bos caffer cottoni (?) from the interior of French Congo.

(Reproduced from P. Z.S. 1910, text-fig. 148, p. 996.)

Text-fig. 43.

Head of Bull of Bos caffer hunti from Southern Nigeria.

238 MR. R. LYDEKKER ON

represented by the cow and young bull from the Congo, figured

on page 321 of Brehm’s ‘Tierleben,’ ed. 3, Siugetiere, vol. iii.

p. 321 *, which are described as being wholly yellowish, with the

exception of the brownish limbs. A full-grown bull of this race

I have never seen.

The ground having been thus cleared, it is now practicable to

discuss Mr. Hunt’s specimens, as represented by the mounted

heads of an adult bull and cow. In the former of these (text-

fig. 43) the general colour of the skin and hair is best described

as dusky, since it is not anything like black. Compared with

the aforesaid head of B. c. planiceros in the Museum, it shows

a much smaller amount of bright orange tawny on the throat and

upper part of the chest, and no upward extension of this on each

side of the head towards a similar patch at the root of the ear,

this patch being absent in the South Nigerian head, in which

the general colour is also lighter. The fringe of long hair on

the upper edge of the ear is reddish orange, but that on the

opposite margin mainly dark. The horns, in addition to being

considerably smaller, differ from those of the male of the Gambian

Buffalo in that their front surface slopes regularly backwards

almost in one plane, instead of curving at first backwards, then

forwards, and finally backwards. The following are the dimen-

sions, in inches, of the Gambian and the South Nigerian horns,

together with those of the Alexander-Gosling skull from the

Ubangui referred to the Lake Chad race :—

Lake Chad. Gambian. S. Nigerian.

Outside length ............... 254 274 204

iRallime breadit lie eres -eeaeee ee 63 a (ci

Tip-to-tip interval ......... ne 212 12

In the head of the South Nigerian cow (text-fig. 44) the colour

is dull tawny, with a dusky tinge on the lower part of the face,

but no bright orange on the throat; the frmge being bright on

the upper margin of the ear, as well as in two flecks lower down,

but elsewhere dark. The front surface of the horns lies wholly

in one plane, instead of curving backwards as in the case of

B. c. brachyceros. The following are the measurements of the

horns in the larger of the two type specimens of the latter and of

the specimen now under consideration :—

Lake Chad. S. Nigerian.

Outside den etli is epee ee eter 174 12

iPalln= breadth) ey 25. see eee eee 4 32

Tip-to-tip interval ve... eee: Gi 94

The foregoing measurements of the horns of the South

Nigerian bull and cow agree, respectively, very fairly with those

of specimens killed by Major A. J. Arnold in the Lokoja district

* The figure is reproduced in the ‘ Royal Natural History,’ vol. 1. p. 208.

DWARF BUFFALOES. 239

of Nigeria, at the junction of the Benue and the Niger, and

quoted by myself * :—

Bull. Cow

Outsidetlens thay v4.0) 43:25 see eed. nee 193 123

IBzailinne wate blit ayrst ws cht. a ane aera eyelet = 43

hip-tosbijpuabernval: Jose ssaeees. sees eee 15 11

These Buffaloes, together with an immature mounted specimen

in the Museum, from Lokoja, although hitherto referred to

B. c. nanus, must, from their locality, be assigned, I think, to

the same race as Mr. Hunt’s specimens. Major Arnold states

that ‘their colour varies from the light red of the younger

animals deepening through the warm rufous red of the mature

beast, to the deep dirty brown of the aged bull.” f

Text-fig. 44.

Head of Cow of Bos caffer hunti from Southern Nigeria.

The above-mentioned female head from the French Congo

agrees with the South Nigerian cow in its generally rufous

colour, but differs by the more abundant and mainly black

fringes to the ears, as well as by the middle portion of the

front surface of the horns being distinctly curved in a backward

direction.

As the result of the foregoing comparisons and descriptions,

it appears that the South Nigerian Buffalo represents a relatively

* “Game Animals of Africa,’ p. 84, 1907.

+ ‘Game Animals of Africa,’ loc. cit.

240 ON DWARF BUFFALOES.

small race of Bos caffer in which old bulls are often dusky coloured

or blackish, with a well-defined bright orange tawny area on the

throat and chest, while adult cows are khaki-coloured, with an

orange fringe on the upper margin of the ears, and a dusky

tinge on the middle of the lower part of the face; the horns in

both sexes being relatively small and lying mainly in one

plane.

That these South Nigerian Buffaloes are racially distinct from

B. c. planiceros and B. brachyceros seems abundantly clear; their

relationship being nearer to the former than to the latter.

They are also distinct from B. c. simpsoni, in which both sexes

are blackish.

That they differ from B. c. centralis, as represented by the type

horns and the two animals figured by Brehm, seems sufficiently

apparent ; and their horns are likewise different in type from

those of Dr. Matschie’s B. c. thierryi from Togoland.

On distributional grounds the South Nigerian Buffalo is un-

likely to be identical with B. ¢. cottoni of the Semliki, and perhaps

the interior of the French Congo; but to remove any doubt on

this point it may be mentioned that in the horns of the type

bull of that race (which came from Kasindi, at the north-west

corner of Lake Albert Edward, and is preserved in Major Powell

Cotton’s Museum at Quex Park, Birchington) the outside span

is 332? inches, against 21 in Mr. Hunt’s Nigerian bull, while

the palm-width in the former is 10 inches, against 7} in the

latter. Moreover, the Semliki bull stood 57} inches at the

shoulder, whereas an adult Nigerian bull shot by Major Arnold

measured only 44 inches. B. c. cotéoni cannot therefore be

termed a Dwarf Buffalo.

As the South Nigerian Buffalo cannot apparently be identified

with any of the named races, I propose to call it Bos (Bubalus)

caffer hunti |P. Z.S. Abstract, 1912, p. 6 (February 25th)], taking

the head of the cow presented to the Museum (text-fig. 44)

as the type.

The occurrence of this race of Buffaloes in Southern Nigeria

renders it practically certain that an adult bull Buffalo from

Ashanti presented to the Museum by Mr. O. Beddington in 1900,

and hitherto identified with B. c. nanus, cannot belong to a race

inhabiting the Congo. Since, moreover, the fringe of hair

on the ears is much less abundant, and the horns are con-

siderably less closely approximated at the tips than in the type

of nanus, I propose to make this bull the type of a new race,

under the name Bos (Bubalus) caffer beddingtoni | P. Z.S. Abstract,

1912, p. 6 (February 25th)]. The height at the shoulder is

514 inches, the outside length of the horns 203, the palm-width

63, the tip-to-tip interval 133, and the outside span 233 inches.

There is no dark dorsal stripe, which apparently occurs in nanus.

In conclusion I give the following tentative “key” to the

named races of West African Dwarf Buffaloes in which the colour

of the coat is known :—

DR. S. F. HARMER ON A HAIR-BALL. 241

A. Both sexes dark; horns regularly curved in a subcircular

form.

Size medium ............ B. c. sinpsoni.

B. Adult bulls often dark, young bulls eae cows ay or eee g

horns more expanded laterally.

. Throat of bulls orange tawny.

a’. Size large ; throat of cows orange ... B. ¢. planiceros.

. Size smaller ; throat of cows coloured ‘like ‘neck ;

horns mainly in one ee ear-fringe of cows

partly orange ..... Geb ooa DOR REG HCO AELIDOUAaaae OH I Aee asa CGE COMA Is

6. Apparently no orange on 1 throat.

Size medium; horns strongly bent backwards; ear-

fringe of cows mainly black ...................:0-::.:... B. ec. brachyceros.

C. Adults of both sexes rufous or ae ons more or Tee

angulated and flattened.

a. Size small; horns strongly angulated; heavy ear-

fringes ; apparently a dark dorsal stripe ............... B. c. nanus.

6. Size apparently larger; horns less strongly angulated ; ¢

less abundant ear-fringes; no dark dorsal stripe ...... B. ce. beddingtoni.

[Since the above was written I have seen specimens which show

that old bulls of B.c. hunti may retain the red coat while younger

ones may be dusky. |

EXHIBITIONS AND NOTICES.

March 4, 1913.

Dr. A. Smita Woopwarp, F.R.S., Vice-President,

in the Chair.

Dr. P. CuAtmers Mircuety, F.R.S., F.Z.8., Secretary of the

Society, exhibited a lantern-slide made from a photograph of a

domestic Donkey (Zquus asinus) taken by Captain C. H. Armitage

in the Gold Coast, and showing a very remarkable pattern of

transversely arranged black stripes on the flank.

Mr. EK. G. Bovutencer, F.Z.S., Curator of Reptiles, made

remarks on the Lung-Fish, Protopterus ethiopicus, which the

Society had recently received from Mr. C. W. Woodhouse, and

which was the only Dipnoan that had not previously been

exhibited alive in Europe.

Dr. 8. F. Harmer, M.A., F.R.S., exhibited a Hair-ball which

was presented to the British Museum (Natural History) by

Mr. A. Dobrée, in January 1911. Alluding to Mr. E. G.

Boulenger’s exhibit, on February 18th last, of spines of a Mada-

gascar Insectivore in the excrement of.a Boa, he pointed out that

this was perhaps another case of the occurrence of spiny hairs in

the alimentary canal of a reptile. Mr. Dobrée’s specimen, which

has been described by Mr. R. Lydekker, F.R.S., in ‘The Field,’

vol. 117, Feb. 25, 1911, p. 383, was found in the Province of

Morondava, West Coast of Madagascar, between Beld on the

242 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

Tsiribihina River and Bevilé; and it has the form of a hair-ball,

consisting largely of the hairs of Tenrecs (Centetes). Bearing

Mr. Boulenger’s remarks in mind, it is possible that the specimen

in question may have come from a Boa; but it seems to be more

probable that it came from the stomach of a Crocodile. The

principal reason for coming to this conclusion is the evidence

supplied by Mx. J. Simao da Costa to the British Museum that

hair-balls of this nature occur in the stomachs of Crocodilians

in N. Brazil, as recorded by Mr. Lydekker in the note above

referred to.

March 18, 1913.

E. G. B. MEADE-Watpo, Esq., Vice-President,

in the Chair.

The SrecrRErARY read the following report on the additions

that had been made to the Society's Menagerie during the month

of February, 1913 :— .

The registered additions to the Society’s Menagerie during the

month of February were 151 in number. Of these, 59 were

acquired by presentation, 27 by purchase, 45 were received on

deposit, 2 in exchange, and 18 were born in the Gardens.

The number of departures during the same period, by deaths

and removals, was 153.

Amongst the additions special attention may be directed to :—

1 Fishing Cat (felis viverrina) 3 ,from Meerpore, presented by

J.S. E. Walker, Esq., on February 15th.

1 Pigmy Hippopotamus (Cheropsis liberiensis) 2 , from Liberia,

new to the Collection, purchased on February 6th.

1 Isabella Gazelle (Gazella isabella) 2 ,from Suakin, and | Persian

Gazelle (Gazella subgutturosa) 3 , from Persia, presented by Oswald

Darke, Esq., on February 14th.

1 Mikado Pheasant (Calophasis mikado), from Mount Arizan,

Formosa, new to the Collection, deposited on February 25th.

2 White-eyebrowed Finches (Sporophila superciliaris), from

Brazil, new to the Collection, purchased on February 5th.

1 Guatemalan Ouzel (Merula infuscata), 2 Cinnamon Tinamous

(Crypturus cinnamomeus), from South America, 3 Lesser Scaup

(Fuligula afinis), 1 Ringed Duck (Fuligula collaris), from North

America, new to the Collection, deposited on February 25th.

1 Kyed Gecko (@dura ocellata), from Australia, new to

the Collection, presented by C. R. Walter, Esq., F.Z.5., on

February 8th.

_ 1 Helen’s Snake (Coluber helena), from Ceylon, new to the

Collection, purchased on February 10th.

6 Argentine Dwarf Toads (Phryniscus nigricans), from Monte

Video, new to the Collection, presented by W. Parkinson Curtis,

Esq., on February 11th.

2 Diamond-Bass (Hnneacanthus gloriosus), from N. America,

new to the Collection, deposited on February 25th.

ON REPTILIAN TAPEWORMS. 243

PAPERS.

20. Contributions to the Anatomy and Systematic Arrange-

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

D.Se., F.R.S., F.Z.8., Prosector to the Society.

[Received January 14, 1913: Read March 4, 1913.]

1X. On a NEw GENuS oF [CHTHYOTANIIDS.

(Text-figures 45-53.)

InDEX.

Page

Account of Solenotenia viperis, gen. et sp. 0. .................. 2438-261

Résumé and General Conclusions...................0e0eceeeeeeee eee 261

A considerable number of examples of a rather large species of

tapeworms clearly referable to the family Ichthyoteniide was

obtained in 1911 from the Crossed Viper (Lachesis alternans). It

is only recently that I have subjected these specimens to a careful

examination, and they prove to represent a species which is most

nearly akin to my recently constituted genus Ophidotenia*, but

present characters which seem to me to justify their inclusion

within a new genus, which I term, by reason of a salient external

character of the ripe proglottids, Solenotenia. This character is

obvious in fully mature proglottids (see text-fig. 49, p. 250) as

a well-marked ventral groove extending along nearly the whole

of the proglottid on the ventral surface. It is generally quite

visible to the naked eye, when it appears to vary in length in

different proglottids, being in fact a mark of maturity.

The general aspect of this tapeworm is that of other Ichthyo-

teniids, and it is quite like Zchthyotenia gabonica, which I have

recently described, so far as external characters go, with the

exception, of course, of the ventral groove, to which I have just

referred. The length is at least 5 or 6 inches, and the breadth

of the ripe proglottids rather more than 2mm. The body does

not narrow much at the head end, and the scolex is more than

1 mm. wide. There is a neck which is at first narrower than

the scolex. In transverse sections the scolex showed no pecu-

liarities that I could detect. There was hardly any rostellar

region, the suckers being very large.

In addition to the four large suckers (which are of course

unarmed, like those of other Ichthyoteniids), there is a very

small apical structure of a somewhat different nature. It is less

cup-shaped and more funnel-shaped than the lateral suckers,

narrowing rapidly at its internal end. At the same time the

* See P. Z.S. 1913, p. 26.

244 DR. F. E. BEDDARD ON

minute structure of this apical organ seems to indicate that it is

to be regarded rather as a sucker than a rostellum. The apical

organ of Acanthotenia*, on the other hand, is, as I think, of the

nature of the rostellum of other tapeworms.

Text-fig. 45.

Transverse section through scolex of Solenotenia viperis.

S. One of four large suckers. s. Small apical sucker showing its minute

size in comparison with large suckers.

The accompanying figure (text-fig. 46) shows several out of

a series of transverse, slightly oblique, sections through this

apical sucker. Being rather oblique they show the external

surface, which is slightly concave. The nuclei, which are chiefly

towards the external surface of the sucker, seem to me to be

rather larger than those in the four lateral suckers. Previous

investigations upon this apical structure are not entirely con-

clusive. Riggenbach } finds an apical depression in [chthyotenia

* P. Z.S. 1913, p. 8, text-fig. 1, R.

+ “Das Genus Ichthyotenia,” Rev. Suisse Zool. iv. 1897, p. 165.

REPTILIAN TAPEWORMS. QA5

fossata, which his figures * show to be, relatively to the lateral

uckers, very much smaller than in Solenotenia viperis. But he

is not able to be decisive as to its minute structure or nature—

that is, whether sucker or rostellum. He quotes the opinion of

Monticelli t that the organ in question is a sucker, and points out

that Lithe = seems to regard it as a rudimentary rostellum.

Text-fig. 46.

Three nearly consecutive sections through minute apical sucker of

Solenotenia viperis.

A. Ata little distance from the surface. B. At the actual apex of the scolex.

C. Near to deep end of apical sucker.

This structure is figured in Jchthyotenia ocellata by Kraemer

in a memoir upon Tvenias from fish §, who regards it as a sucker.

* Loe. cit. pl. vii. figs. }, 3, 4.

+ Boll. Soc. Nat. Napoli (1) v., 1891.

ft Zool. Anz. 1894,

§ Zeitschr. wiss. Zool. liii. 1892, pl. xxviii. fig. 40.

246 DR. F. E. BEDDARD ON

In tabulating the characteristics of the fish Teenias, Kraemer

writes thus: * “‘Der Mangel eines Rostellums, anstatt dessen

Text-fig. 47.

Upper figure a transverse section through part of a proglottid of Solenoteni

viperis ; lower figure a longitudinal section through part of a proglottid.

1. Longitudinal muscles. ¢. Transverse muscles.

* Zeitschr. wiss. Zool., loc. cit. p. 718.

REPTILIAN TAPEWORMS. OAT

hiiufig ein den Scheitel des Scolex einnehmender, kleiner fiinfter

Saugnapf entwickelt ist. Das Vorhandensein eines scheitel-

stiindigen Saugnapfes ist wohl ein urspriingliches, den niederen

Formen der Tanien eigenes Verhalten.” The balance of opinion,

therefore, seems in favour of regarding an apical organ as charac-

teristic of the Ichthyotentidex, but to leave it for the present

doubtful whether we are to describe this organ as a sucker or as

a rudimentary rostellum. It appears to me that there is evidence

in favour of believing that both structures are present among

the members of the family Ichthyoteniide, and that Acanthotenia

is to be characterised by the possession of a rostellum while

Solenotenia has an apical sucker. But it may be that both

structures have a common origin.

In transverse sections of the neck region muscular fibres,

forming a longitudinal layer, are visible. But these fibres do

not form a compact layer ; they are scattered singly through the

parenchyma, and not massed into bundles, as in Acanthotenia,

or into a definite layer, as in some other Ichthyoteniids. Further

back the fibres of the longitudinal layer are arranged in bundles ;

these bundles are small, consisting of but few fibres and with

considerable interspaces, where there are no longitudinal fibres

delimiting the cortical from the medullary layer. The position

of the bundles, and therefore the length of the interspaces, are not

regular, and the fibres of each bundle (in which I have counted

about 4-7) are apt to be placed one on top of the other, thus

forming a line at right angles to the long axis of the transverse

section. Longitudinal sections through the proglottids illustrate

also the scattered condition of the small bundles of longitudinal

fibres ; for there are to be seen frequent gaps where the cortical

layer is not separated from the medullary layer by these fibres.

There are apparently, as will be seen in text-fig. 47, transverse

muscular fibres in the medullary layer.

The generative system of Solenotcenia is like that of other

Ichthyoteniids, with certain peculiarities of its own and features

that it shares only with Ophidotenia. The accompanying text-

figure (text-fig. 48) is a view of a mature proglottid mounted

entire, and it shows the general arrangement of the various

parts of the generative system. The marginal generative pores

lie quite at the middle point of the segment, and the cirrus is

seen to be occasionally protruded. As in other Ichthyotzniids,

the male pore is sometimes in front of and sometimes behind the

vaginal pore.

The witelline glands ave very plain in such preparations, and

form the usual strips extending through the greater part of the

length of the proglottid. Marotel * has represented the vitelline

glands of Ichthyotenia calmetiei as consisting of numerous and

separate follicles. I find in the present species in certain regions

an obvious separation between the adjacent follicles ; elsewhere,

* Arch. de Parasitol. 1899, p. 34,

248 DR. F. E, BEDDARD ON

on the other hand, the individual glands are connected into a

more continuous mass. This is shown in the accompanying

figure (text-fig. 48).

Text-fig. 48.

View of an entire proglottid of Solenotenia viperis stained and mounted

as a transparent object.

g, opening of cirrus-sac; 2, opening of vagina.

o. Ovary. wu, Uterus showing open groove on surface of body and irregularly paired

diverticula. v.gl. posterior region of vitelline gland, which is of greater

diameter than the anterior region and which gives off one or two processes

towards the middle line of the body. v.g1.’ Anterior thinner region of the

vitelline gland.

The same text-figure also shows a peculiarity which is ap-

parently, at least, not universal among these tapeworms.

Towards the posterior end of the proglottid the marginal strip

of vitelline gland gives off one or two processes of vitelline tissue

which pass to the interior of the proglottid in the direction of the

REPTILIAN TAPEWORMS. 249

ovary. These ingrowths of the vitelline tissue are limited to the

posterior region of the gland. Furthermore, the preparation

which is represented in the text-figure shows that the vitelline

gland is distinctly thicker posteriorly in the region of the ovary,

and that it tapers somewhat anteriorly. These apparently minute

facts are Important when we consider the characteristics of the

vitelline glands among the Cestoidea. Among the Tetracotylea

(=Cyclophyllidea), to which group Jchthyotenia and its imme-

diate allies have been by some referred, the vitelline gland is —

single and compact, and lies nearly always behind the ovary,

7.e. quite posterior in the segment, but always near to the ovary.

On the other hand, the Tetraphyllidea, to which Jchthyotenia

and its allies are more generally referred, have strip-like vitelline

glands situated laterally.

At first sight Ichthyotzeniids appear to conform entirely with

the latter definition. It seems to me, however, that Solenotwnia

shows quite intermediate characters. The thinning of the

vitelline strip anteriorly, and the projections posteriorly directed

towards the middle line of the proglottid, if they were carried

farther—.e. to the disappearance of the anterior part of the

glands and the junction of the posterior regions—would produce

the vitelline gland characteristic of the Tetracotylea. Another

matter in reference to the vitelline glands remains for con-

sideration. Riggenbach*, in his account of the anatomy of

Ichthyotenia abscisa, distinctly states that the vitelline strips lie

outside of the water-tubes and the nerve-cord, while Schwarz f

writes absolutely to the same effect of 7. nattereri: ‘ Die Dotter-

stécke legen wie bei allen Ichthyotenien ganz randstiindig,

ausserhalb der Nerven und der Liingsgefaisse und reichen vom

vordern Gliedrande bis zum hintern.” This obviously implies

their position in the cortical parenchyma.

In defining the Tetraphyllidea, in which group he includes the

Ichthyotenide as a family, Prof. Max Braun t writes: “ Dotter-

stécke in zwei seitlichen Feldern resp. in randstindigen Lings-

treifen—in der Rindenschicht.” This phrase seems to imply an

agreement with the opinions just quoted, 7. e., that the strips of

vitelline follicles lie in the cortical layer outside the nerve-cord.

Nevertheless in a figure § (copied from Kraemer) it is distinctly

shown that in “ Tenia filicollis” (an Ichthyotenia) the vitelline

strips lie within, not only the nerve-cord, but the water-vessel.

In Solenotenia there 1s no doubt that the vitelline strips lie to

the znside of the nerve-cord though outside of the water-vascular

tube, and are thus obviously medullary in position. They also

certainly lie within the nerve-cord in other Ichthyotzniids which

I have myself || had the opportunity of examining.

* Rey. Suisse Zool. iv. 1897, p. 210.

+ “Die Ichthyotenien der Reptilien,” Inaug.-Diss. Univ. Basel, 1908, p. 22.

+ Bronn’s ‘ Thierreichs,’ Bd. iv. Abth. 1 B, p. 1699.

§ Loe. cit. Taf. lv. fig. 5.

|| P. ZS. 1913, p. 4.

Proc. Zoo. Soc.—1913, No. XVII. ihe

250 Dk. F. E. BEDDARD ON

The groove upon the ventral surface of the mature proglottids,

referred to above, cannot, as I think, be compared with the

ventral and dorsal grooves upon the proglottids of Piychobothrium

Text-fig. 49.

Transverse section through middle region of proglottid of Solenotenia viperis,

to show the ventral uterine groove.

d. Diverticulum of uterus containing an embryo (e). The opening into the uterus of

the corresponding diverticulum is not shown on the opposite side. se. Sub-

cuticular layer. J. Testis. U. Uterus. 2. Vagina.

REPTILIAN TAPEWORMS. 251

belones *. For in that Bothriocephalid the uterine pore or genital

pore opens into one or other of the grooves. In Solenotewiia, on

the other hand, the groove is actually the interior of the median

stem of the uterus, which is exposed to the exterior by a splitting

of the integument. The origin of this state of affairs is a question

to which I shall recur later. The extent and depth of the groove

is indicated in text-fig. 48, when seen from below in a proglottid

mounted entire, aml in fend fig. 49 in transverse section. On

the superticial view the actual groove is seen to occupy as nearly

as possible the median ventral line of the proglottid, and to be not

broader than 1/7th to 1/10th of the diameter of the proglottid.

The variation in breadth is of course due to the amount of gape

of the slit, which may be quite coextensive with the breadth of the

median stem of the uterus.

In transverse sections (see text-fig. 49) the gape of the ventral

groove is often seen to be fully as wide as the lumen of the

uterus. The diverticula open into this canal on either side. On

the whole they form two more or less regular rows, one on each

side (right and left) of the median groove. I have, however, seen

them more crowded, two diverticula opening close together on one

side and one on the other. There is thus no perfect regularity,

as is indeed generally the case with the uterine diverticula in

the Ichthyoteeniids.

It is very noteworthy that the continuous slit later nowhere

shows signs of tearing or bursting; its edges are smooth and

regular. This, of course, might be explained by the pre-existence

in earlier stages of the development of the uterus of a series of

uterine pores, increasing slightly in longitudinal diameter, and

thus leading eventually to a smooth and easy tear, such as is

effected in sheets of postage stamps. To this matter I shall

again draw attention in considering Immature stages of the

uterus. I have examined carefully the end of the ventral

groove, close to the ovary, where the uterus of course

commences.

The conditions met with at this end of the uterus are very

suggestive of a pre-existing pore. There is, in fact, as shown in

the accompanying drawing (text-fig. 50, p. 252), a distinct and

funnel-like in-pushing of the pubeuticuler layer of cells (and over-

lying cuticle), which opens into the cavity of the median stem of

the uterus. The subcuticular cells in the funnel-lke in-pushing

come to lie at right angles, or nearly so, to the subcuticular cells

upon the general surface of the body. The narrow aperture of

communication thus formed, widens out in subsequent sections

to the wide gape already described. It must be admitted that

this funnel-like involution (as it apparently is) is not altogether

like the uterine pores which I have recently described and figured

* Lonnberg, “Anatomische Studien tiber skandinavische Cestoden,’ K. Svensk.

Vet.-Ak. Handl. Bd. xxiv. (1891) No. 6; for a figure, see Bronn’s Thierreichs, Bd. iv.

Abth. 1B, Taf. xlvi. fig. 4.

Was

252, DR. F. E. BEDDARD ON

in Ophidotenia*, where there is not a very marked funnel-shaped

ingrowth of the subcuticular layer to meet the cellular tube which

grows out from the uterus. It will be noted, however, that there

Text-fig. 50.

Transverse section through middle of a proglottid of Solenotenia viperis at

posterior end of uterine groove.

d. Diverticula of uterus containing embryos. P. External pore which forms the

commencement of the ventral uterine groove. U. Uterus. V. Vagina, which

is seen cut through twice, being coiled in this region near to the ovary.

is some comparison possible between the conditions that I have

just described and those asserted to exist by Zschokke in certain

allied genera of Cestodes.

* P, Z. 8. 1913, p. 30, text-fig. 9.

REPTILIAN TAPEWORMS. 253

A salient character of the uterus of this genus Solenotenia is

afforded by the diverticula. This character is shared by Ophido-

tenia, and the diverticula appear to me to be quite similar in

both of these genera of Ichthyoteniide. The peculiarity of

these diverticula is, first of all, their clear separation from the

uterine stem into which they open, often by a very narrow mouth,

and, secondly, the delicate long-stalked glandular cells which beset

them externally (text-fig. 51). As I have already pointed out in

the case of Ophidotenia. these cells seem to be exactly like those

of the shell-gland in other tapeworms, and I can here say the

same of those of Solenotenia. It appeared te me to be right in

the case of the allied form Ophidotenia naiw* to correlate the

Text-fig. 51.

A transverse section of one of the uterine diverticula of Solenotenia viperis highly

magnified to show the long-stalked glandular cells which form the greater

part of its walls.

presence of these cells, distinguishing the diverticula of the

uterus of this genus from those of [chthyotenia, with the absence

of a well-marked shell-gland, which is very conspicuous in many

species of Ichthyotenia. I find, however, in Solenotenia viperis

a short thick-walled region of the oviducal apparatus, which I

take to be the equivalent of the shell-gland. This is far less

conspicuous than the shell-gland of Jehthyotenia, where I have

seen it; and the cells are not stalked, and have thus not the

characteristic appearance so well known in Cestodes,

In less fully mature proglottids, where there is no ventral

groove upon the ventral face of the body, the uterus is a simple

* P_Z.S. 1913, p. 33.

254 DR. F. E. BEDDARD ON

tube of approximately circular contour in transverse section. At

its termination its walls are quite thin, and consist of little but a

membrane. But elsewhere, although there are no diverticula as

in the ripe uterus, the walls are laterally marked by a thick layer

of cells (text-fig. 52), which are the forerunners of the pear-shaped

cells which deck the diverticula of the mature uterus ; so, at least,

I presume. ‘These cells form a heap on each side, and are not

markedly pear-shaped. Rather, indeed, are they circular in

outline. .In many sections this uterine tube is seen to he at

some distance from the ventral body-wall, and a certain thick-

ness of cortical parenchyma lies between it and the glandular

subcuticular layer. In other regions of the proglottid and for a

considerable number of consecutive sections, the shape of the

uterus in transverse section becomes pear-shaped, the tube thus

extending right up to the subcuticular layer, which is, however,

as far as I could ascertain, not perforated anywhere.

The cavity of the uterus is here not large, and it is not swollen

with eggs, of which, indeed, but few are pe be observed at this

stage in its cavity, and those not mature

I have recently described in inde gabonica t prolonga-

tions of the uterus towards the exterior, the histological details

of which offer reasons for believing that they are rudimentary

(perhaps incipient) external uterine pores like those so plain in

Ophidotenia. In the former species, however, there was not

always so marked an extension of the cavity of the uterus

as of the tissues forming its walls. In Solenotenia, on the

other hand, what is generally met with is rather an extension

of the uterine cavity which often came to underlie considerably

the subcuticular layer. A weak line is thus formed, which by

splitting forms the ventral groove so characteristic of the mature

proglottids of this Ichthyoteniid. Occasionally, however, the

approach of the prolongation of the uterus to the cuticle, is

associated with the disappearance of the subcuticular layer for a

brief space. Such spots seem to me to represent the uterine

pores of Ophidotenia. The existence of the remains of an

external pore is, I think, well shown in text-fig. 52. This

illustrates two out of about six sections through the uterus of a

not fully mature proglottid close to it, beginning at the ovarian

end of the proglottid. The uterus consists in each of these few

sections of a circular area suspended by a thin stalk from the

subcuticular layer. The stalk is not hollow but composed of a

nucleated layer of cells, continuous with the walls of the uterus,

and very distinctly not formed as a specialised tract of the

medullary parenchyma. In one of the two sections selected for

figuring here it appears to me that this duct, as I regard it, could

be followed as far as to the cuticle. I could not, however, see

* T am, indeed, not quite certaim that they are eggs.

+ P. Z.S. 1913, p. 157, text-fig. 34.

REPTILIAN TAPEWORMS. 255

anywhere a lumen in this cord of cells. That fact, nevertheless,

ean hardly be regarded as militating against the view that I take

of their nature. Following this series of sections to which I have

just referred onwards the cord of cells ceases absolutely, and the

uterus is to be seen as a circular cavity lying some way away

from the subeuticular layer. Very soon, however, it grows out

towards the cuticle with a lumen throughout which reaches the

subcuticular layer. There is no obvious suggestion here of a

duct. In fact, the condition of the ripe uterus in Solenotenia is

not altogether easy of comparison in detail with that of Ophido-

tenia.

Text-fig. 52.

Two nearly consecutive sections through unripe uterus of Solenotenia viperis

showing (in the left-hand figure) a rudimentary external pore. The lateral,

thickenings of the uterine tube are probably the commencement of the lateral

diverticula.

The above facts in the structure of the uterus of Solenotenia

show clearly that it is to be referred to the same type as that

which characterises the genus Ophidotenia, differing therefore

from that of Ichthyotcenia sensu stricto, though one species of

Tchthyotenia, I. gabonica, shows certain points of likeness to both.

It is possible, however, that in a revision of the family Ichthyo-

teenlidee, Ichthyotenia gabonica may ultimately be given generic

rank. The resemblance, however, is somewhat masked by the

artificial opening of the uterus, presumably caused by splitting

along a ventral line where the uterus, then swollen with eggs,

approaches close to the integument. The slit thus formed is not

exactly a concrescence of numerous uterine pores, of which, how-

ever, there are plenty of traces in Solenotenia; it is largely

independent of them—perhaps altogether so. On the other hand,

256 DR. F. E. BEDDARD ON

they may assist in forming the ventral groove or slit. Of this I

have not positive evidence one way or the other. This condition

is, as I think, to be directly compared with the cases described by

Zschokke, whose statements as to their significance have been

denied, as I have already pointed out, by others. Details are

wanting as to whether Zschokke’s types show evidence of the

pre- existence of uterine pores in younger stages.

The question of the absence or presence of a separate uterine pore

in the Tetraphyllidea is not uniformly answered in the text-books,

Thus Sedgwick, in his ‘Student’s Text-book of Zoology,’ * remarks

that “in the Bothriocephalide and other forms (possibly in all

Cestodes except the Treniadee) the uterus has a special opening

of its own on to the exterior.” On the other hand, Benham

divides the Dibothridiata (containing only the Bothriocephalide

and forms with grooves instead of proper suckers) from the

Tetrabothridiata (the rest of the Merozoic Cestoidea) by the

occurrence of the uterine pore in the former and its absence in

the latter t. Probably the balance of opinion is in favour of the

view held by Dr. Benham. Leuckart {, however (at any rate in

the revised English edition of his work), was apparently of opinion

that uterine pores are found in the whole group of the Bothriade,

by which group he understands “the species which remain after

the exclusion of the Treniade,” though, as Prof. Braun has

pointed out in his account of the Tapeworms in Bronn’s

‘Thierreichs,’ Leuckart admitted that but few species had been

examined from this point of view.

Prof. Zschokke, in his most extensive work upon the Cestoidea §,

has brought forward certain facts which he regards as pointing to.

a retention of the Bothr iocephalid uterine pores among the higher

Tapeworms. This orifice in Calliobothriwm coronatum i is descr “ibed

as “un enfoncement en entonnoir de Ja cuticle, auquel correspond

un relévement interne de la paroi de Vutérus.” He observes,

further, that there is no sign of tearing, and a perfect continuity

in the walls of this tube. The orifice is figured 4] without

histological detail. On pl. v. fig. 81 of the same memoir is

represented the corresponding pore of another species, C’. leuckarti,

also without histological detail, and of it a similar description is

given. In another! species algo a uterine aperture is said to be

formed when the contained ova are ripe. Of Onchobothrium

uncinatum thesame fact is stated**. In the genus Anthobothrium tT

a similar uterine pore is stated to exist in the ripe uterus; but

* Vol. i., London, 1898, p. 249.

+ A Treatise on Zoolog ey, edit. by E. Ray Lankester, pt. iv. London, 1901, pp. 116

& 118.

+ ‘The Parasites of Man,’ Edinburgh, 1886, transl. by W. E. Hoyle, p. 317.

§ Mém. Soc. Inst. Geney. xvii., 1889.

|| Loe. cit. p. 196.

* Loe. cit. pl. iv. fig. 71.

** Toc. cit. p.-250.

+7 Loe. cit. pp. 260, 272, 281.

REPTILIAN TAPEWORMS. 257

no details or figures are given. Dr. Zschokke has not given

any figures of transverse sections through these pores, which

are represented in his drawings of them in full face as rather

slit-like, and thus suggestive of a tear due to the pressure of eggs.

There has been some criticism of the conclusions which I have

just briefly abstracted.

Pintner, criticising* these statements as embodied in an

earlier paper f of a preliminary nature, definitely denies their

truth. “ Fiir simmtliche mir bekannte Arten,” he writes, ‘‘ der

Gattung Calliobothrium, fiir Anthobothrium musteli, fir Phyllo-

bothrium gracile, fir Echenetbothriwm und noch fiir manche andere

von mir untersuchte Tetrabothrien, trifft das ganz entschieden

nicht ¢ zu.” Pintner goes on to observe (and his remarks are

given by Braun) that a definite slit is formed on the ventral

surface of the proglottid shaped‘ like a “ gothic door,” out of

which the eggs pour. This is an artefact due to the internal

pressure of the swollen uterus, which lies nearer to the ventral

than to the dorsal surface. This slit can be recognised in carefully

handled proglottids as a median line. Doubtless, this latter

may be a vestige of the uterine opening of the Pseudophyllidea ;

but that there is no actual natural pore Pintner is satisfied.

The existence of uterine pores has also been dealt with by

Kraemer §, and in forms more nearly allied to Ophidotenia and

Solenotenia, viz. in Ichthyotenia itself. For to this genus (at any

vate sensu lato) we are, I believe, to refer Tenia filicollis,

7’. ocellata, and 7. torulosa, the anatomy of which is described by

_Kyvaemer. Of the first-named species the author remarks that

the eggs are divided by a break in the body-wall, that is to say,

through a secondarily appearing uterine opening, which is

illustrated by a figure || very like those given by Zschokke and

referred to above. A more elaborate account is given of “ Tenia”

torulosa. But the secondarily formed slit appears to be formed

in the same way as that of the other species, and no suggestion is

made that it is in any way a permanent structure, or even a last

vestige of the real uterine pore of the Bothriocephalids. Kraemer

investigated the slit-like pore by means of sections, and traced it

into continuity with the cavity of the uterus, observing (if I

understand him rightly) that there is no difference in form which

would imply a differentiated duct.

Prof. Braun does not accept the views of Zschokke with

reference to a uterine pore 4 in Calliobothriwm, ete., and in the

* “Neue Untersuchungen tiber den Bau des Bandwurmkorpers,” Arb. Zool. Inst.

Wien, viii. 1889, p. 6 (of memoir) footnote.

y+ “Studien uber den anatomischen und histologischen Bau der Cestoden,”

Centralbl. Bakt. u. Paras. Bd. i.

{ Italics, author’s.

§ “ Beitrage zur Anatomie und Histologic der Cestoden der Stisswasserfische,”

Zeitschr. wiss. Zool. liii. 1892, p. 647.

|| Loe. cit. Taf. xxviii. fig. 42.

4 Bronu’s Klassen und Ordnungen des Thierreichs, Bd, iv. Abth. 1 B, p. 1441.

258 DR. F. E. BEDDARD ON

systematic list of families and genera distinguishes the Tetra-

phyllidea from the Pseudophyllidea by the character (among

others) ‘‘ Keine praeformierte Uterusmiindung” *; and again,

with reference to the Cyclophyllidea f uses as part of his diagnosis

the words “ Keine Uteruséffnung.” Nothing can be more definite

than these statements, which appear to express the general

opinion upon the matter. In detailed criticism Braun points out

that the alleged uterine pore differs in its position in various

proglottids, which is not consonant with the idea of a fixed and

definite structure. He points out, further, that the constancy of

what he considers to be merely a point of rupture to the ventral

surface is associated simply with the ventral situation of the

uterus, and perhaps, furthermore, reminiscent of the ancestral] and

ventrally situate uterine pore. Prof. Max Braun, in fact, entirely

associates himself with Pintner’s criticisms of Prof. Zschokke.

The statements of so expert a student of Cestodes as Prof.

Zschokke must not be passed over without attentive examination.

It is to be noted, however, in the first place, that the conditions

which he describes and figures in Calliobothriwm (and allied

forms) are different from those which I find in Ophidotenia and

Solenotenia, and that these observations do not therefore precisely

confirm those of Zschokke, though they certainly increase their

probability on general grounds. It may be that Zschokke’s two

figures refer to a stage comparable to that which I have figured

in a comparatively young proglottid of Ophidotenia russelut. I

am not, however, disposed to think that this is the case, for the

same state of affairs exactly appeared to occur in all of the many

species enumerated by Zschokke. There would not be this exact

repetition were the circumstances as I have described them in

Ophidotenia, and in the genus which forms the subject of the

present paper. Iam, therefore, of opinion that a further study

of Calliobothrium, Anthobothrium, etc., is necessary before they can

be accurately compared with Ophidotenia and Solenotenia.

At the time when I was occupied with the study of the species

of Ichthyotzeniids, the results of which have been communicated

to the Society, the volume of the ‘ Zoological Record’ published in

1912 had not come into my hands. I regret, therefore, to have

missed a preliminary notice § by Mr. G. La Rue of a forthcoming

revision of the Ichthyoteeniide (or, as he terms the family, Proteo-

cephalidze), in which an important fact is recorded which I had

thought to have put forward for the first time. In the definition

of the family Proteocephalide, the following sentence occurs :—

“ Uterus median, ventral, with lateral outpocketings, and with

one or more preformed ventral openings,” and in the description of

five new species of Proteocephalus (all from fishes), the number of

Bronn’s ‘ Thierreichs,’ Zoe. cit. p. 1698.

Loe. cit, p. 1707.

P. Z.S. 1918, text-fig. 35, p. 163.

Zool. Anz, Bd. xxxyii. p. 473.

REPTILIAN TAPEWORMS. 259

these ventral openings is mentioned; thus, e.g., he writes of

Proteocephalus singularis, sp. n.: ‘ Uterus with 20-25 lateral

pouches on either side and with 2—4 ventral pores.” The same

statement (save for the actual number of the pores) is made of one

species of Ophiotcenia *, a new genus described by the author.

This comprises all that is written in the preliminary note

concerning the uterine pores. There is enough, however, as I

believe, to prove that Mr. La Rue and myself have observed

identical structures; but it is, on the other hand, conceivable that

the conditions characteristic of the various species described by

La Rue are similar to those which I have described above in

Solenotenia, and that the ‘ventral openings” of the uterus may

be merely the commencement of ruptures which end in fusing

together and forming a ventral groove. This view would account

for the variable numper of the openings in a given species, of

which I have quoted an instance above.

The testes of this species are very numerous, and really extend

quite across the body except for the middle regon occupied by

the uterus, and this is relatively quite narrow. Posteriorly the

testes reach the ovary, and are found at the side of that gonad

which is not greatly extended laterally. The most posterior

region of the proglottid is, however, free of testes ; not however

the most anterior, for they extend quite to the anterior limit of

the proglottid. In younger proglottids, where the uterus is

simply represented by a narrow median cord, the testes are more

obviously disposed in two lateral bands, leaving a considerable

gap in the region of the uterus, which is to be occupied by the

latter when fully mature and provided with its diverticula.

In mature proglottids I have seen quite ten testes on either

side of the uterus; but they are not arranged in regular rows.

The coil of the vas deferens before it enters the cirrus-sac is

denser than I have seen it in some allied forms. The coil

extends quite to the uterus, and the lumen of the tube is wide.

The cirrus-sac (text-fig. 53) seems hardly to possess a definite wall

of itsown. The wall of the cavity in which the coiled cirrus lies is

practically a layer of the network of the medullary parenchyma.

This region of the cirrus-sac is hardly wider than that situated

nearer to the external pore. Further in the latter direction the

bands of longitudinal muscle-fibres come to lie actually on the wall

of the cirrus where, however, they are obviously merely adven-

titious and undergo no modification, such as increase in thickness,

to assist in the protrusion of the cirrus. The cirrus-sac is not so

markedly flask-shaped as it isin many other Cestodes; but there

is a narrower neck region towards the external pore. This region

has marked but not very thick muscular walls, in which the

fibres trace a more or less circular direction, thus forming a

* T do not know whether this genus is exactly coextensive with my genus Ophido-

tenia. The preliminary definition given of Ophiotenia is not sufficient. The

accidental similarity of names is a coincidence which might well have been expected.

260 DR. F. E. BEDDARD ON

constrictor ; but I am not able to assert whether this is or is not

a local modification of the general longitudinal layer of the body.

The interior of the cirrus-sac is packed with a delicate fibrous

Text-fig. 53.

Transverse section through a portion of a proglottid of Solenotenia viperis, showing

the cirrus-sac, ¢.s., which is seen to have no definite walls. In the interior

of the sac is shown the terminal wider part of the cirrus and three sections of

the narrower part of that tube. sp.d. Coiled region of sperm-duct.

network, with elliptical or spherical interspaces, which seems to me

to present no differences from the general parenchyma of the

body. The cirrus increases in diameter towards the external

REPTILIAN TAPEWORMS. 261

pore, and this thicker proximal region lies straight within the

sac. The cirrus is invested with a layer of glandular cells. This

layer is continuous with the subcuticular layer of the body. The

male orifice, aS is general among the Ichthyoteeniids, is some-

times in front of, and sometimes behind the vaginal pore. There

is nothing remarkable about the ovary or the vagina.

§ Résumé and General Observations.

The foregoing account of the worm Solenotenia viperis proves

the existence in a New World snake of an Ichthyoteniid, most

nearly allied to my genus Ophidotenia, which is Old World in

range, and at present known only from India. This distributional

fact gives at once a certain @ priori reason for justifying the

distinctness of the two genera. I do not, however, attempt to

justify my conclusion by the geographical fact only. The large

size of the scolex, the pecular apical sucker, and the ventral

groove upon the mature segments seem to me to be sufficient

facts to go upon in the present state of our knowledge of the

Ichthyotzniide. I cannot, however, state the absence of this

apical sucker in Ophidotenia. The most valid reason is the

gaping of the uterine slit to form a ventral groove upon the

mature segments. In this respect Solenotenia has advanced a

stage further than Ophidotenia. For while there are traces of

separate and pre-existing uterine pores, these are replaced by an

opening up (however caused) of a ventral median area of the

uterus and integument. This leads the way to—if it be not

identical with—the conditions recorded by Zschokke and others in

Calliohothrium and other genera. It appears to me that inas-

much as my specimens of Ophidotenia were furnished with fully

mature proglottids (as regards ova and uteri), a ventral slit is not

developed in that genus, and is therefore a legitimate character

in the definition of Solenotenia.

The existence of this ventral uterine slit still further justifies

the relationship between the Ichthyotzniids and Calliobothrium,

and other ‘‘ Bothriadz,” and their removal from the Tetracotylea,

with which they have been placed, though sometimes only doubt-

fully. It seems probable that their definite inclusion by Max

Braun (in Bronn’s ‘ Thierreichs’) with the Tetraphyllidea is

correct.

262 DR. W. A. CUNNINGTON ON

21. Zoological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, 1904-1905.

—Report on the Branchiura. By Witiiam A.

Cunnineton, M.A., Ph.D., F.Z.8. ;

[Received February 11, 1913: Read March 4, 1913.]

(Plates XLI.-XLYV.*)

INDEX.

Systematic: Page

Argulus: Key to African Species of .................. 267

AR GULUS DECISIS, SDs Meee eerie eee REA ee eee LOS

Al GVARTO IIS RYDs 18: Sade ats adeeineoed soos spvseneasee: 6, eSATA!

Aisemiguiis: Spe mie) ies, ee ee cee a na 272

A NONGUSTLCEDS, Spin este eee en TO

PASENLALUSS (SPs. Deseo, if socae te ceeeemeeMasteta secs Meee eT

APUDESCENS:SP cle pe cse cathe one eee a ee LO.

ID RS ERO UOD, URNS: CIE ccoe coe odo can ccoenoan {ke vanandenenasess RO

1. Introduction.

Through the collections made by Stuhlmann in Lakes Victoria

Nyanza and Albert Edward Nyanza, and by Filleborn in

Lake Nyasa, we have for some time known of the existence of

Argulide in the waters of those lakes. The material collected

during the Third Tanganyika Expedition proves that these

parasitic copepods also occur commonly in Tanganyika, from

which lake, indeed, far more species are now known than

from any of the other great African lakes. As many as

363 specimens were obtained by this Expedition, the great

majority of them from Tanganyika, with a few from Victoria

Nyanza. There are nine species represented in all, and it is

interesting to remark that, while two species—both previously

described—were found in Victoria Nyanza, the same two forms

were associated with no fewer than seven new species in Tan-

ganyika.

Since the return of the Expedition there have come into my

hands specimens of Argulids from Nyasa and from Albert

Nyanza; and as these constitute new records and add to our

knowledge of distribution, it seems desirable to refer to them

also in the present paper. Including the two types collected

by Fulleborn in Nyasa, the following is thus a complete list

* For explanation of Plates, see pp. 282-283.

I Zos> WOH, JPL SILI.

W.A.C. Photo. London Stereoscopic Co. imp.

1, 2. ARGG6EES.INCISUS. 3°5. A. RUBROPUNCTATUS.

Osea AL ae RSONAT Us:

Ps; WIZ, IP, SGI.

W.A.C. Photo. London Stereoscopic Co. imp.

OP ARGUES ExIGUIS: HO), Wis” No VAGUS TIGIEIHS:

12-14. A. STRIATUS. TOMO. VA IK BIE SENS:

1) OSS IE LN

W.A.G. del.

Huth sc.et imp

1718. ARGULUS INCISUS. 19,20.ARUBROPUNCTATUS.

21,22.A.EXIGUUS.

I ZS LS), PIII,

W.A.C. del. Huth se.et imp.

28-20 /velG UILAU'S igi SONAIUS, 26,4 2 AJANNIGIUS TUGIaIE Ss),

FAS) OMS JEN CLI

SD A

ATARI GRAN A ARS

W.A.C. del. Huth sc. et imp.

ee SO ARGUuLUS STEIATU Ss, Si so VAC ISSUE) S| CiEINS

BRANCHIURA FROM LAKE TANGANYIKA. 263

of the species at present known to occur in the lakes in

question :—

TANGANYIKA.

Dotlops ranarum Stuhlmann.

Argulus incisus, sp. n.

» rubropunctatus, sp. n.

AS personatus, sp. Ni.

5 + exGUes, Sp. N.

He angusticeps, sp. 1.

op striatus, sp. n.

» rubescens, sp. n.

» africanus Thiele.

Victoria NYANZA.

Dolops ranarum Stuhlmann.

Argulus africanus Thiele.

Nyasa.

Dolops ranarum Stuhlmann.

Argulus africanus Thiele.

Chonopeltis inermis Thiele.

ALBERT NYANZA.

Argulus africanus Thiele.

By the kindness of Dr. Calman I have been permitted to

examine the African specimens belonging to the British Museum.

They are of species already known to science, but as, in one

instance, they afford a record of a new locality, and as no notice

of them has been published, they will be referred to in their

place in the systematic part of this paper.

To illustrate the general appearance of the new species, I have

made use of photographs taken with a low-power lens from the

actual specimens themselves. The difficulties in the way of

obtaining reasonably good results from objects which are by

no means flat are very considerable, but I think that the photo-

graphs in question are really more satisfactory for the purpose

than drawings would be. They were made with a special photo-

micrographic lens in the Zoological Laboratory of King’s College,

London, and I have to express my indebtedness to Prof. Dendy

for allowing me to work in the laboratory, and also to him and

to Mr. R. W. H. Row for affording me the benefit of their

experience with the apparatus.

2. Systematic Notes and Description of New Species.

Family ARGULID&4.

DoLoPs RANARUM Stuhlmann.

Gyropeltis ranarum Stuhlmann, Zool. Jahyb., Syst. Bd. vi.

1891, p. 104.

264 DR. W. A. CUNNINGTON ON

Dolops ranarum Bouvier, Bull. Soc. Philom. sér. 9, tome 1.

ISG To las

Dolops ranarum Wilson, Proc. U.S. Nat. Mus. vol. xxv. 1903,

p. 737.

Dolops ranarum Thiele, Mitt. Zool. Mus. Berlin, Bd. ii.

Heft 4, 1904, p. 17.

This species was originally obtained by Stuhlmann at Bukoba,

on the western shores of Victoria Nyanza, and described by him

under the name (Gyropeltis ranarwm. The specimens were

found attached to the surface, and occasionally at the entrance

of the branchial chamber, of a number of tadpoles, and on that

account received their specific name. It does not appear a very

fortunate title, however, since specimens were obtained by the

present Expedition from the bodies of four different kinds of

fish, and most of the specimens, as a curious coincidence, actually

from Bukoba itself. It is, indeed, no new fact that certain

species of Argulidz may be found on a variety of hosts, the

well-known European type, Argulus foliaceus, being an excellent

example, as its host-list also includes frog-tadpoles as well as

a number of fish. The literature of the subject contains other

cases of similarly inappropriate specific names derived from those

of the hosts, and as long ago as 1875 Claus* deprecated this

obvious but unsatisfactory manner of naming species.

It may be no more than coincidence, but it is remarkable how

cften this form was found associated with Argulus africanus

on a common host.

TANGANYIKA.—Dolops ranarum does not appear to be common

in Tanganyika, since it was obtained on only one occasion, and

then from a single large fish. The coloration of these specimens

differs somewhat from that stated hitherto, and, in fact, to some

extent from the Victoria Nyanza specimens in the present col-

lection. The upper surface of both carapace and abdomen is

irregularly covered by arborescent pigment-spots of a grey

colour, which are most numerous on the carapace lobes. In

addition, there is an irregular series of small white spots,

scattered without any apparent relation to the grey pigment-

spots already mentioned.

Occurrence.—Kala, 18/11/04. From mouth-cavity and gill-bars

of large Lates microlepis (length 910 mm.). Three male and

three female specimens. Associated with them were specimens

of Argulus africanus.

Vicrorta Nyanza.—This is the only species of Argulid hitherto

known from the lake, and still seems to be the commonest,

although specimens of another widely distributed form (drgulus

africanus) were obtained during this Expedition. The specimens

of D. ranarum taken in this lake differ slightly in colour from

those collected in Tanganyika. They are of a more uniform

brown colour, and show little of the arborescent grey spots above

* Zeitschy. f. wiss. Zool. Bd. xxv. 1875, p. 278.

BRANCHIURA FROM LAKE TANGANYIKA. 265

mentioned, though these are faintly indicated in some cases.

The white spots are clearly visible.

Occurrence.—Bukoba, 24/4/05. From surface of body, mouth,

and gill-chambers of a large Siluroid (? Bagrus degeni)—native

name, Vfui. Ten male and eleven female specimens. <A

specimen of Argulus africanus was associated with these.

Bukoba, 24/4/05. Associated with specimens of Argulus

africanus from surface of body and inside mouth of large Proto-

pterus cethiopicus (length 1040 mm.). Three males and one

female.

Bukoba, 24/4/05. From surface of body, mouth, and gill-

chamber of large Siluroid (¢ Clarias anguillaris)—native name,

Nshonzi. Two male and three female specimens. An indi-

vidual of Argulus africanus occurred with these.

A single female specimen in the collection of the British

Museum has been more recently obtained from the lake. The

particulars recorded are:—‘On skin of fish (‘ W/alé’) ? catfish.

Lake Victoria. Collected by G. D. H. Carpenter, 6. xi. 11.”

This was associated with specimens of Argulus africanus.

Nyasa.—I am indebted to my friend Capt. E. L. Rhoades, of

the British Naval Service on Lake Nyasa, for a small collection

of Argulide from this lake. The specimens all proved to belong

to this species, and not to either of those obtained by Fiilleborn

in the lake. They are thus an addition to the number of forms

known in Nyasa, and at the same time a record of still wider

distribution of the species. There are two male and four female

specimens, the only particulars given on the label being :—

“* Wish-lice, ex. ‘ Suwngwa’: Lake Nyasa, 1908.”

River Niie.—In the British Museum collection are six male

and ten female specimens from yet another new locality. With

them was associated a single female specimen which I place, with

some hesitation, under Argulus africanus. They were obtained

by Mr. L. Loat, who gives the following details :—-“* Parasites

out of the mouth of a large ‘ Hala’ (Heterobranchus bidorsalis),

caught at the mouth of Lake No, White Nile, Jan. 30, 1901.”

Genus ARGULUS.

All the remaining Argulids in the collection belong to the

genus Argulus itself. The number of species included in this

genus is fairly considerable, yet they are, on the whole, reason-

ably distinct, and there seems to be no grounds for subdividing

the genus. As the number grows, it becomes at the same time a

matter of increasing difficulty to diagnose a species accurately,

save at some length, since the differences are often of a minor

nature, concerning numerous parts of the external anatomy.

These facts render it more than ever difficult to offer to other

workers in this field a satisfactory key to the species.

Wilson *, in 1903, made a serious attempt to provide a key to

* Proc. U.S. Nat. Mus. vol. xxv. 1903, p. 701.

Proc. Zoot. Soc.—1913, No. X VIII. 18

266 DR. W. A. CUNNINGTON ON

the whole of the then known species of Argulus, but without

achieving a great success. He was unfortunate in choosing for

the purpose certain characters which clearly depend upon the

state of contraction of the specimen and, in addition, are difficult

to distinguish with exactness. It is not easy, for example, to

determine how far the swimming-legs are covered by the

carapace, as this is influenced by the relative states of con-

traction, as well as by any chance displacement. Again, while

it is easy in extreme cases to point out forms in which the

carapace lobes overlap, or do not reach the abdomen, it 1s

virtually impossible to distinguish accurately those with “ Cara-

pace lobes just reaching the base of the abdomen.” Most keys

are open to the objection that they are purely artificial, and

Wilson, it is true, fully admits the artificiality in this case.

Nevertheless, a key in which in so many cases male and female

specimens of the same species are separated under merely

arbitrary standards, falls very short of what is to be desired.

It is certainly a matter of great difficulty in this genus to

discover characters which satisfactorily lend themselves to the

purpose of key-making. Where male and female differ markedly

- In external shape, as is sometimes the case, it seems perhaps

inevitable that the sexes should be artificially separated; but

this might be avoided to a greater degree than has been done by

Wilson. While objecting to the form of the key above men-

tioned, we feel that Wilson is to be commended for attempting

to tabulate the species ; and since he included all the then-known

forms, it at first seemed wise to incorporate in his key the seven

new species described in this paper. It was on going into the

matter with this end in view that the unsatisfactory features

already mentioned revealed themselves, and, in addition, certain

regrettable inaccuracies * which only added to the difficulties of

incorporation. Under these circumstances it was finally decided

to prepare a key to include only the African species ; and this is

the more desirable, since the one species of this genus hitherto

known from the continent is now increased to eight.

Before proceeding to give the key and the descriptions of new

species, a few words of caution may be offered. The characters

employed in diagnosing the forms belonging to this group, while

perhaps the most obvious, are in some cases profoundly influenced

by artificial contractions and contortions, and in addition by

the age of the individual. Proportions of length to breadth

are seldom quite constant, size of suckers may vary with degree

of expansion, and certain spines and processes may be more

* Op. cit. p. 702. On this page, for example, under the heading 6., 4. niger is

said to have an “abdomen narrow, ovate.’ Under the heading 7., A. pugettensis is

contrasted with this as having ‘*‘ abdomen wide, elliptical.” Actual measurements

of both these species are given in the body of the work, and may be written breadth

over length as follows :—niger = pugettensis ee Expressing these in the form of

percentage of breadth to length, they become :—niger 81:2, pugettensis 75°4. Thus

it is evident that the abdomen of niger, described as “narrow, ’ is in reality broader

than that of pugettensis, described as “ wide.”

BRANCHIURA FROM LAKE TANGANYIKA, 267

prominent in some individuals than others. An extensive series

of measurements undertaken by the writer has shown that, while

sizes and proportions may be stated in general terms, a con-

siderable degree of variation is commonly met with, and that

young specimens as a rule deviate somewhat from the normal

specific characteristics.

Concerning the actual figures given in the specific descriptions,

it is to be noted that the proportions are not always precisely

those adopted by Wilson. The Jatter gives the relative length of

the abdomen as a fraction of the entire length of the animal

exclusive of the abdomen. This has seemed to me less satisfactory

than to express it as a fraction of the total length of the animal.

What is wanted is to indicate briefly what proportion of the total

length is constituted by the abdomen, and this mode of expression

has therefore been chosen in the following descriptions.

Again, in the matter of the suckers it seems likely that Wilson’s

fractions are not strictly comparable with those given in this

paper, although the same plan has been adopted of expressing the

size as a proportion of the breadth of the carapace. My earlier

sucker measurements were all made on a basis of the over-all

diameter contrasted with the carapace breadth. They gave

decimal fractions which seemed to vary somewhat erratically,

and it was, moreover, most difficult to decide what might be

legitimately considered as the edge of the sucker in an average

state of expansion. The well-marked chitinous ring to which

the membranous border of the sucker is attached seemed to offer

a much more definite structure for investigation, the dimensions

of which could be clearly observed. A further series of measure-

ments, in which the size of this chitin ring was taken as a

standard of comparison, gave results, when expressed in fractions,

which were distinctly more uniform; and this method has con-

sequently been adopted for arriving at the figures given in the

specific descriptions. The diameter of the suckers and the

corresponding decimal fractions are both smaller, of course, than

if the over-all dimensions had been taken; and as the figures

obtained are smaller, on the whole, than those given by Wilson,

it appears highly probable that his are based on the maximum

diameter of the suckers.

Key to the African Species of Argulus.

a. Carapace orbicular, about as long as broad.

6. Antero-lateral depressions very pronounced ; no accessory

spines or processes behind the antennules and maxil-

lipeds_........ .. incisus.

b’, Aneta: lateral depressions slight; ‘accessory spines pr resent

behind the antennules and maxillipeds ....................- rubropunctatus g.

’. Carapace elliptical, distinctly longer than broad.

en Basal segment of maxilliped bearing spines.

e. Second segment of antennule with anterior hook.

d. Abdomen long elliptical, lobes bluntly pointed and

separated by moderately broad sinus .................. personatus.

d’, Abdomen broadly elliptical, lobes obtusely pointed

and separated by broad sinus; carapace with char-

acteristic reddish spots.............::ccceeee ee rubropunctatus 2

18*

268 DR. W. A. CUNNINGTON ON

c'. Second segment of antennule with anterior spine.

d. Abdomen slightly longer than broad, sinus less

than one-quarter of its length; eyes very large ... exiguus 6.

d'. Abdomen. distinctly longer than broad, sinus one-

half of its length; eyes and suckers situated close

together and far forward . .. angusticeps ?.

. Basal segment of maxilliped bearing road | processes.

ce. Abdomen long elliptical, at least one and a quarter

times as long as broad.

d. Abdominal lobes pointed, sinus one-third or more

of length of abdomen ; ees lobes character-

istically striated . striatus.

. Abdominal lobes rounded, ‘sinus one-sixteenth of

length of abdomen.. .. rubescens &.

. Abdomen broadly elliptical, “less than one and a

quarter times as long as broad.

d. Basal segment of antennule bearing broadly ronnded

process, second segment with small posterior spini-

. Basal segment of “antennule _ bearing ‘somewhat

=punded = spine, second segment with stout pos-

terior spine; carapace with characteristic black

PUGWOCMUE-SIWOUS coos soganedaccos season qiecancenbeesoncocssn bas GYsMMG MUS:

Ancimus Incisus, spn. (Pl. XOUIS figs. 1572) Ply xen

figs: 17, 185)

Description.—Carapace orbicular, about as long as broad, with

antero-lateral depressions very pronounced, so that portions of

the antenne may be visible from above. Posterior lobes broad

and rounded, only covering bases of first and second swimming-

legs, and separated bya broad sinus about one-fifth the length of

the carapace. The edge of the carapace below is only moderately

armed with small spines, which extend to the region of the

swimming-legs. There are no such spines in the anterior region

in front of the eyes. Abdomen in the male twice as long as

broad, about two-fifths * entire length of body; posterior lobes

pointed, and separated by broad sinus two-fifths length of

abdomen. Abdomen in the female more than one and a half

times as long as broad, about one-third entire length of body ;

posterior lobes bluntly pointed and separated by broad sinus

one-half length of abdomen. The furcal appendages in both

sexes lie at the bottom of the anal sinus, Antennules and

antenne small and ill-armed, approaching closely to one another

in the mid line. The second segment of the antennule only

is armed with an inconspicuous anterior spine and slender lateral

hook. Noaccessory spines or processes close behind these appen-

dages. Eyes rather large, in the males less than one-quarter, in

the females less than one-fifth breadth of carapace apart, and

situated somewhat far forward. Suckers rather large, in the

males 14-15, in the females ‘157 of breadth of carapace.

Maxillipeds of moderate size, rather poorly armed; basal

segment produced into three flattened processes; ‘“ area ”

* Proportions not to be confused with those given by Wilson and differently

expressed, vide supra.

+ Diameter of sucker measured at chitinous supporting ring and not at extreme

edge.

BRANCHIURA FROM LAKE TANGANYIKA. 269

rather large, roughly pear-shaped and armed with small spines.

No accessory processes are present between the bases of the

maxillipeds. Flagella present on the first two pairs of swimming-

legs. In the male, vesicula seminalis present on the third leg,

copulatory peg and accessory cushion on the fourth. In the

female, basal segment of fourth leg produced postero-laterally

into pointed lobe. Colour (in spirit) light brown, the testes

covered above by large irregular blotches of dark brown.

Dimensions as follows :—

Adult male (largest specimen). Adult female (largest specimen).

mm. mm.

diotalaleme-bhiees sere 3 Notallilenethew ass .-ce. 11-6

Length of carapace ... 4:1 Length of carapace ... 6:2

Breadth of carapace... 4:3 Breadth of carapace... 6-7

Remarks.—The most striking characteristic of this species is

the unusually deep antero-lateral depression or sinus, which

gives occasion for the specific name. This feature does not

occur in any other of the African species—a fact which naturally

suggests that excessive shrinkage might be the sole cause. Such

an explanation is obviously impossible in this case. The spe-

cimens were all of them taken from a single fish, in association

with A. striatus, but the species were not distinguished on the

spot, and all the specimens were treated alike and were brought

home in the same bottle. On examination, the deep antero-

lateral incision first afforded a palpable distinction between the

forms, which proved to be specifically different to a marked

degree. Another important feature in which this type differs

from the other African species, and indeed from the great

majority of the known species, is the complete absence of

accessory spines or processes behind both the antennules and

maxillipeds.

Occurrence.—Rumonge, 16/2/05. From mouth and surface of

body of large specimen of Auchenoglanis occidentalis, var. tangant-

canus. Ten males and thirteen females, some ovigerous. These

were associated with a large number of specimens of Argulus

striatus.

ARGULUS RUBROPUNCTATUS, sp. n. (PI. XLI. figs. aM

Pi XGLIE. figs. 19,20:)

Description.—Cavapace in the male orbiculav, slightly tmgedlen

than long; in the female elliptical, slightly longer than broad.

Antero-lateral depressions slight. Posterior lobes broad and

rounded, covering bases of all but fourth swimming-legs, and

separated by a broad sinus about one-fifth the length of the

carapace. The edge of the carapace below is well armed with

small spines, which extend into the region of the swimming-legs.

Abdomen in the male nearly two and a half times as long as

broad, about three-sevenths entire length of body; posterior

270 DR. W. A. CUNNINGTON ON

lobes pointed and separated by fairly broad sinus one-half length

of abdomen. Abdomen in the female as long as or slightly

longer than broad, less than one-fourth entire length of body;

posterior lobes obtusely pointed and separated by broad sinus

almost one-half length of abdomen. The furcal appendages

in both sexes lie at the bottom of the anal sinus. Antennules of

moderate size, basal segment with stout posterior spine; second

segment with stout anterior hook and rather slender lateral

hook. Antenne with stout spine on basal segment. Immedi-

ately behind the insertion of the antennule is situated a stout

accessory spine. Eyes somewhat small, about one-quarter breadth

of carapace apart. Suckers in the males rather small, -12—14 of

breadth of carapace; in the females rather large, 15-16 of

breadth of carapace. Maxillipeds fairly strong and moderately

armed; basal segment produced into three stout spines; “‘ area”

of moderate size, oval shape, distinguished by red-brown pigment,

but only sparsely armed with spines. A pair of stout accessory

spines is situated between the bases of the maxillipeds, and

a second pair a short distance behind them. Flagella present

on the first two pairs of swimming-legs. In the male, vesicula

seminalis present on the third leg, copulatory peg and accessory

cushion on the fourth. In the female, basal segment of fourth

leg produced postero-laterally into sharply pointed lobe. Colour

(in spirit) light brownish grey, covered by an irregular series of

red-brown spots, which in part are aggregated together to form

definite blotches. Testes covered by irregular blotches of the

same pigment.

Dimensions as follows :—

Adult male (largest specimen). Adult female (largest specimen).

; mm. mm.

Motalslenethiess cece. 12-2 Motal Ventless, -e-cecee 10-6

Length of carapace... 7-1 Length of carapace ... 8-1

Breadth of carapace... 7:7 Breadth of carapace... 7:4

Remarks.—This species affords a good example of the striking

differences in form of male and female specimens which are not

uncommon, although extremely inconvenient for incorporation in

any key. Both the shape and dimensions of the carapace, as

well as the nature of the abdomen, are markedly dissimilar in

the two sexes. While the red spots and blotches are certainly

characteristic of the species, it must be admitted that they are

not equally obvious on all specimens. They show little on

certain specimens in the collection, which are uniformly of a

darker brown colour, and they are less conspicuous on young

individuals. The posterior lobes of the carapace, especially in

the female, may sometimes nearly meet in the middle line, thus,

of course, altering the shape of the sinus and also covering the

bases of the fourth swimming-legs.

Occurrence.—Kibwesi, 19/12/04. From surface of head (one

BRANCHIURA FROM LAKE TANGANYIKA. 271

specimen on eyeball) of very large Lates microlepis (length

1340 mm.). Thirteen male and five female specimens.

Maswa, 15/1/05. Mostly from the surface of the body of a

number of Lates microlepis. Fifteen males, seven females.

Ndanvie, 8/2/05. From surface of head and body of large

specimen of Lates microlepis. Seventeen males, three females.

As will be seen from the above, this form has always been

found infesting specimens of /ates, and never in association with

any other species of Argulid.

ARGULUS PERSONATUS, sp. n. (Pl. XLI. figs. 6-8; Pl. XLIV.

figs. 23-25.)

Description.—Carapace elliptical, longer than broad, with

rather pronounced antero-lateral depressions. Posterior lobes

broad and rounded, covering bases of all but fourth swimming-

legs: in a female distended with eggs the third swimming-legs

are only partially covered. The lobes are separated by a mode-

rately broad sinus, about one-fifth the length of the carapace or

sometimes more. ‘The edge of the carapace below is well armed

with very small spines, which extend to the region of the

swimming-legs. Abdomen in the male two and a half times as

long as broad, about two-fifths entire length of body ; posterior

lobes bluntly pointed and separated by moderately broad sinus

two-fifths length of abdomen. Abdomen in the female about

one and a quarter times as long as broad, more than one-quarter

entire length of body; posterior lobes bluntly pointed, and

separated by moderately broad sinus more than one-half length

of abdomen. The furcal appendages are small and inconspicuous

and lie at the bottom of the anal sinus. Antennules of moderate

size, basal segment with stout posterior spine; second segment

with small anterior hook, posterior spiniform process, and lateral

hook of medium size. Antenne with small spine on _ basal

segment. Immediately behind the insertion of the antennule is

situated a stout accessory spine. Hyes rather large, about one-

quarter breadth of carapace apart. Suckers of moderate size, in

the males -13—-15, in the females -10—"18 of breadth of carapace.

Maxillipeds strong and well armed; basal segment produced into

three stout spines; “area” large, pear-shaped, and armed with

small spines. A pair of stout accessory spines is situated

between the bases of the maxillipeds, and a second pair, rather

slighter, a short distance behind them. Flagella present on the

first two pairs of swimming-legs. In the male, vesicula seminalis

present on the third leg, copulatory peg and accessory cushion on

the fourth. In addition, a small backwardly directed process is

present near the base of the second leg, and a considerable conical

projection on the anterior face of the third leg. In the female,

basal segment of fourth leg produced postero-laterally into pointed

lobe. Colour (in spirit) light yellowish brown, the testes covered

by irregular blotches of darker brown.

72 DR. W. A. CUNNINGTON ON

Dimensions as follows :—

Adult male (largest specimen). Adult female (largest specimen).

mm. mm.

oval lemothieecrs meus 6°6 Motal Veniotnteeses-ca ee 11-4

Length of carapace ... 3:8 Length of carapace... 7°

Breadth of carapace... 3:4 Breadth of carapace... 6:0

Remarks.—While exhibiting no very outstanding character-

istics, this species is sufticiently well defined by a number of

minor differences affecting several parts of its external anatomy.

There are only a few specimens in the collection, and, as is usual,

the small ones differ to a certain degree from the large. In the

smaller specimens the antero- lateral depressions are less pro-

nounced, and the anterior hook on the antennules is less marked

or even absent.

In default of a name more obviously suggested by its anatomy,

the mask-like appearance of the carapace is alluded to in its

specific name.

Occurrence.—Ndanvie, 7/2/05. From mouth-cavity of several

specimens of Bathybates ferox. ‘Three males, two females.

ARGULUS EXIGUUS, sp.m. (Pl. XOLID digs 9);. Pl_ exes

figs. 21, 22.)

Description.—Carapace (in the male) elliptical, considerably

longer than broad, with slight antero - lateral depressions.

Posterior lobes moderately broad, rounded, covering bases of all

swimming-legs, and separated by moderately broad sinus one-

third the length of the carapace. The edge of the carapace

below is armed with small spines which extend to the region of

the swimming-legs. Abdomen in the male slightly longer than

broad, about one-quarter entire length of body; posterior lobes

bluntly pointed and separated by moderately broad sinus almost

one-quarter length of abdomen. The furcal appendages lie at the

bottom of the anal sinus. Antennulesand antenne rather small,

but moderately armed, approaching somewhat to one another in

the mid line. Basal segment of the antennule with posterior

spine; second segment with blunt anterior spine, posterior

spiniform process, and slender lateral hook. Antenne with slight

spiniform projection on basal segment. Immediately behind the

insertion of the antennule is situated a stout accessory spine.

Eyes very large, a little more than one-quarter breadth of

carapace apart and somewhat far forward. Suckers of moderate

size, "13-15 of breadth of carapace. Maxillipeds strong and

well armed; basal segment produced into three stout spines;

“area” of fair size, roughly oval, and armed with small spines.

A pair of stout accessory spines is situated between the bases of

the maxillipeds, and a second pair, rather slighter, a short

distance behind them. Flagella present on the first two pairs of

swimming-legs. In the male, vesicula seminalis present on the

third leg, copulatory peg and accessory cushion on the fourth.

BRANCHIURA FROM LAKE TANGANYIKA. Ze

Colour (in spirit) a reddish brown; in one specimen, the testes

are covered by irregular blotches of a darker colour.

Dimensions as follows :—

Adult male (largest specimen).

HRotallslemert hesausscreeeeectase eel 2-7

Weng thvor (carajace)-a eee t-es cl 2°1

Breadth of carapace ............ 16

Remarks.—This species is unfortunately only represented by

two specimens, both of the male sex. hey appear to be adult,

however, although they are small, and there seems no reasonable

doubt but that they represent a distinct species. The compound

eyes are unusually large, and are actually larger in proportion than

in the species which has beennamed A. megalops. The swimming-

legs in these specimens are so backwardly directed, that the bases

of at least the last pair are, strictly speaking, covered by the

abdomen and not by the carapace lobes.

Occurrence.—Mpala, 31/12/04. From surface of body of small

Simochromis diagramma. One male specimen.

Mpala, 31/12/04. From outside body of medium-sized Haplo-

chilus tanganicanus. One male.

ARGULUS ANGUSTICEPS, sp.n. (PI). XLII. figs. 10,11; Pl. XLIV.

figs. 26, 27.)

Description.—Carapace (in the female) elliptical, considerably

longer than broad, with antero-lateral depressions well marked.

Posterior lobes moderately broad, rounded, covering bases of all

but fourth swimming-legs, and separated by a somewhat narrow

sinus, more than one-third the length of the carapace. The edge

of the carapace below is well armed with very smali spines which

barely extend to the region of the swimming-legs. Abdomen in

the female slightly longer than broad, about one-quarter entire

length of body; posterior lobes somewhat pointed but with

rounded tips, and separated by moderately broad sinus one-half

length of abdomen. The furcal appendages, which are very small

and inconspicuous, lie at the bottom of the anal sinus. Antennules

and antennz small, but moderately armed, approaching closely to

one another in the mid line. Basal segment of the antennule

with posterior spine; second segment with stout anterior spine,

posterior blunt spiniform process, and slender lateral hook.

Antenne with slight spiniform projection on basal segment.

Immediately behind the insertion of the antennule is situated

a stout accessory spine. Eyes large, one-sixth breadth of carapace

apart, and far forward. Suckers small, -11 of breadth of carapace.

Maxillipeds strong and well armed; basal segment produced

into three stout and sharp spines; ‘‘area” of fair size, roughly

oval, and armed with small spines. Both suckers and maxillipeds

are situated unusually far forward. A pair of stout accessory

spines is situated between the bases of the maxillipeds, and a

274 DR. W. A. CUNNINGTON ON

second pair a short distance behind them. Flagella present on

the first two pairs of swimming-legs. In the female, basal

segment of fourth leg produced posteriorly into pointed lobe.

Colour (in spirit) a uniform brown, but somewhat darker in the

region of the egg-masses.

Dimensions as follows :—

Adult female (only specimen).

Motalllemoti es yes suse cee) set NOR

length of carapace ............ 78

Breadth of carapace ............ 56

Remarks.—-It is unfortunate that this species has to be

described from a single specimen. The latter is, however, an

ovigerous female which exhibits a number of distinctive features

and cannot be classed with any of the known forms. Prominent

characteristics are the narrow head region, rather well marked

off, and the large eyes placed close together. The eyes, suckers,

and maxillipeds are also situated remarkably far forward. The

carapace lobes appear curiously thick and fleshy—an appearance

which might possibly be due to shrinkage. The specimen was

obtained from an uncommon source, namely, from a basket of

small fish, but was attached to none of the latter. Thus, although

the specimen may well have been dead, there is no reason for

supposing that it was dried up to any extent, and indeed, its

general condition seems to preclude such a suggestion.

Occurrence.—Uvira, 5/3/05. Found among a number of small

fish of different kinds—at the moment adhering to none. A

single female, distended with eggs.

ARGULUS STRIATUS, sp.n. (PI. XLII. figs. 12-14; Pl. XLV.

figs. 28-30.)

Description.—Carapace elliptical, longer than broad, with

slight antero-lateral depressions. Posterior lobes fairly broad ~

and rounded, covering bases of all but fourth swimming-legs ;

in a female distended with eggs, the third swimming-legs are

only partially covered. The lobes are separated by a broad sinus

one-fifth the length of the carapace. ‘The edge of the carapace

below is armed with small spines, which extend to the region of

the swimming-legs. Abdomen in the male nearly twice as long

as broad, about two-fifths entire length of body ; posterior lobes

pointed, and separated by broad sinus one-third length of abdomen.

Abdomen in the female about one and a quarter times as long as

broad, about one-quarter entire length of body; posterior lobes

pointed and separated by broad sinus one-half length of abdomen.

The furcal appendages in both sexes lie at the bottom of the anal

sinus. Antennules of moderate size, basal segment with blunt

posterior process; second segment with stout anterior spine,

posterior small spiniform process, and lateral hook of medium

size. Antenne with short blunt process on basal segment.

BRANCHIURA FROM LAKE TANGANYIKA. 270

Immediately behind the insertion of the antennule is situated a

stout and moderately sharp accessory spine. Eyes rather large,

a little more than one-quarter breadth of carapace apart.

Suckers, in the males, rather small, -12--14 of breadth of cara-

pace; in the females, rather large, *14—-18 of breadth of carapace.

Maxillipeds strong and moderately well armed ; basal segment

produced into three broad rounded processes; ‘‘area” of fair

size, pear-shaped, and armed with small spines. A pair of

accessory processes, also broad and rounded, is situated between

the bases of the maxillipeds, and a second pair, rather slighter,

a Short distance behind them. Flagella present on the first two

pairs of swimming-legs. In the male, vesicula seminalis present

on the third leg, copulatory peg and accessory cushion on the

fourth. In the female, basal segment of fourth leg produced

postero-laterally into pointed lobe. Colour (in formol) light

brownish grey, sometimes with a pronounced reddish tinge, the

lateral and posterior regions of the carapace lobes showing a

distinct radial striation, most clearly seen by transmitted light.

Testes covered above by irregular blotches of brown pigment.

Dimensions as follows :—

Adult male (largest specimen). Adult female (largest specimen).

mm. mm.

Motalilemo taper ese 10-2 Motallene tines ceeee ito!

Length of carapace... 6°7 Length of carapace... 7:5

Breadth of carapace... 6°1 Breadth of carapace... 6°5

Remarks.—It is not unusual among the Argulide for the

posterior carapace lobes to exhibit some striation, which is due,

no doubt, to the branching of what Wilson calls the side-pouches

of the stomach. This appearance is, however, so outstanding a

feature of these specimens, that striatus at once suggested itself

as a suitable specific name. From the large number of specimens

contained in the collection, this would seem to be the commonest,

and perhaps the most widely distributed of the Tanganyika

species. ‘There are a few small individuals the sexual characters

of which are less obvious, and yet others which must be classed

as larval forms and do not show the secondary genital modifications

typical of the adult.

Occurrence.—-Mbete, 25/9/04. From gill-arches and roof of

mouth of large male specimen of Dinopterus cunningtoni (length

1020 mm.). Twenty-two males and twenty-five females.

Sumbu, 13/10/04. From gill-arches and roof of mouth of

large Clarias robecchii. One female, associated with specimens

of A. africanus.

Vua, 29/10/04. From surface of head and mouth of very large

female specimen (length 1550 mm., weight 30:6 kilogr.) of a

Siluroid (? Clarias lazera)—native name Muomi. A single male,

associated with specimens of A. africanus.

Kasawa, 1/11/04. From mouth and gill-bars of large Dino-

topterus cunningtont. Three male and two female specimens.

276 DR. W. A. CUNNINGTON ON

Kibwesi, 19/12/04. From head, mouth, and gill-bars of several

large specimens of Chrysichthys brachynema. One male and one

female, associated with specimens of A. rubescens.

Rumonge, 16/2/05. From mouth and surface of body of large

Auchenoglanis occidentalis, var. taunganicanus. Sixty-four males

and fifty-six females (some larval) associated with specimens of

A. incisus.

ARGULUS RUBESCENS, sp.n. (Pl. XLII. figs. 15,16; Pl. XLV.

figs. 31, 32.)

Description.— Carapace elliptical, longer than broad, with

moderate antero-lateral depressions. Posterior lobes broad and

rounded, covering bases of all but fourth swimming-legs, and

separated by a broad sinus, more than one-fifth the length of the

carapace. The edge of the carapace below is armed with small

spines, which extend to the region of the swimming - legs.

Abdomen, in the male, nearly one and a half times as long as

broad, about two-fifths entire length of body; posterior lobes

rounded and only separated by an insignificant sinus one-sixteenth

length of abdomen. Abdomen, in the female, slightly longer

than broad, about one-quarter entire length of body ; posterior

lobes rounded and separated by somewhat narrow sinus almost

one-quarter length of abdomen. ‘The furcal appendages in both

sexes he at the bottom of theanal smus. Antennules of moderate

size, basal segment with rounded posterior process; second

seginent with blunt anterior spine, posterior small spiniform

process, and lateral hook of medium size. Antenne with rounded

process on basal segment. Immediately behind the insertion of

the antennule is situated an accessory rounded process. Hyes

of moderate size, a little more than one-quarter breadth of

carapace apart. Suckers, in the males, rather large, :15 of breadth

of carapace; in the females, of moderate size, *13—14 of breadth

of carapace. Maxillipeds fairly strong and moderately armed ;

basal segment produced into three broad flattened processes ;

“area” of fair size, oval shape, and armed with small spines.

A pair of accessory flattened processes is situated between the

bases of the maxillipeds, anda second pair, rather slighter, a

short distance behind them. Flagella present on the first two

pairs of swimming-legs. In the male, vesicula seminalis present

on the third leg, copulatory peg and accessory cushion on the

fourth. In the female, basal segment of fourth leg produced

postero-laterally into pointed lobe. Colour (in spirit) reddish,

but with lighter areas anteriorly and round the eyes. Testes

covered above by irregular blotches of reddish brown.

Dimensions as follows :—

Adult male (largest specimen). Adult female (largest specimen).

Total length ............ 3:54 ‘Total length /15....98 5-4

Length of carapace... 2°3 Length of carapace ...° 4°

Breadth of carapace... 1:9 Breadth of carapace... 3

BRANCHIURA FROM LAKE TANGANYIKA. NU

Remarks.—This species, which in some features comes near to

the preceding one, may be readily distinguished from it by the

rounded nature of the abdominal lobes. The reddish coloration,

which is striking in the larger specimens, is less evident or almost

entirely absent in smaller ones. All the male specimens in the

collection, which are rather small, show little colour, except, of

course, the characteristic blotching of the testes. It will be

noticed that this form has only been obtained from the Siluroid

Chrysichthys brachynema, which is itself an endemic species.

Occurrence.—Moliro, 24/10/04. From roof of mouth of fair-

sized Chrysichthys brachynema. One female.

Kibwesi, 19/12/04. From head, mouth, and gill-bars of several

large Chrysichthys brachynema. 'Three males and nine females,

associated with specimens of A. striatus.

Mrumbi, 27/12/04. From inside mouth and outside body of

large Chrysichthys brachynema. One male and one female

specimen.

ARGULUS AFRICANUS Thiele.

Argulus africanus Thiele, Zool. Anz. vol. xxiii. 1901, p. 47.

Argulus africanus Wilson, Proc. U.S. Nat. Mus. vol. xxv. 1903,

1, Call

Argulus africanus Thiele, Mitt. Zool. Mus. Berlin, Bd. 2,

Heft 4, 1904, p. 37.

A preliminary description of this form was given by Thiele in

1901, and was followed by a more complete account with figures

in 1904. It is the only species of the genus Argulus which had

been previously obtained from the continent of Africa, and it

still remains the only species known outside the confines of Lake

Tanganyika. ‘Thiele speaks of it as a form widely distributed in

eastern Africa, and it is now possible to add other localities to

those hitherto known.

As I have examined more than 60 specimens from different

pavts of Africa, perhaps I may add a few remarks in modification

of Thiele’s detailed description. Between the average male and

female specimens there is by no means so clear a contrast as he

indicates in the relative proportions of the carapace (cephalo-

thorax). It is a fact, as I have determined by actual measure-

ments, that the carapace of a large female is broader in proportion,

and so less closely resembles the male condition, but except

in extreme cases of large females contrasted with small males,

there is but little difference in this respect between the two

Sexes.

Again, the carapace lobes in the female are stated to cover the

whole of the swimming-legs, leaving but the tips of the last two

pairs visible from above. ‘This may, perhaps, be the condition

of things in an individual greatly distended with eggs, but is

neither normal nor typical. In the great majority of the

females in this collection, the tips of all four pairs of swimming-

legs project from beneath the carapace, and in a few cases the

278 DR. W. A. CUNNINGTON ON :

bases of the fourth pair of legs are also visible much as in the

male.

The proportions of the abdomen are likewise indicated as

affording a distinction between the species, yet it appears that,

as a rule, the female abdomen is really longer than broad, the

measurements agreeing closely with those of the male, though

of course the shape is distinctly different in the two cases.

While these additional observations slightly modify the account

given by Thiele, there can be no reasonable hesitation in

identifying the species, which is readily distinguished by the

conspicuous pigment spots and the nature of the antenne and

maxillipeds.

TANGANYIKA.—The species is apparently quite common in this

lake, as the collection contains more than thirty specimens

obtained from various localities. It does not appear to be

confined to one host, and it was in some instances found in

association with the endemic species of Argulus.

Occurrence.—Sumbu, 13/10/04. From gill-arches and roof of

mouth of large specimen of Clarias robecchi. Four males and

four females. A female of A. striatus was associated with these.

Sumbu, 20/10/04. From branchial arches and roof of mouth

of two smallish specimens of Clarias robecchti. Six males and

six females.

Vua, 29/10/04. From surface of head and mouth of very

large female specimen (length 1550 mm., weight 30°6 kilogr.)

of a Siluroid (? Clarias lazera)—native name Muomi. Five

males and five females, with which a single A. striatus was

associated.

Kala, 18/11/04. From mouth-cavity and gill-bars of large

Lates microlepis (length 910 mm.). Two females, associated with

specimens of Dolops ranarum.

Ndanvie, 8/2/05. From surface of head and body of large

Lates microlepis. One male. Specimens of A. rubropwnctatus

were associated with this.

VicroriA Nyanza.— From this lake, too, specimens of

A. africanus were obtained by the Expedition for the first time.

All of them, as it happens, were collected at Bukoba, on the

western shore, the place at which Stuhlmann first discovered

Dolops [Gyropeltis| ranarum. Curious to relate they were also

associated in every case with specimens of the latter, living with

them on a common host.

Oceurrence.—Bukoba, 24/4/05. From surface of body, mouth

and gill-chambers of a large Siluroid (? Bagrus degeni)—native

name WVfui. One female specimen, associated with a number of

Dolops ranarwm.

Bukoba, 24/4/05. From surface of body and inside mouth of

large Protopterus cethiopicus (length 1040 mm.). Four males,

five females, associated with Dolops ranarum.

Bukoba, 24/4/05. From surface of body, mouth, and _ gill-

chamber of large Siluroid (? Clarias angwillaris)—native name

BRANCHIURA FROM LAKE TANGANYIKA, 279

Nshonzi. A single female, associated with specimens of Dolops

ranarum.

In the collection of the British Museum are other specimens

which have been subsequently obtained. They were also

associated with a Dolops ranarum. The label reads:—‘‘ On

skin of fish (‘Male’) @catfish. Lake Victoria. Collected by

G. D. H. Carpenter, 6.xi.11.” Five male and fourteen female

specimens. |

Apert Nyanza.—Asthe result of a rather unusual occurrence,

we now know of the existence of Argulide in this lake. Ina

tow-netting from the lake which was placed in my hands for

examination by my friend Dr. R. T. Leiper, Helminthologist

to the London School of Tropical Medicine, I found very un-

expectedly, a young Argulid. The material was collected by

Dr. Leiper in July 1907, when he accompanied the expedition

despatched to Uganda by the Egyptian Survey Department, and

is of special interest at the present time, being the only col-

lection of plankton as yet made in the lake. The tow-netting

was taken at the north end of the lake opposite Magungo.

While we know of the free-swimming habit of young Argulids,

which would render them liable to be taken in the tow-net, it is

certainly very exceptional to capture them in this manner, as

is evidenced by their complete absence from the extensive series

of tow-nettings made during the Third Tanganyika Expedition.

It is thus a very happy accident which gives us this additional

record.

I believe the specimen to bea male larval form of A. africanus.

The blunt rounded spines or processes on antenne and maxillipeds

which are characteristic of the adult, are represented by sharper

structures in this individual, but I think there is evidence that

this is usual in young specimens.

River Nrue.—In the British Museum collection, associated

with the specimens of Dolops obtained by Mr. Loat, and referred

to above, there is a single female which I regard as belonging to

this species. There are, it is true, points of difference from a

typical specimen, yet in view of its general similarity, and since

this form is known to occur in the Nile and is commonly found

in association with Dolops, it seems desirable to place it here.

It was taken from the buccal cavity of a large Heterobranchus

bidorsalis, caught at the mouth of Lake No, White Nile.

3. General Remarks.

With singular uniformity, all the collections made in Tanganyika

prove, to a greater or less degree, the unique character of the

organisms inhabiting the lake. The Branchiura are no exception

to the rule, for, as a result of this Expedition, we now know

that while two species are widely distributed in Africa, they are

associated in Nyasa with a single endemic form, but in Tanganyika

with no less than seven, Thus in this case again, Tanganyika

280 DR. W. A. CUNNINGTON ON

is shown to possess not only a considerable number of character-

istic species, but a much richer fauna than the other great

African lakes, with which we may reasonably compare it. In

the adjoining table of distribution, these facts are expressed in

graphic form.

Tabie of Distribution.

Tato. | Other

Name of Species. Tanganyika. Rear Nyasa. eee parts of

! Atrica

| |

| | St peo

Dolops ranarum ........--.+-+.| + + + +

Argulus imeisus ............+-.| + |

35 rubropunctatu + | |

2p Persomatus ......... ot |

0 CEXUGWUS ........2.-+-- ar |

3 angusticeps ......... ar

3 SAPICUBUID, ann snecee p00 205 +

| ay PUDESCENS ..........-. + |

a CH FUOHDONS 34,020 0n0 0% + oF + + +

Chonopeltis inermis ......... | ge sake |

| |

10 species. 9 2 | Shy ulead 2

Taking a general survey of the specimens which have been

examined, there are a few points which call for mention.

Speaking broadly, there is no great disparity in the numbers of

the two sexes in the collection. Only in the case of Argulus

rubropunctatus was there a marked preponderance of one sex,

a total of 45 males accompanying 15 females. There were

occasional isolated individuals, but both sexes were usually

associated on one host. As a rule, females were larger than

males, which seems to be commonly the case.

It is difficult to understand why, in certain species, the relative

size of the suckers is different in the sexes. The most striking

example is that of A. striatus, where a series of measurements

ave the following results:—-Sucker diameter in males -12—14

of breadth of carapace: in females, -14—18 of breadth of cara-

pace. Such a degree of variation cannot, in this case, be merely

an index of carapace diversity in the sexes—short and broad,

for instance, contrasted with long and narrow—since these males

and females are very similar in this respect. Asa rule, where

we have sufficient evidence to go upon, the male appears to have

a smaller sucker in proportion than the female.

Concerning the relations of these forms to their hosts, some

further observations may be offered. Both in Victoria Nyanza

and Tanganyika, the smaller fish appear to be relatively little

infested by the parasites. These were most frequently observed

upon large fish, and particularly on Siluroids, the scaleless nature

of which must render them especially liable to attack. Large

BRANCHIURA FROM LAKE TANGANYIKA. 981

specimens of the Tanganyika Serranid Sates microlepis also

usually bore some examples attached to the surface of the head

or within the mouth-cavity.

It is commonly considered that these parasites do not constitute

a menace to the fish on which they prey. This was equally the

impression made in the present instance. Cases have been

recorded, however, in which it seems clear that an excessive

number of Argulids infesting a fish may actually cause the

death of their host. What appeared to be a striking instance of

this came to my notice during this Expedition, and certainly

merits a few words of description. Nearing the north-eastern

shore of Tanganyika one afternoon, after a long voyage, a large

fish was noticed floating on the surface of the water. My men

fetched it on board, and it proved to be a specimen of the

“kavungwe” (Auchenoglanis occidentalis, var. tanganicanus),

absolutely covered with Argulide. There was literally hardly a

spot on the body not infested with the parasites, and in the head

region and within the mouth they were positively overlapping

and lying on top of one another. Instead of detaching isolated

specimens, as had usually to be done, a layer of writhing Argulids

could be scraped from most parts of the body. The fish was not

only perfectly fresh, but was still feebly making convulsive

movements, and one could not resist the conviction that it was

succumbing to the fierce attack of such a swarm of parasites.

The latter, in this instance, did not all belong to one species.

There were specimens of both Argulus striatus and A. incisus,

the last-named species, strange to say, being obtained on this

occasion only.

It was no very unusual thing on Tanganyika to discover the

bodies of large fish floating far from shore. The crew of my dhow

kept a sharp lookout, and made a point of picking them up when

possible, as it meant a little addition to their fare, unless the

fish were too putrid for even their taste. I never saw Argulide

under these circumstances, except in the case detailed above,

but in no other instance was the fish still living. The suggestion

nevertheless occurs, that these also may have been victims of

such parasites as appear to have caused the death of the

Auchenoglanis.

No doubt, as is usually the case, the different types of Argulid

are not rigidly confined to one host, though they may have a

preference for a particular fish. Thus Argulus rubropunctatus,

taken on different occasions and in different localities, was

obtained only from Lates microlepis ; similarly, A. rwbescens was

found only on Chrysichthys brachynema. On the other hand,

A. striatus, perhaps the commonest and most widely distributed

Tanganyikan form, was obtained from a number of hosts, whilst

A. africanus and Dolops ranarum, owing to their extended

- distribution, clearly have still larger host-lists.

The question suggests itself—do.the Tanganyika endemic

species infest endemic fish @ Since the number of endemic types

Proc. Zoot. Soc.—1913, No. XIX. 19

282 ON BRANCHIURA FROM LAKE TANGANYIKA,

of fish in Tanganyika is so great (some 101 out of 115) it is not

surprising that they constitute a majority of the actual hosts,

Still there is no reason to suppose that there is any significance

in such a connection from what we know of the habits of these

parasites, and we find, in fact, an endemic Argulus. living on a

widely distributed host, and widely distributed Argulids (Dolops

ranarum and Argulus africanus) living in Tanganyika on endemic

fish.

Not infrequently two species were found living together on a

common host. It is remarkable that Dolops ranarwm was almost

invariably found associated with Argulus africanus. On the

only occasion on which the former was obtained in Tanganyika,

it was, as usual, accompanied by individuals of the latter.

While one is hardly prepared to say that there is a real signi-

ficance in this association, it is certainly strange that these two

widely distributed forms should appear to range together in the

localities from which they come. There-is no such constancy

in the occurring together of other types. A. rubropunctatus,

though taken on several different occasions, was always found

alone. .

The question of distribution within the limits of each lake

was hardly likely to prove interesting. Apart from forms

obtained on a single occasion, there is no evidence that any of

the species are confined to a particular region.

EXPLANATION OF THE PLATES.

Pratt XLI.

Fig.1. Argulus incisus (p. 268). Male, dorsal view. XX 53.

2. 55 Female, dorsal view. X 53,

3. Ar ulus rubropunctatus (p. 269). Female, dorsal view. X 54.

4. 5 os Male, dorsal view. X 5.

5. Male, ventral view. X 65.

6. Argulus personatus (p. 271). Male, dorsal view. X 6.

the is 35 Female, dorsal view. x 44.

8. ) SS Female, ventral view. X 42.

Piate XLII.

Fig. 9. Argulus exiguus (p. 272). Male, dorsal view. X 6.

10. Argulus angusticeps (p. 273). Female, dorsal view. X 53.

11. eens ventral view. X 53.

12 Ar ulus striatus (p. 274). Female, dorsal view. X 6.

13. 5 33 Male, dorsal view. X 6.

14. Male, ventral view. X 63.

15. Argulus rubescens (p. 27 6). Female, dorsal view. X 63.

16. ie, a Male, dorsal view. X 63.

Pruate XLITI.

Fig.17. Argulus incisus (p. 268). Antennule and antenna. > 60.

18. 3 Maxilliped. X 50.

19. Ar gulus rubropunctatus (p. 269). Antennule and antenna. X 40.

20. Maxilliped. X 30.

21. Argulus ewiguus (p. 272). Antennule and antenna. X 100.

22. 9 3 Maxilliped. > 100.

ON PLANKTON FROM NEW BRUNSWICK. 283

Prats XLIV.

Fig. 23. Argulus personatus (p. 271). Posterior three swimming-legs to show

accessory copulatory apparatus. xX 60.

24. He a5 Antennule and antenna. X 60.

25. 59 35 Maxilliped. X 60.

26. Argulus angusticeps (p. 273). Antennule and antenna. X 60.

yet (aes 5 Maxilliped. X 60.

Pratt XLV.

Fig. 28. Argulus striatus (p. 274). Posterior two swimming-legs to show

accessory copulatory apparatus. > 40.

29, 5 > Antennule and antenna. X 60.

30. - - Maxilliped. x 60.

31. Argulus rubescens (p. 276). Antennule and antenna. X 60.

32. S :s Maxilliped. x 60.

22. Notes on Plankton collected across the mouth of the

St. Croix River opposite to the Biological Station at

St. Andrews, New Brunswick, in July and August 1912.

Jar seusamstore Naoussa, IDES es Wiclawon, 110 Se.

McGill University, Montreal.

[Received January 8, 1913: Read March 4, 1913.]

(Text-figures 54 & 55.)

InDEx.

Geographical Zoology : Page

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Clad Oceana eer erwtes arte cea enn Maen Se ne nectar Bae techs OE

TECEOA KH COUIID Rech cee BaeMDHaO SAR ENO ts yAOBEES AOROL TOS HDAE HOE SCODncnaEL OnE eld)

Ethology :

Brac hio lanier scsedseceressvenna tse sae Patan ceo ee ete OO

Morphology :

Bolina and position of Céenoplana.........ccccc0cveeeeec ee, 288

Systematic :

Actinotrocha and Phoronis brownei Sélys-Longchamps 290

The marine Plankton of the Atlantic coast of Canada has been

examined qualitatively by Professor Ramsay Wright (‘The

Plankton of Eastern Nova Scotia Waters,” 39th Ann. Rep. Dep.

Mar. & Fish.: Further Contributions to Canadian Biology 1902-

1905. Ottawa, 1907, pp. 1-19,7 plates); and the Phytoplankton

more particularly by Professor L. W. Bailey (‘The Marine and

Estuarine Diatoms of the New Brunswick Coasts,” Bulletin Nat.

Hist. Soc. New Brunswick, No. 28, vol. vi. 1910, pp. 219-239,

2 plates).

The estuarine or tidal Zooplankton off the mouth of the

St. Croix river does not seem to have received special attention

hitherto. As for the quantitative determination of this tidal

plankton, and the systematic tabulation of its periodical fluc-

tuations, an endless field of work is offered to the investigator.

Samples taken in a tow-net weighted to a depth of 3-5 fathoms

Loe

284 DR. ARTHUR WILLEY ON

are often very rich in individuals of Copepoda, Cladocera,

Diatoms, and Dinoflagellates. The dominant constituent of the

plankton-at all times is the Diatom, Coscinodiscus, which adheres

so tenaciously to the bottom of the watchglasses that it obscures

the remaining contents. This Diatom genus, together with

Biddulphia, is recorded as forming the main part of the Phyto-

plankton in the brackish water of the Weser river (Ch. Brockmann,

1906; see A. Steuer, ‘ Planktonkunde,’ Leipzig and Berlin, 1910,

p. 36).

; mh character of the water in the tract under survey is defined

by the predominance of Coscinodiscus. and the value or interest

of the records accordingly depends upon this circumstance. Of

special note in this regard was the occasional appearance of

Fritillaria borealis Lohmann, an Arctic Appendicularian asso-

ciated bionomically with the Copepod, Calanus jfinmarchicus.

The Fritillaria was always more or less damaged by the force of

its impact with the tow-net. The Calanws was very rare and

always immature; each ramus of the fifth pair of legs was

bi-articulate instead of being 3-jointed, as it is typically.

H. Lohmann (‘ Die Appendicularien der Plankton-Exped.’ 1896,

Taf. viii. f. 6, p. 49) has recorded #. boreahs also from the

Antarctic Ocean, but it is not found in the intervening warm

regions. L. W. Williams (‘“ Notes on Marine Copepoda of Rhode

Island,” Amer. Nat. vol. xl. No. 477, Boston, 1906, pp. 639-660)

says that C. fiumarchicus appeared abundantly in tows taken in

Naragansett Bay in January, but was found at no, other time.

Professor Ramsay Wright (op. cit. p. 13) says it was very

abundant in the earlier part of the summer at Canso. At

St. Andrews I noted its occurrence specifically and singly on

July 30th and August 5th; the length was 3 mm.

The characteristic and abundant Calanoids at this time were

Acartia clawsi Giesbrecht, Tortanus discaudatus Thompson &

Scott, and Hurytemora herdmant Thompson & Scott. Of these,

A. clausi was the most abundant; and this is noteworthy, imas-

much as this species was not found at Woods Hole during July

and August 1899, although a related species, 4. tonsa Dana, was

one of the commonest copepods in tow taken from the wharf of

the U.S. Fish Commission at that station (W. M. Wheeler, ‘The

free-swimming Copepods of the Woods Hole Region,” Bull. U.S.

Fish. Comm. vol. xix. pp. 157-192. Washington, 1901). L. W.

Williams (op. cit. 1906) found Acartia tonsa abundant throughout

the summer in Charlestown Pond, Rhode Island, where it was

the predominant copepod in the tow; he also found A. clausi

abundant in Naragansett Bay in January and February.

Thompson and Scott have recorded A. clawsi from the Gulf of

St. Lawrence. It is often coloured with blue spots in pairs

below; and it has a large quivering eye with two lenses.

Tortanus discaudatus is chavacterised by its dark brown caudal

furea with variable and unequal rami, the right larger; often

PLANKTON FROM NEW BRUNSWICK, 285

with the basal lobe of the right outer seta enlarged to form a

conspicuous process upon the right ramus. Professor Ramsay

Wright found it exceedingly common at Canso from the end of

July to the middle of August; he offers an explanation of the

asymmetrical tail based upon the mode of attachment of the

spermatophore (op. cit. p. 14). This is very likely correct, but I

cannot confirm the statement that the asymmetry is greater in

the female than it is in the male. It was not so in my obser-

vations at St. Andrews. The species was originally described

from the Gulf of St. Lawrence in 1897 (I. C. Thompson and

A. Scott, * Notes on new and other Copepoda in Plankton col-

lected continuously during two traverses of the North Atlantic,”

Proc. and Trans. Liverpool Biol. Soc. vol. xii. 1898, pp. 71-82,

pls. 5-7). It was again described as Corynura bumpusii by

W.M. Wheeler, from Woods Hole and Vineyard Sound (op. cit.

1901). L. W. Williams (op. cit. 1906) first suggested the identity

of Corynura bumpusit with Tortanus discaudatus; but he de-

scribed an allied form, abundant in Naragansett Bay and

Charlestown Pond, as a new species, 7’. setacaudatus, differing

from discaudatus, as it would appear, chiefly in the character of

the fifth legs in the female, which carry spines in the former and

are without spines in the latter species. The eye of 7’. discuu-

datus resembles that of Acartia clausi and quivers in like manner.

It would be worth while to make a biometrical study of its

remarkable caudal furca. Length of ¢ 1:75 mm.

Eurytemora herdmani Thompson and Scott, is to be distin-

guished from allied species of the genus, especially from #. hirun-

doides, by the structure of the fifth legs in the female. Females

with ovisac were noted on August 12th; and on the same date,

males with the tumefied central part of the right antenna scarlet.

L. W. Willams (op. cit. 1906) recorded three species from the

Rhode Island region: #. americana, sp. n., LH. hirwndoides, and

L. herdmani.

The ebbtide plankton of August 9th was largely composed of

Chetoceras and Rhizosolenia, Tintinnoids, Syncheta, Acartia,

Nauplu, some Pritidlaria borealis, and the Cladocera: Huadne and

Podon. Podon occurred in almost every tow, and often contained

mature embryos in the dorsal brood-sac. Professor Ramsay

Wright mentions two species of each genus as occurring at Canso

(op. cit. 1907, p. 13). In the same year, what he claimed to be

the first American records of Hvadne nordmanni Lovén, and

Podon polyphemowles Leuckart, were made by L. W. Williams

(‘‘ List of the Rhode Island Copepoda, Phyllopoda, and Ostracoda,

with new species of Copepoda.” Special Paper, No. 30, 1907.

Reprinted from 37th Ann. Rep. of the Commissioners of Inland

Fisheries of Rhode Island, pp. 69-79, 3 plates).

Of Dinoflagellates, Peridiniwm divergens Khrb. was sometimes

very abundant. In the peripheral part of the cytoplasm there

286 DR. ARTHUR WILLEY ON

were numbers of relatively large, bright red bodies with a

variable and irregular contour. These bodies were not mentioned

in Doflein’s treatise on the Protozoa ; but were duly described by

F. Schiitt (‘Die Peridineen der Plankton-Expedition,’ i. Theil.

27 plates. Kiel und Leipzig, 1895). This author calls the red

bodies in P. divergens, hygrosomes (p. 84), and attributes their

formation to similarly shaped plastids, termed hygroplasts (p. 74).

He found that their contents are of a fatty nature, blackening

with osmic acid and dissolving in ether (p. 85). It is possible

that they may exhibit the property of phosphorescence, though I

had no opportunity of testing them for ee quality.

L. Plate (“ Pyrodinium bahamense, n. g., n. sp.,” Arch. Protis-

tenk. Bd. vii. 1906 ; see Steuer, op. ai. p- "307) suggested that the

phosphorescence of Pyr odininum depended upon the oxidation of

the numerous oil-drops at the hinder end. Biuitschli mentions

numerous fat-drops in Peridiniwm divergens, which, according to

Pouchet, often form an annular zone parallel with the transverse

groove; and this same species has been designated the luminous

Peridinium of the Gulf of Trieste.

Brachiolaria, the larva of Asterias, appeared in the tow from

August 10th, and attracted my particular attention on account of

the three papilliferous adhesive processes with a median sessile

sucker between them, upon the preoral lobe. ‘This Brachiolaria

is virtually identical with that of <Asteracanthion pallidus

(= Asterias vulgaris Stimps.) as figured by Alexander Agassiz

(‘Embryology of the Starfish,’ 1864; see Selections from

Embryological Monographs compiled by Alexander Agassiz,

Walter Faxon,and E. L. Mark. II. Echinodermata. Cambridge,

Mass., 1883). Agassiz stated that A. vulgaris reproduces only in

the third week of August at his more southerly station.

Behind each of the three adhesive processes, which are called

‘‘brachiolarian arms,” there is a long tentacle-like process which

may be referred to as a bipinnarian arm. As the term implies,

the ciliated band is continued upon the bipinnarian arms but not

upon the adhesive processes. On the other hand, the hydroceel,

which can be traced forwards from its pore-canal, sends a diver-

ticulum into each of the adhesive processes but not into the

bipinnarian arms.

The adhesive processes, or “‘ brachiolarian arms,” are structures

sui generis, but they are paralleled by the three adhesive processes

of the Ascidian larva. There is this in common, that they are

developed upon the preoral region of the larval body; they serve

for the permanent (Ascidian) or temporary (Asterias) fixation of

the larva, and they consist of one median dorsal process and two

lateral processes. It would be hard to find a closer convergence

(parallel adaptation) in the category of adhesive organs. Without

presuming that they stand for anything more than physiological

adaptations, it seems legitimate to infer that some indirect

support, by way of analogy, is given to my interpretation of the

PLANKTON FROM NEW BRUNSWICK. 287

anterior lobe which carries them in the Ascidian larva, as a

preoral lobe (** Studies on the Protochordata,” 1893, Quart. Journ.

Micro. Sci. vols. 34 & 35); a suggestion which the late W. K.

Brooks was unable to accept.

Apart, however, from questions of morphology, the comparison

between the three adhesive processes of Brachiolaria and those of

the Ascidian tadpole cannot fail to be appropriate. I am not

aware that it has been made before.

Th. Mortensen (‘ Die Echinodermenlarven der Plankton-

Exped.’ 1898) figured the Brachiolaria of the European Asterias

rubens, and adds that it is so much like that of A. vulgaris that a

separate description is unnecessary ; but as the one larva occurs on

the North American coasts, and the other on the North Kuropean

coasts, no confusion is possible. I do not know of any data

which would enable a distinction to be made between the larva

of A. vulgaris and that of the associated species, A. forbesit.

Text-fig. 54.

Semidiagrammatic view of Brachiolaria attached to a piece of seaweed.

The larva swims through the water with the three anterior

bipinnarian arms curving backwards, the remaining eight bipin-

narian arms trailing behind. The anterior medio-dorsal adhesive

process is directed forwards; the lateral adhesive processes may

be bent forwards or backwards. The median process develops in

advance of the lateral processes and is already formed, with its

contained branch of the hydroccel, whilst the lateral processes

are represented by rudimentary ectodermal thickenings with a

minute projection of the hydroceel below each of them.

288 DR. ARTHUR WILLEY ON

I observed the fixation of the larva to a fragment of red sea-

weed by means of the adhesive processes, and kept it overnight,

alive and attached. The bipinnarian arms became much wrinkled

and constricted ; the adhesive processes were stretched wide apart

so as to allow the median suctorial pad to touch the seaweed

between them. The quinque-radiate dise swayed vertically and

freely in the water. The accompanying semidiagrammatic sketch

(text-fig. 54) will give a better idea of the attached larva than will

a verbal description.

A figure from another aspect, after Johannes Miiller, of a

Brachiolaria fixing itself in a different attitude, is reproduced in

the Cambridge Natural History (Vol. i. 1909, Echinodermata,

by E. W. MacBride, see p. 613).

I was at first puzzled by the statement in Delage et Hérouard

(Traité de Zoologie Coneréte, T. iii. Les Echinodermes. Paris,

1903, see pp. 80-81) that the median epithelial pad, between the

bases of the brachiolarian arms, is a ciliated sucker (ventouse

ciliée,) as I had observed no sign of ciliary action upon it. Ina

subsequent investigation Delage himself found no indication of

ciliation of the median suctorial pad (Yves Delage, “ Elevage des

larves parthénogénétiques d’Asterias glacialis,” Arch. zool. expér.

sér. 4, 11. 1904, see p. 38).

In larvee of Asterias glacialis raised by Delage from unfertilised

eges treated with carbonic acid, the paired adhesive arms

appeared first, towards the middle of the third month of the

culture. Brachiolaria which had reached the fourth month

in Delage’s experiments, showed clearly the initial phenomena

of the metamorphosis; they ceased to swim, the bipinnarian

arms became shrivelled, and the larve adhered by the mobile

brachiolarian arms, occasionally shifting their position.

Amongst the numerous larve seen by me at St. Andrews,

many of which were sketched from life, I only observed the

fixation to occur on the part of the oldest, when the asterodise

was most opaque.

The larva does not remain fixed irrevocably at the first spot

where it alights upon the bottom; but it never swims again.

Therefore it cannot be said that the adhesive arms serve merely

for occasional attachment. They represent a definite stereotropic

mechanism.

Bolina.—On August 14th at low tide,about 7 a.m., Dr. Huntsman

found quantities of Bolina alata at the foot of the wharf, adrift

in shallow water. I embraced the opportunity thus afforded of

making a fugitive study of this remarkable and fragile Ctenophore

which defies all methods of preservation.

L. Agassiz (‘Contributions to the Natural History of the

Acalephz of North America.” Mem. Amer. Acad., Boston, 1849)

described the various attitudes of Bolina, including one in which

the characteristic mantle-lobes are spread out lengthwise, at

right angles to the secondary axis in which the tentacles lie, and

PLANKTON FROM NEW BRUNSWICK. 289

parallel with the axis of the mouth and polar sensory field. The

two pairs of ciliated auricles are a distinctive feature of Bolina.

' The auricles and the reduced inconspicuous tentacles between

_ them occur on either side of this extended pallial axis. When

seen from the aboral aspect, as figured by Agassiz, with the lobes

flattened out, the organism exhibits a pronounced biradial sym-

metry, the mouth-cleft and the polar field being parallel to one

another in the direction of the longer axis, which coincides with

the plane of the flattened stomodeum, whence it is called the

stomodezeal axis.

In his memoir on Pleurobrachia, Agassiz, in spite of some

textual confusion, clearly considered the stomodeal axis to cor-

respond with the sagittal axis of bilateral animals, the tentacular

axis with the transverse. Later writers continued the discussion.

Korotneff’s genus Ctenoplana, which I obtained for the second

time in New Guinea waters (“On Otenoplana,” Quart. Journ.

Micro, Sci. vol. xxxix. 1896), seemed to me at that time to be

capable of throwing light on this question. Professor Arnold

Lang, of Zurich, had compared the Ctenophore with the Polyclade

organization, and I, for one, was penetrated with the conviction

that biradial symmetry must have preceded bilateral symmetry.

It is now clear that there is no such consecutive necessity ; and

it appears that the dorsiventrality of Ctenoplana, approximating

it to the facies of a flatworm, is a very interesting piece of

convergence.

The question has been re-opened quite recently by Th.

Mortensen (‘‘ Ctenophora.” Danish Ingolf-Exped., Copenhagen,

1912), apropos of the new sessile Ctenophore, 7jalfiella tristoma

rtsn.

Incidentally Mortensen throws doubt upon the aecuracy of my

figures of sections showing the peculiar genital ducts of Ctenoplana

korotne#it ; but in so doing he is hardly justified*. I may say,

however, that if the morphological comparison between the

secondary axes of Radiata (s. sé.) and the principal axes of

Bilateralia were not fallacious, then the relations exhibited by

Bolina would point to the opposite identification to that which I

advocated on the basis of Cienoplana (tentacles in the sagittal

plane), but to that which Agassiz advanced on the basis of

Pleurobrachia (tentacles in the transverse plane).

R. Woltereck (“ Wurmkopf, Wurmrumpf, und Trochophora,”

* In Ctenoplana korotneffi there were four testes only, in two pairs, paired

about the tentacle axis. In one individual there were twelve ducts distributed

equally among the four testes. Of these ducts I observed the actual opening to the

exterior in six instances, in section. In another individual seven ducts altogether

were counted in a series of sections.

With regard to the systematic position of Ctenoplana, Abbott says “it must be

conceded that Ctenoplana stands midway between Celoplana and the Cydippid

Ctenophores, in regard to its primitiveness or its degeneracy.” And again: “The

weight of the morphological evidence bears out the conclusion that Ceeloplana is a

very highly specialized Ctenophore derived from the Cydippida.” (James Francis

Abbott: “ The Morphology of Celoplana,” Zool. Jahrb., Anat. xxiv. 1907, pp. 41-

70, pls. viii.—x. and 7 text-figures.)

290 DR. ARTHUR WILLEY ON

Zool, Anz. xxvili. 1904, pp. 273-822), disposing of excellent data,

maintained that the change from a radial to a bilateral form is

best understood by the transition from the pelagic to the

benthonic habit, on the assumption that Bilateralia are to be

derived from pelagic, radiate, marine animals. He reconstructs,

in imagination, the ancestral type of Annelids, as a pelagic,

spherical, octoradiate, bipolar Ccelenterate, to which he applies

the phylogenetic designation: Octoradiata-Bipolaria. His idea

of the divergent descent of Ctenophora and Trochophora may be

expressed In graphic form :—

Bipolaria.

Bee cipaa ee

erase ee

Trochophora. Ctenophora.

Woltereck refers quite simply to Ctenoplana as a “ creeping

Ctenophore.” As a creeping Ctenophore its most conspicuous

distinctions are a permanent dorsiventrality, the habit of

reptation, the presence of pairs of organs, and the possession

of a circular mouth. These four qualities are shared by the

Polyclade Turbellarians, with bilaterality in addition. Never-

theless, in comparison with the flatworms, these features of

Otenoplana are the result of convergence; it is not in the least

degree related to the flatworms because of its flatworm habit.

To those who have not seen Cfenoplana this may have trifling

significance; but after seeing it in the living animal, it becomes

fraught with meaning, bearing upon the principles and effects of

adaptation.

Actinotrocha, the larva of Phoronis, appeared in the tow, a

solitary example, on August 19th, from about five fathoms. The

typical long tentacles were marked with brown pigment; and

behind their bases were the small replacing tentacles, indicating

that the metamorphosis was at hand. On the preoral lobe or

hood, there was a projecting ectodermal knob, a papilla, ciliated

and presumably sensory, a little ventral to the apical plate. Such

an ectodermal knob on the hood is figured by Mare de Sélys-

Longchamps in Actinotrocha browner from Plymouth, England,

the larva of an undiscovered Phoronis. This larva bears a striking

but resemblance to the one at St. Andrews, and [ have no doubt but

that it isthe same species. [M.de Sélys-Longchamps, ‘“‘ Phoronis,”

Fauna u. Flora des Golfes von Neapel, 30th Monograph. Berlin,

1907; see p. 190 and pl. xi. fig. 31.]

In front of the anal peritroch, on the ventral side, was an

oblique depression, marking the site of the invagination of the

metasome into the interior of the larva.

The eversion of the Phoronis-body or metasome took place

about 3 p.m. in the watchglass. As I had not expected such a

PLANKTON FROM NEW BRUNSWICK. 291

rapid transformation, it happened whilst I was not looking.

Brown-coloured débris appeared in the intestine; and the

definitive Phoronis-tentacles were without pigment.

A conspicuous blood-vessel with red corpuscles traversed

the length of the body of the young Phoronis, transmitting

branches into the tentacles.

The aboral extremity of the body was very mobile, adhering

with active, metabolic movements to the glass. Retractile,

digitiform, vascular processes, termed cecal capillaries, were

pushed out in rapid succession from the main vessel into the

mobile foot-dise and were withdrawn again at frequent intervals

(text-fig. 55). I am not aware that the retractility of these

first-formed aboral cecal capillaries has been noted previously.

Text-fig. 55.

49,

0,0,

Nene

00,

“OM

“

)

Mee

Posterior end of a young Phoronis immediately after the metamorphosis, showing

the retractile vascular processes in the mobile and adhesive foot.

The above-mentioned foot-dise corresponds with the terminal

portion only of the ampulla of the fully-formed Phoronis. The

excal capillaries of the ampulla are figured by Sélys-Longchamps

in a young individual of Phoronis sabatiert (see his pl. viii.

fig. 13). This author describes, on p. 35 of his monograph, the

special and independent contractility of the terminal portion of

the ampulla. The peculiar mobility of this part is associated

with a situs inversus of the muscular layers, the longitudinal

292° MISS EDITH E. BAMFORD ON THE

fibres being external and the circular fibres internal (op. cit. pl. v.

fig. 4). As a consequence, the lower end of the body exhibits

great variability of form, sometimes presenting the same diameter

as the rest of the ampulla, at other times dilating to a sphere,

and again attenuating to an elongated process. Its special

function in the adult is to bury the animal in the mud of the

sea-floor.

The function of the foot-dise of the newly-transformed Phoronis

seems to be the attachment of the body to the substratum. It is

desirable to distinguish between a postlarval and an adolescent

phase. So far as I am able to judge from the work of M. de

Sélys-Longchamps, the peristaltic contractility of the hinder

division of the ampulla in the mature or submature Phoronis is

not identical with the metabolic contractility of the foot-dise in

the postlarval phase, when it is associated with, though not

in any way caused by, the retractility of the cecal capillaries at

this stage. In other words, the formation of the adhesive

foot-dise precedes that of the burrowing ampulla.

IT believe this is the first record of the occurrence of Phoronis

on the eastern coast of Canada. No specific description can be

based upon a single specimen which has just undergone the meta-

morphosis; but if my identification of the larva is correct, then

the species may be named provisionally Phoronis brownet.

In conclusion J desire to acknowledge the kindness and

attention which I received from Dr. A. G. Huntsman, the

energetic and obliging Superintendent of the Station.

23. Variations in the Skeleton of the Pectoral Fins of

Polypterus. By Epvrru EK. Bamrorp, Newnham College,

Cambridge *

[Received January 22, 1913: Read March 18, 1913.]

(Text-figures 56-60.)

InDEX.

Page

Variation and Altiology ............sceseeceeeeecereeeeeeees 293

It was suggested to me by Dr. Gadow that the examination

of the individual variations of the ample material of voung and

adult Polypterus collected by Budgett in his Gambia and Niger

Expeditions, now the property of the Cambridge University,

might account for the discrepancies which are found in de-

scriptions of the pectoral fins of Polypterus. After examining

the material, it becomes evident that these discrepancies are

mostly the result of the many individual variations and of the

former scarcity of material. A description of the variations

found in the ample material at my disposal is of future value ;

* Communicated by Dr. H. GApow F.R.S., F.Z.S.

PECTORAL FINS OF POLYPTERUS. 293

if can be only a question of time for the ontogeny of this

important type of fins to be studied exhaustively, whilst embryo-

logical research cannot easily take up the question of the extent

of the amplitude of variations.

The material examined consisted of 102 specimens of Polypterus

lapradit and Polypteruws senegalus, varying in length from 12

to 43 cm.

Specimens have been previously described by Gegenbaur™*,

Klaatsch ft, Pollard, and Budgett9$. Unfortunately of larval

Polypterus only one specimen is known; this was found and

described by Budgett. In his description, he says: “The shape

of the fin-blade is triangular, the metapterygial border forming

with the distal border an acute angle, while the pr opterygial

border forms with the distal border an obtuse angle.” None of

the specimens [I examined had the larval shape; the youngest,

12 em. long, had already acquired the characteristic form of the

adult, the distal border of the mesopterygium being rounded

and produced near the metapterygium some distance beyond the

latter, thus forming a projection at this place. It seems probable

that the peculiar form of the pectoral fin, which has given

rise to so much controversy and which caused Gegenbaur and

Klaatsch to think it consisted of a stem carrying a biserial

arrangement of radials (a biserial archipterygium), is in a large

measure due to the partial ossification of the fin and the sub-

sequent unequal rate of growth of the bony and cartilaginous

parts, the cartilaginous parts growing more quickly.

The radials showed variations both in number and form :—

I. Variations in Number.

Smallest number

of Radials.

Largest number

of Radials.

No. of Fins examined} Average number

(right or left). of Radials.

Number of

specimens examined.

Specimens with equal

number of Radials in right

and left Fins.

(two with 15, one with 16).

Greatest difference in

;number of Radials in right

and left Fins.

(12 in the left,

16 in the right).

* Morph. Jahrbuch, Bd. xxii. 1895, p. 119.

+ Festschr. fiir C. Gegenbaur, Part I. 1896, p. 261.

t Zoologische Jahrbiicher, v. 1892 2, p. 387.

5 Trans, Zool, Soc. London, vol, xvi, pt. vil., October 1902.

294 MISS EDITH E. BAMFORD ON THE

Il. Variations in Form.—Some radials were broadened distally

(text-fig. 56 A), some had two distal epiphyses, others were

more or less deeply forked distally (text-fig. 56 B), and some

were incomplete (text-fig. 56 C). A perfect gradation was found

from the broadened to the deeply forked radial. In 67 fins

examined, it was found that 52 per cent. had either forked or

incomplete radials or both variations. Of these, 54 per cent.

had forked radials only, 23 per cent. had incomplete radials only,

and 23 per cent. had both forked and incomplete radials. The

incomplete radials were perfect distally, but had no proximal

epiphyses and did not reach to the mesopterygium, There seems

Text-fig. 56.

Radials of Polypterus.

A. Broadened Radial, showing indications of forking. B. Forked Radial.

C. Incomplete Radial.

to have been some difference of opinion concerning the kind of

variation that occurred in the radials. Wiedersheim* showed a

dividing radial, but Klaatsch found only incomplete radials in

two specimens, and he doubted the occurrence of dividing radiais ;

but that both forms are found (and frequently) is now evident.

No forked or incomplete radial is shown in Budgett’s larval

Polypterus.

Judging from the position and shape of the proximal end, an

incomplete radial in one specimen appeared to be one of the

* Lehrbuch der vergleichenden Anatomie, Erster Theil, 1882, p. 179.

PECTORAL FINS OF POLYPTERUS. 295

forks broken off from a forked radial (text-fig. 57 A), but none of

the other incomplete radials showed signs of having been derived

in this way. It seems highly improbable that the forked radials

are two radials fused proximally; the proximal epiphyses and

ends are of normal size, the peculiar variation (text-fig. 57 B)

where there is probably fusion of two radials shows that forking

can occur, for they have divided distally into three. But fusion

of radials has almost certainly been found in three fins in the

radials next to the propterygium, and these are wide proximally

as well as distally. In the table ‘‘ Fins with Forked Radials

only,” the one in position 15 seems to be of this kind.

Text-fig. 57.

| oe

Radials of Polypterus.

A. Incomplete Radial, apparently formed from one of the forks of a

forked Radial.

B. Radial, probably formed of two fused Radials divided distally

into three.

The positions of these forked and incomplete radials on the

mesopterygial edge and the relative number of the radials

occupying the different positions are of interest. In the tables

given below the radials are numbered from the metapterygium,

and 1 represents the position of the first radial on the meso-

pterygium counting from the metapterygium ; 3/4 indicates

that an incomplete radial is found between radials 3 and 4.

296 MISS EDITH E. BAMFORD ON THE

Fins with Forked Radials only.

Position of Forked Radials Number of Forked Radials De tacneaen

on Mesopterygium. found in this position. ai

1 1 4-5

2 3 13°6

5) 8 3674

4 3 13°6

5 4 18:2

6 il 4-5

12 1 4-5

15 1 4:5

Fins with Incomplete Radials only.

Position of Incomplete Number of Incomplete Pavccntas

Radials on Mesopterygium. | Radials found in this position. : Bees

3/4 A 50

4/5 2 25

6/7 2 25

In the eight fins where both forked and incomplete radials

were found this was their position on the mesopterygium :—

Forked Radials. Incomplete Radials.

TL 12 3/4

Tit, 10 5/6

TIL. 3 2/3

IV. y 4/5

7 Ve 7/8

VI. 3 5/6

Vi 5 4/5

Vil i 3/4

Tt will be seen that the third radial is most frequently forked

and that the incomplete radials most frequently occur between

radials 3,4 and 5. From this position there is a gradual decrease

in the frequency of the occurrence of forked and incomplete

radials on either side. The region where these forked and

incomplete radials ave most usually found is the distal projection

PECTORAL FINS OF POLYPTERUS. 297

of the mesopterygium, and the most frequent position 3; 3/4 is

approximately the apex of the projection.

I found no superficial indications of a former division of the

mesopterygial cartilage into a number of blocks, as in Chlamydo-

selachus and as described by Pollard *. Blood-vessels pass

through the mesopterygial foramina, but I found no nerves going

through them as indicated in Wiedersheim’s‘ illustration. The

Text-fig, 58.

1 a

GDS Vern 9 2 hy

Z Api Ht tas (\

4h UY

Fins of Polypterus.

A. Fin with forked metapterygium.

B. Fin with epiphysis of propterygium continuous with mesopterygium,

and small cartilage attached to propterygiumn.

number and arrangement of the foramina vary: from one to six

were found, of these one or two occur in the mesopterygial

ossification, the rest in the cartilaginous part. The ossified

* Zoologische Jahrbiicher, v. 1892, p. 416.

+ Lehrbuch der vergleichenden Anatomie, Erster Theil, 1882, p. 179.

Proc. Zoou. Soc.—1913, No. XX. 20

298 MISS EDITH E. BAMFORD ON THE

plate in the mesopterygium encloses numerous fat-cells, and

branches from the blood-vexsels passing through the foramina

ramify among them. The left pectoral fin of a specimen 24 em.

long had no indications of an ossified plate, the right pectoral fin

being normal. Ossification of the mesopterygium is not always

confined to the large ossified plate: I have often seen bone-cells

in the proximal cartilage between the converging propterygrum

and metapterygium, and also occasionally very small ossified

plates similar to the large one; calcification of the mesopterygial

cartilage in specimens measuring from 21 to 30 em. is frequent.

The metapterygium showed little variation; one adult Poly-

pterus had a forked metapterygium (text-fig. 58A).

Variations were frequent in the distal epiphyses of the pro-

pterygium. Klaatsch describes the fin of a young Calamoichthys

Text-fig. 59.

Propterygium of fin of Polypterus, with ossified outgrowth and cartilaginous

epiphysis in the position of the small cartilage generally found at this

place.

18 cm. long, in which the epiphysis is not separated from the

mesopterygium, and one 25 cm. long, in which it is separated.

He says he has never found this union in a Polypterus fin.

Wiedersheim’s illustration of a Polypterus fin shows a distal

cartilaginous epiphysis, and succeeding it a piece of cartilage

continuous with the mesopterygium. I have very often found

the epiphysis of the Polypterws fin continuous with the meso-

pterygium in specimens varying from 13 to 43 em. in length

(text-fig. 58, A, B), but I have never seen a distal epiphysis suc-

ceeded by cartilage continuous with the mesopterygium as shown

by Wiedersheim. The propterygium in a few cases showed

indications of a forking of the ossified distal end.

PECTORAL FINS OF POLYPTERUS. 299

I can find no explanation of the small cartilage attached to the

external side of the propterygium about two-thirds of the length

from its base. It is not shown in Budgett’s larval form, but in a

Polypterus 13 cm. long I found it similar to that in the adult.

In one specimen, 18 cm. long, there was an ossified outgrowth

een oe in a cartilaginous epiphysis at this place (text-

fig. 59).

The small distal cartilages have many variations. Usually

they alternate with the radials and do not appear to be their

terminal segments, as was thought by Gegenbaur when he tried

to establish the homology of the radials and the propterygium

Text-fig. 60.

Fin of Polypterus with proximal fusion of propterygium and metapterygium.

and metapterygium; but when a multiplication of the carti-

laginous segments occurs, as frequently happens, these additional

ones are terminal (text-fig. 58, A, B).

In a specimen 35 cm. long the propterygial and metapterygial

proximal epiphyses of the left fin were completely united; the

propterygium and metapterygium were also completely fused for

a distance of 5mm. ‘This is an interesting variation in support

of Klaatsch’s theory that the pentadactyle limb is derived from

the Polypterus type of fin, the humerus being formed from a

20*

300 MR. H. H. STIRRUP ON

fused propterygium and metapterygium*. The articulation with

the pectoral girdle was normal, and there were 17 complete and

1 incomplete radials (text-fig. 60). The fin on the right side was

’ typical, the epiphyses only being united; it had 17 radials.

Budgett + has described and illustrated a small flange of

cartilage on the-external side of the metapterygium in the larval

Polypterus, which suggests traces of a biserial arrangement of

radials on the metapterygium. He says, ‘‘On the free edge of

the metapterygium, at its distal end, isa slight flange of cartilage,

seemingly forming a rudimentary continuation round the distal

end of the radial cartilage.” I saw no traces of this cartilaginous

flange, but on removing the dermal skeleton from the fins I

found a flange in the position described, and it was apparently a

continuation of the small distal cartilages. But on examining

sections it proved to be connective tissue without any cartilage.

24. A Descriptive Study of an Oligochete Worm of the

Family Enchytreide ; with an Appendix on certain

Commensal Protozoa. By H. H. Srrrrup, B.Sc.

(Birm.), Lecturer in Agricultural Biology, Hast

Anglian Institute of Agriculture, Chelmsford f.

"Received March 10, 1913: Read March 18, 1913. ]

(Plates XLVI.-XLIX.§ and Text-figures 61-67.)

INDEX.

Pages

Enchytreus pellucidus, Anatomy of .................. 303-316

@ommensal) Protozoan pees -b-e eee eee OL OKO]

Summary of Observations... .............:.c::ceee ee 817-319

INTRODUCTION.

The object of the following paper is to give as full a description

as possible of a typical Enchytreid or ‘‘ white worm.” In spite

of the descriptions of Enchytreids given by Vejdovsky (16),

Michaelsen (8, 9, 10), Friend (5, 6), Southern (11, 12, 13), etc.,

there seems to be need for such a straightforward description,

especially of certain points about which much vagueness and

difference of opinion exists. The knowledge of our British

Enchytraids is increasing: rapidly every year, chiefly owing to

Southern in Ireland and Friend in England, but almost all

recent work in connection with Enchytreids has been purely

systematic, species new to the British Isles and also new to

science being discovered quite frequently.

* Festschr. fiir C. Gegenbaur, Part I. 1896, p. 295.

-+ Trans. Zool. Soc. London, vol. xvi. pt. vil. p. 329.

+ Communicated by Prof. F. W. GamBre, F.R.S., F.Z.S.

§ For explanation of the Plates see pp. 320-321.

156 Aja (Sq oA IS), test SIL WAL

TUT VN UA

H.H.S.del.ad nat. West,Newman lith.

ANATOMY AND PPOTOZOAL PARASITE OF FNCHYTRZID WORMS.

YS. 1918, Pl xvi

F.H.S.del.ad nat. West, Newman lith.

ANATOMY AND PROTOZOAL PARASITE OF ENCHYTRAID WORMS.

12, BS. WS}, 1 XLNWVAML,

Cob. of sep. gl.

oe.

Seala?

FLH.S.del.ad nat. West,Newman ith.

ANATOMY OF ENCHYTRAID WORMS.

IP SAS IPL, SO

av;

CUE.

soecees OVE

Tier

sed

Tantei CROTIN GE

g ie ule Hit P 3 b. J Ve d. V“

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ST TT

H.H.S.del.adnat. West, Newman lith.

ANATOMY OF ENCHYTRAID WORMS.

AN OLIGOCH ETE WORM. 301

Goodrich (7) gives a good description of the nephridia and

ecelomic corpuscles of a species of Hnchytreus, but says very little

about any other organs.

The Enchytreids are amphibious in their habits. They may

be found in comparatively dry places, as in soil or leaf-mould ;

but they are also found in certain places where they are practically

living an aquatic mode of life. A common resort of these Enchy-

treids is on the sea-shore, but when found here they are most

abundant at the point where fresh water is running into the sea.

Consequently, they are not purely land or aquatic animals, nor

yet are they purely fresh-water or marine.

The Enchytreids or ‘“‘ white worms” have been found in recent

years to be much more plentiful than was formerly supposed.

They are found most abundantly wherever there is decaying

vegetable matter, such as in leaf-mould, dead and decaying plants,

ete. But they are also found, sometimes in large quantities, at

the roots of living plants, and ultimately cause the death of these

plants. Hewitt (18) found that an Enchytreid, viz. Mridericia

bisetosa Levinsen, was the direct cause of the death of a number

of larch seedlings. It became a great pest in the nursery, killing

off the seedlings by eating away the living cortical tissue of the

main root.

I, also, have found Enchytreids at the roots of living plants.

In this case, the plant attacked was Antirrhinuwm from a garden

in Edgbaston, Birmingham. The particular Enchytreid causing

the damage was Hnchytreus argenteus Mich. As in the ease of

the larch seedlings, the Enchytreids were found at the roots,

destroying the living cortical tissue and leaving the central woody

cylinder exposed.

~ On account of this the Enchytreids are probably of great

interest from an agricultural point of view, and this paper is a

preliminary attempt to discover the economic significance of these

‘¢ white worms.”

The Enchytreid I have chosen for this paper is Enchytreus

pellucidus Friend (5), chiefly because I have an abundant supply of

these worms, but also because I think this species is a good type

of the whole genus Hnchytreus.

There is a difficulty in determining some species of the

Enchytreide, since the specific differences are sometimes very

small. After having closely examined many specimens of

Enchytreus pellucidus and also specimens of Hnchytreus albidus

Henle, which I was able to obtain in large quantities, | have come

to the conclusion that #. pellucidus cannot be regarded as a

distinct species, but only as a variety of 1. albidus, and differing

only slightly from it. According to Michaelsen (10), thirteen

authors have, in eighteen papers, given to this worm, Hnchytraus

albidus, five generic and twelve specific names, so that it 1s

evidently fairly common and probably varies in different

localities.

Two of the chief distinctions between JZ. albidus and

302 MR. H. H. STIRRUP ON

E. pellucidus lie in the structure of the brain and spermathece.

The brain of £. albidus is definitely concave behind (Pl. XLVI.

fig. 1), whilst that of ZH. pellucidus is rounded or convex behind

(fig. 2); but this is a variable character, according to the state of

tension of the worm, as I have seen the brain of £. albidus quite

straight behind with no concavity at all, and also the brain of

E. pellucidus with slight indications of a concavity behind. The

spermatheca of H. albidus has a definite rosette of glands around

the external opening of the duct (fig. 3), whilst E. pellucidus

is described as having no such glands. But the duct of the

spermatheca of H. pellucidus is irregular in outline, and in my

sections it is seen that these irregularities are due to simple

glands, which are not, however, in the form of a definite rosette

as in FZ, albidus. Again, the habitat of these two species is the

same, viz. old stable-manure and lJeaf-mould. In all other respects

they are very similar.

Short Description of the most easily observed Characters of

E. pellucidus.—It is a white transparent worm, from 15 to

20 mm. in length. The number of segments varies, but is

usually about 60. The setze are straight with the innermost ends

slightly curved, giving them the appearance of hockey-sticks ;

the number of set.is not absolutely constant, but is usually 4 in

each bundle in front and 3 behind the clitellum. The brain is

twice as long as broad when stretched to the fullest extent, and

convex behind with pigment-spots at the posterior end. ‘The

nerve-cord is equally broad throughout. Salivary glands (pepto-

nephridia) are present ; they are not branched, but simply tubular

and slightly coiled or undulating; the free ends are slightly

swollen. Blood colourless; sperm-duct very long, often extends

10 or 12 segments behind the clitellum; it is internally ciliated

very strongly along its whole length. Spermathecse in segment 5

(external opening at intersegment 4/5), communicate with ali-.

mentary canal; ampulla large, filling up the whole of segment 5;

no rosette of glands around the external opening of the duct.

Habitat.—My material of F. pellucidus was found abundantly

in a heap of refuse which consisted for the most part of leaf-

mould. In the heap were autumnal sweepings of leaves, grass cut

from the lawn, soil, stones, and occasionally bundles of soaked

brown paper. Although single worms were common throughout

the whole heap, I often came across large quantities of these

Enchytreids quite unexpectedly. When I did so, these worms

always looked bigger and healthier, and a large percentage of

them was always mature. I have always been able to get mature

worms from this heap, whatever the month of the year; this

points to the fact that they may be mature all the year round,

In the laboratory I kept the Enchytreids in tin boxes with the

leaf-mould in which they were found. They lived quite well

here, but I always found that the number of mature worms

AN OLIGOCHZTE WORM. 303

steadily decreased until, in about a month’s time, I could not find

a single mature worm. This heap of leaf-mould, which was

situated in a garden in Edgbaston, Birmingham, has been my

chief source of material. In the same heap were also numbers of

the ‘“ Gilt-tail,” Dendrobena subrubicunda ; this is interesting, as

Friend has also noticed that certain Enchytreids are often

associated with certain of the larger earthworms. A favourite

retreat of the Enchytrzids was between 2 or 3 decaying beech-

leaves that had been tightly compressed, a handful of such decaying

leaves often yielding 40 or 50 worms, many of which were quite

mature. But I have also found groups of these worms between

sheets of damp compressed paper in the heap, although one

would be surprised if they obtained much nourishment from

that. They were also fairly abundant on the under side of stones.

They were very rarely present in any lcose material, but seemed

to prefer compressed materia]. Occasionally I have seen these

Enchytrexids boring their way inside blackened decaying leaves

between the upper and lower epidermis, They were feeding on

the soft tissue of the leaf, as an examination of the contents of

the gut showed, and this suggests that Enchytraids may be quite

an important factor in the formation of leaf-mould. These

Enchytreids are also very gregarious in their habits; one often

wonders if this is due to the worms collecting together at some

point where there is a special advantage, e, g, nourishment, or

whether it is due in some manner to rapid multiplication, owing

perhaps to a similar advantage, I have isolated numbers of

specimens with a view to seeing whether Enchytrzeids are capable

of fission, but so far with a negative result,

ANATOMY,

The Pharyna, which is just behind the buecai cavity and

occupies segment 2, is interesting because of a large pharyngeal

ingrowth, This is an ingrowth of cells from the dorsal side of

the wall of the pharynx. It occupies segments 2 and 3, and is

composed of elongated spindle-shaped cells, which run dorso-

ventrally. The nuclei of these cells are large, elongated, and

distinct (Pl. XLVI. fig. 4), This pharyngeal ingrowth almost

obliterates the cavity of the gut, reducing it to a small ventral

channel. There is some connection between this ingrowth and

the septal glands, but this will be referred to later. In this

Enchytreid there is nothing in the nature of a stylet attached

to the pharyngeal ingrowth, which has been described in Enchy-

treus buchholzii Veja. (16). This stylet was said to be used for

clinging to the roots of plants. The function of the pharyngeal

ingrowth is not quite plain, but it may be some kind of sensitive

organ, as these Enchytreids, when irritated, often evert the whole

of the pharynx spasmodically, the pharyngeal ingrowth then.

hanging out like a tongue. The pharynx is extremely muscular,

the muscles running obliquely backwards from it to the body-

304 MR. H. H. STIRRUP ON

wall. These are the muscles used when the pharynx is everted.

and then withdrawn. There is another ingrowth of cells from

the ventral side, anterior to the pharyngeal ingrowth, but this is

much smaller (fig. 4).

The Esophagus gradually merges into the intestine; the actual

point where csophagus ends and intestine begins cannot be

defined.

The Intestine can be seen quite plainly owing to its darker

colour, due partly to the food in it and also to its external covering

of cells which contain many oil-globules. It is constantly under-

going peristaltic contractions, the waves running from behind

forwards. The intestine is covered over its whole length with

chloragogen-cells. When these cells break away from the gut-

wall, they are seen to be spherical, and contain numerous oil-

globules of various sizes. ‘The cells are about 16 pu in diameter.

The intestine is internally strongly ciliated. Owing to the

chloragogen-cells which cover the gut, I have never been able to

see whether the ciliary current in the intestine begins at the anus

and runs forward, or whether it runs backwards to the anus.

Stephenson (17) has recently published some interesting con-

siderations on the phenomena of antiperistalsis and ascending

ciliary action in the intestine of aquatic Oligochetes, He has

never observed ascending ciliary action from behind forwards in

the Enchytreide, but thinks that it is very possible that such an

action will be found to occur in this group. His reason for this

is that ‘‘ the inhalent function of the intestine is common in the

aquatic Oligocheta, and is evidenced by a widely occurring

ascending ciliary current in the intestine” (17, p. 74).

The problem of the nutrition of these Enchytreids is highly

important. As already mentioned, I have seen Enchytrzids

situated inside decaying leaves and literally eating their way

through the soft tissue of the leaf. When the contents of the

gut is examined, it is found to consist chiefly of disorganized

vegetable cells. These are usually brown and dead, but I recently

examined the contents of the gut of an Enchytreid which had

been feeding on the green living cells of a plant. In this case

the contents of the gut was green in colour, and a greenish fluid

was also slowly oozing out from the anus whilst the worm

was under pressure. Green vegetable cells could be recognized,

although they were being gradually disorganized. These were

the soft parenchymatous cells of the leaf or stem. In contrast

with these cells, the long vessels and tracheides of the vascular

bundles were always extruded quite whole and apparently little

changed. This would be due, no doubt, to their thick walls

resisting the action of the digestive juices.

The epithelial cells of the gut are strongly ciliated (Pl. XLVI,

figs. 4, 5). The nuclei of these cells stain deeply, and there are

present numerous large vacuoles or spaces, which often contain,

masses of material; this material has no affinity for any stains,

but is almost colourless. They suggest to me that nutrition may

AN OLIGOCHATE WORM. 305

take place, at least partly, by actual ingestion as well as by dif-

fusion of liquid nutriment. To try to prove this I have made

numerous experiments, but they were all unsuccessful. I kept

worms alive for a time in a weak solution of methylene-blue,

and then examined the cells of the gut to see if any of the

solid indigestible particles had been ingested by the cells, but

this was unsuccessful. I also tried feeding them on the scum

from a hay infusion, which contained numerous cysts, but this

was again unsuccessful. If the almost colourless material inside

the vacuoles is not actually ingested from the gut, I can only

think that it is some digestive liquid, formed by the epithelial

lining of the gut in these vacuoles, and then gradually forced into

the gut-cavity. This would mean that the vacuoles would be in

the nature of small digestive glands, Still, the fact that the cells

of the gut are so strongly ciliated, and the presence of these large

spaces in the wall of the gut filled with masses of almost colour-

less material, point at least to the possibility of nutrition being

partly by ingestion.

Salivary Glands.—The salivary glands extend as far back as

the first pair of septal glands, which are situated in segment 4.

They enter the esophagus just behind the pharyngeal ingrowth

(fig. 4). They are paired structures, each being a simple, undu-

lating, tubular gland. ‘Transverse sections show that each gland

is of almost uniform thickness, ending in a slight swelling or

ampulla. Although these two simple tubular glands converge

towards the same point just behind the pharynx, they open into

the cesophagus quite separately on the dorsal side.

The salivary glands have been called “ peptonephridia.” The

main characters which distinguish these glands from nephridia

are (1) there is no opening to the ccelom, (2) there is no trace

of a funnel, (3) the tubes are not ciliated, and in no Oligo-

chetes are there nephridia which are entirely without cilia.

On the other hand, the segments occupied by the salivary glands

are devoid of other nephbridia ; and Beddard (1, p. 47) found that

in Octochetus multiporus, the study of the development revealed

the fact that the “salivary glands” were undoubtedly formed by

the fusion of at least two pairs of nephridia, which were at

first distinct and each provided with its own coelomic funnel which

is subsequently lost. This suggests that it is quite possible that

in the earlier stages of the development of Enchytreids, the

“salivary glands” might have had an opening into the celom, and

if so, this would do away with one of the chief arguments against

the nephridial nature of these “salivary glands.” On the whole,

however, it seems best to leave the nephridial or non-nephridial

nature of the “ salivary glands” an open question.

Sete.—As before mentioned, the sete are straight with the

innermost ends curved like a hockey-stick. The sete are

slightly swollen in the middle (text-fig. 61 A). New bundles of

sete are constantly being formed by the side of the old ones.

Text-fig. 61 B shows three sete just beginning to be formed, and

306 MR. H. H. STIRRUP ON

text-fig. 61 C shows three setze almost formed with the three old

sete close beside them.

Blood-system.—Vhe intestine is surrounded by a blood-sinus

which is in very close contact with the epithelial lining of the

gut. In segment 15 the dorsal portion of this blood-sinus

becomes swollen and is gradually pinched off to form a large dorsal

vessel. The fact that this dorsal vessel is postclitellar in origin

is an important generic character distinguishing the genus

Enchytreus from most of the other genera in the family

Enchytreide. This dorsal vessel is continued forward until it

reaches segment 1. It then divides into two, each branch going

backwards to the ventral side and uniting to form the ventral

vessel. This ventral vessel continues, just ventral to the gut, to

the posterior end of the worm. Both dorsal and ventral vessels are

Text-fig. 61.

Enchytreus pellucidus.

A. Three sete. B. Three new setz just being formed,

C. Three setze almost formed with the three old setze close to them.

joined at intervals to the gut-sinus by small vessels (Pl. XLVII.

fig. 10). There are three pairs of lateral vessels also in the

anterior blood-system ; two pairs are given off close together and

join the two branches which form the ventral vessel, and one pair

is given off just behind this and joins the ventral vessel proper.

Text-fig. 62 represents this anterior blood-system.

The dorsal vessel, which is very large at its point of origin, is

partly covered with chloragogen-cells, which also extend round

the gut. At the point where the dorsal vessel arises, it swells

and contracts rhythmically, sending the blood forward. I have

seen no trace of the cardiac body, ‘‘a solid rod of cells attached to

the ventral side of the dorsal blood-vessel, and extending along

its whole length” (1, pp. 77-78), in Lnchytreus pellucidus. This

cardiac body was first described by Michaelsen in the genus

Mesenchytreus. He suggested that this solid rod of cells served

to ease the contractions of the dorsal vessel. The blood is a

colourless fluid with no blood-corpuscles.

AN OLIGOCH ETE WORM. 307

Nervous System.—The brain is dorsal in segment 1. Anteriorly

it gives off two connectives which unite ventrally to form the

ventral nerve-cord. The connectives can be traced in sections ;

they are in close approximation with the body-wall of the 1st

segment and seem to be connected at this point with a ring

of tissue which appears to be nervous. It may be this nervous

ring of tissue which makes the Ist segment, including the

prostomium, so extremely sensitive.

The brain is twice as long as broad when stretched to its

fullest extent, and oval in shape. Posteriorly it is rounded, but

occasionally shows signs of a concavity. The brain itself seems

to be composed of two parts, a central core and an outer portion

(Pl. XLVI. figs. 1& 2). The outer portion is cellular in structure ;

Text-fig. 62.

Anterior blood-system of Enchytreus pellucidus.

d.v., dorsal vessel ; v.v., ventral vessel.

the cells are small and numerous, and each one has a prominent

nucleus. In preparations, the outer portion stains very deeply

owing to the large number of nuclei. The central core is

fibrous in structure with no nuclei showing.

Pigment-spots in the Brain.—At the posterior end of the brain

there are numerous black spots in the outer cellular portion

(fig. 2). These vary in number, usually 4 to 8, and are not

regularly or symmetricaliy placed. Although these bodies have

been mentioned by some authors, I have never seen them fully

described, nor have I seen any suggestions as to their possible

function. If the brain be carefully dissected out and examined

under the high power of a microscope, the pigment-spots seem to

be dark, granular or oily, with a clear spot in the centre. They

are always present in this worm. Each black spot is composed

308 MR. H. H. STIRRUP ON °

of a number of small, rounded, highly refractive bodies. From

their appearance one would almost think they were ocelli of some

kind. However, their function is, I believe, unknown, and more

work is required upon them before they could definitely be called

ocelli. 4

In #&. pellucidus there is nothing in the nature of winged

expansions to the nerve-cord, which have been described in some

Enchytreids (6). The so-called “copulation-glands,” which is

the name given to masses of large cells which almost surround

the nerve-cord in segments 13 and 14 (14), are not present in

Enchytreus pellucidus.

The ventral nerve-cord sends out numerous branches to the

body-wall. Fig. 12 (Pl. XLVII.) shows a longitudinal section of

the body-wall passing through the nerve-cord. Here the nerve-

branches are very thick strands and can be seen passing through

the longitudinal muscles to the epidermis.

The nuclei of the nerve-cord are always situated in the ventral

region only; the rest of the nerve-cord is fibrous in structure

and has no nuclei.

In transverse sections, one almost invariably sees a number of

clear spaces, usually two or three, in the nerve-cord. These have

every appearance of being giant fibres running longitudinally

along the cord.

It would be as well to describe here some curious bodies in the

nerve-cord, which I have only seen twice in the large number of

Enchytreids I have examined. I saw them first in a small

immature Enchytreid in each of seements 8, 9,and 12. Just

recently I have again seen an exactly similar body in a fully

mature specimen of /. pellucidus; in this case it appeared in

segment 14. Under the high power they appeared to be very

granular, definite, oval bodies, quite distinct from the nerve-cord

itself, but embedded in it (Pl. XLVII. fig. 11 a, 6). I have seen

no description of these bodies elsewhere, and am inclined to think

that they were of a parasitic nature, but since I have only seen

them twice, I have been unable to do more than describe them.

Celomic Corpuscles.—The coelomic corpuscles in F. pellucidus

are comparatively fewin number. In some Enchytrzids they are

so numerous that it is very difficult to examine the various

organs in the living worm; but Z. pellucidus is so transparent

that the nephridia, brain, etc., can be seen quite easily.

My observations on the coelomic corpuscles agree very closely

with those of Goodrich in his description of H. hortensis (7).

(a) By far the commonest and largest type of corpuscle is oval

in shape and flattened (text-fig. 63.A). It varies from 47 p to

23 in length and from 12, to 8, in breadth. It is very

granular, with a clearer spot in the centre, the nucleus. It is

usually rounded at one end and at the other end drawn out

into one or two threads, the points of attachment to the ccelomic

epithelium. Whilst examining the living worm, one can always

see a number of these corpuscles attached to the walls of the

AN OLIGOCHATE WORM. 309

celomic cavity and the free ends moving about with the flow of

the ceelomic fluid. The pressure of the cover-slip on the worm

is all that is necessary to obtain these corpuscles, which are forced

out through the head-pore to relieve the pressure. The head-pore

is situated between the prostomium and the Ist segment in the

dorsal median line. Very often, also, the coelomic contents are

Text-fig. 63.

A. Celomic corpuscles of Enchytreus pellucidus.

B. Corpuscle just dividing into two.

n., nucleus ; th. of a., threads of attachment to the coelomic wall.

forced out through a pore near the anus. The presence of

this pore near the anus is not so generally known as that of the

head-pore, but it is certainly there, although rather difficult to

see. Goodrich (7) has never seen any cases of multiplication

by division of these corpuscles, but I have often observed these

corpuscles in various stages of division (text-fig. 63 B).

(6) The second type of cclomic corpuscle (text-fig. 64) is

Text-fig. 64.

Three stages in the disintegration of a ccelomic corpuscle

of Enchytreus pellucidus.

Ref.bod., refringent body ; coil.th., coiled thread ; n., nucleus.

smaller and not uniformly granular, but certainly seems, as

Goodrich suggests, to be a variety of the first kind of corpuscle.

In it there is a colourless and very refringent body, whilst the

rest of the corpuscle is granular. The curious fact is that when

310 MR. H. H, SY!IRRUP ON

this corpuscle comes into contact with a strange fluid, such as

distilled water, the colourless body swells and begins to show

that it is really formed of a long thread of transparent substance.

This long thread is very much coiled, and as it swells out the rest

of the cell disintegrates. Text-fig. 64 shows three stages in the

disintegration of one of these corpuscles. Salt solution is much

the best medium in which to examine the ccelomic corpuscles, as

they disappear very rapidly when put into distilled water.

This is a small, almost spherical corpuscle, with a number of

clear pseudopodia. Text-fig. 65 shows a number of drawings

of the same ameeboid corpuscle taken at intervals of about half a

minute.

Text-fig. 65.

Drawings of the same amceboid corpuscle ot wnchytreus pellucidus taken at

intervals of half a minute.

Pseud., pseudopodia.

In the living worm one often sees the free ceelomic contents

moving apparently through the septa from segment to segment.

I often wondered whether there were any definite apertures in

the septa through which these corpuscles could travel, and, if so,

where they were. In transverse sections, I happened to get a

section through a septum which showed quite plainly that there

were at least two definite holes in the septum. These were

situated dorso-laterally and were semicircular in shape. They

were large and distinct, and .the cclomic corpuscles were

especially crowded round the edges of the septum adjoining the

hole.

Nephridia.—Goodrich (7) has given a full account of these

organs in £. hortensis, a species closely allied to, if not identical

with, H. albidus, and I have been able to confirm his excellent

description in most particulars.

There are no nephridia in the first six segments, the first

nephridium occurring in segment 7. The nephridium is a lobed

flattened body (Pl. XLVII. fig. 13a). It is composed of three

parts, an anteseptal portion, the flattened body of the nephridium,

AN OLIGOCH ZTE WORM. 311

and the duct leading to the exterior. The anteseptal portion

is small and consists of the funnel (fig. 136). This funnel is

composed of two lips. From the extremity of the upper lip are

given off numerous very fine cilia, which wave about independ-

ently in the body-cavity. From the inner surface of the upper

lip there also arises a number of long cilia, but unlike the external

cilia these move simultaneously and rhythmically, forming an un-

dulating bunch. The function of this bunch of undulating cilia

is undoubtedly to propel liquid down the canal which leads from

the funnel. The flattened body of the nephridium consists of

granular cells, through which runs the nephridial canal. At

numerous points (usually about 4 or 5), but not continuously

through this canal, one can see bunches or “flames” of cilia

similar to the bunch in the funnel. All this ciliary action can

be observed in the living worm. The duct leading to the exterior

is broad, and is given off from the posterior end of the body of

the nephridium. The nephridiopores are arranged definitely on

a band of tissue running longitudinally along each ventro-lateral

surface, and the nephridiopore is always situated on this band

just anterior to the sete of each segment (text-fig. 66).

Text-fig. 66.

Drawing to show the position of the nephridiopores in Enchytreus pellucidus.

Sep., septum ; se¢., sete (ventral) ; 2.p., nephridiopore.

Septal Glands.—The septal glands are organs of unknown

function situated in segments 4, 5, and 6, and attached to the

posterior septum-of each of these segments. They are always

present, both in quite young Enchytreids and in old mature

ones. Inthe living worm they appear as lobed organs surrounding

the gut. Each gland consists of four lobes, which project

forwards and surround the gut, being united on the dorsal side

but not on the ventral. Fig. 14 (Pl. XLVIII.) shows a semi-

diagrammatic drawing of these organs im situ, which has been

obtained by reconstruction from a series of transverse and

longitudinal sections. The septal glands are solid organs with

no central cavity. Ihave examined numerous series of transverse

sections, but have never been able to discover any opening into

the gut. The gland itself (Pl. XLVIII. fig. 15a & 6) consists of

two parts; one part, which comprises the bulk of the gland, is

composed of solid blocks of tissue. These are remarkable for

the readiness with which they take up stains such as Hemalum,

312 MR. H. H. STIRRUP ON

Hematoxylin, etc. The other part is in the nature of a connecting-

link, and might be called the “core” of the septal glands. It

consists of two solid nucleated strands, one on each side of the

gut, which only stain slightly and connect together all the septal

glands. Now, if these connecting-strands were hollow or tubular,

one could easily understand that the septal glands were, in

reality, glands, and that the connecting-strand was a duct to

carry off their secretions. But the connecting-strands are solid,

and appear quite incapable of carrying any liquid secretion.

I have also been struck by the resemblance which these

connecting-strands bear to nervous cords or strands, but in tracing

them through series of sections I have been unable to find any

connection with the main nervous system.

In segment 4, these solid strands leave the substance of the

first septal gland and travel forwards parallel and close to the

salivary glands. They then move towards the dorsal side and

finally enter the substance of the large pharyngeal ingrowth on

the dorsal side, first dividing into four or five branches (fig. 14). °

Significance of S i

upon the development of these organs there is considerable doubt

as to their morphology, and still more as to their function.

Beddard (1), quoting Vejdovsky, shows that they are of widespread

occurrence in Oligochetes. They even occur in the early stages

of Lumbricidz, but disappear in the course of development. Up

to the present time these glands have been regarded as single

and not compound structures. The above description shows that

they contain two distinct anatomical components.

The strands connecting the septal glands with the athena:

are not muscular, and therefore it seems as if they are not

merely supporting strands. Neither are they ducts, and there-

fore, if the septal glands have a glandular function, one wonders

how this internal secretion is transmitted to its destination. I

am inclined to the view that the connecting solid strands are

nervous, and if some connection with the central nervous system

could be found, it would make the problem of solving what these

organs are, considerably easier. If the connecting-strand is an

epithelial structure, it is quite possible that it may originate as

an outgrowth from the pharynx (cf. Outgrow ths 1 in Capitellids of

Hisig), but even this will not explain the “core,” which has such

a delicate fibrillar structure. Nor cana simple septal origin te

accepted, since in that case it would be difficult to understand

the ‘‘ gland” opening into the pharynx.

Reproductive Organs.—All existing accounts of the reproductive

organs of Enchytreids are vague and incomplete. Certain of

the reproductive organs are well understood, e. g. the male

efferent apparatus consisting of the sperm-funnel and sperm-duct ;

but, on the other hand, the question of the female opening, the

oviduets (if present), and the problem as to how such large ova as

are found in Enchytreids escape by such insignificant female

pores, are still unsolved.

AN OLIGOCH ETE WORM. Sil

According to Michaelsen (9), ‘“ The reproductive apparatus of

Enchytreids varies very little. The testes are formed on the

posterior side of the septum between segments 10 and 11, and

the ovaries on dissepiment 11/12. Two sperm-funnels are present

in segment 11. The sperm-ducts lead from the sperm-funnels,

and each opens in a penis-like swelling on the ventral side ot

segment 12. A pair of oviducts open on the ventral side

of segment 13. The spermathece lie in segment 5 and open

externally on the intersegmental furrow, segments 4/5. The

clitellum covers segment 12 and often parts of segments 11

ace hoe.

Spermathece.—In Enchytreus pellucidus the spermathece are

present in segment 5. In the fully mature worm they are large

organs, filling up practically the whole of the body-cavity in that

segment. They open at intersegment 4/5.

The ampulla of the spermatheca is large and pear-shaped,

with the narrow end anterior. Ina mature worm, after copulation

has taken place, the spermatozoa can be seen through the

walls of the ampulla, giving it a definite golden-brown colour.

The duct leading to the exterior is not so long as the ampulla,

and in #. pellucidus has not gota definite rosette of glands around

the external opening. Fig. 3 (Pl. XLVI.) shows the sperma-

theca of H. albidus, with the rosette of glands round the external

aperture—it had been dissected away from the worm, and there-

fore the connection with the gut is not seen. To show how this

spermatheca differs from the spermatheca in other species, I give

a figure of the spermatheca of a mature specimen of Hnchytreus

argenteus. In this worm the ampulla of the spermatheca is

merely a slight swelling of the spermathecal tube.

Text-fig. 67.

Spermatheca of Enchytreus argenteus, for comparison with fig. 3 (PL. XLVI.)

spz., sperms in the ampulla ; amp., ampulla; op. to g., opening of

spermatheca into the gut.

The spermatheca always communicates with the gut (Pl. XLVIIJ.

fig. 16), sometimes by a long connection and sometimes by a

short one. In Z£. pellucidus the connection with the gut is

short and wide. The fact that the spermatheca communicates

Proc. Zoou, Soc.—1913, No. X XI. 21

314 MR. H. H. STIRRUP ON

with the gut has been known for some time, but it has never

been satisfactorily explained why this is so. My own observations

show that spermatozoa are occasionally seen in the tube con-

necting the ampulla with the gut, but by far the majority are

stored in the ampulla with their heads buried in the wall and

their tails free in the cavity of the ampulla. In transverse

sections, at the point where the spermatheca opens into the gut,

I have occasionally seen a number of spermatozoa actually in

the gut. Although they were quite near to the spermathecal

opening, I think this observation supports the view that this

connection between spermatheca and gut has some definite

function, probably to allow the excess spermatozoa to escape from

the spermatheca.

One can always see where the gut ends and the spermatheca

begins, because the epithelial cells of the gut are ciliated, whilst

those of the spermatheca are not.

Some time ago I had the good fortune to observe two Enchy-

treids in copulation. They were lying quite still, with the

ventral surfaces very close together, so that the spermathecal

opening in each worm was directly in touch with the male

opening of the other worm. ‘They were so tightly fixed together

that even whilst killing and fixing them they did not separate,

and so I was able to obtain sections through this region. Fig. 17

(Pl. XLVIIT.) shows a drawing of the ventral clitellar region.

From this it is seen that there is a shallow depression, overhung

on each side by a lip. Underneath each lip and in the groove

is the male opening. During copulation the fifth segment of

one worm fits closely into this shallow depression and is held in

position partly by the overhanging lip, but also probably by the

secretion of certain glands in this region, to be described later.

Fig. 19 (Pl. XLIX.) shows a drawing of a section through these

two copulating Enchytreids. The ampulla of the spermatheca

is seen on one side, whilst on the other side is the external

opening of the other spermatheca in close approximation with

the penis-like termination of the sperm-duct, which is protruding

through the male opening.

Sperm-funnels—The sperm-funnels of #. pellucidus occupy

the clitellar region. They are very long, sometimes being as

much as seven or eight times as long as broad, and are often

bent upon themselves. They lie free in the body-cavity. The

mouth of the sperm-funnel is large and has a swollen ridge,

in which are embedded the heads of numerous spermatozoa.

The tails of the spermatozoa hang out stiff and straight like

the bristles of a brush, and when viewed in the living worm

are of a beautiful golden-brown colour. The sperm-duct leads

from the sperm-funnel and is very long and coiled. It is very

strongly ciliated internally along the whole of its length. The

coils of this sperm-duct are free in the body-cavity and often

extend a dozen segments behind the clitellum before finally

ending in the male opening on segment 12.

AN OLIGOCH ETE WORM. BiH)

We now come to some gland-like bodies which are present in

the clitellar region, and which I have not seen described before

(Pl. XLIX. fig. 20). These bodies protrude into the body-cavity

in the ventral region of segment 12 on both sides of the male

openings. They are continuous with the epidermis, which in the

ventral region is composed of true epidermal cells, as the clitellar

epithelium does not extend over the ventral region. These

bodies are usually solid and composed of a few cells; they are oval

in shape and the cells have prominent nuclei. I have occasionally

seen signs of slit-lhke cavities in them, but even then there has

been nothing approaching a duct to the exterior. One can see

quite clearly that the cells forming these glands are continuous

with the epidermal cells, and they have evidently been formed as

impittings from the epidermis. The muscular system in this

region is also slightly modified. Surrounding these glands are

strong muscles, which are continued obliquely across the body-

cavity to the lateral wall. These oblique muscles are not found

anywhere else, except in the region of these glands. I think

the most probable solution of the function of these glands is that

they form some kind of secretion which helps to make the worms

adhere together closely during copulation.

Male Opening.—The actual male opening is worthy of note. The

sperm-duct ends in a peculiar chitinous-like thickening (Pl. X LIX.

fig. 21). This penis-like thickening of the sperm-duct opens to

the exterior in a groove. It does not pass through the so-called

penial bulb, but only skirts the edge of it. The penial bulb

(fig. 21) has every appearance of being simply one of these ventral

clitellar glands just described, but it is always rather bigger.

The cells composing it are continuous with the epidermis, and it

has a very strong muscular coat.

Sperm-sacs (Vesicule Seminales).—Beddard, in his ‘ Mono-

graph of the Oligocheeta,’ says: ‘‘ In only one family of Oligocheetes

are the sperm-sacs nearly always absent; this family is the

Enchytreide. The genus Mesenchytreus, however, has paired

sperm-sacs which originate from the septum bounding posteriorly

the segment in which the male gonads lie.”

In £. pellucidus there are two large sacs filling up almost the

whole of the body-cavity in segments 10 and 11. These sacs are

filled with sperm mother-cells and masses of spermatozoa in

various stages of development. Since ‘sperm-sacs’ is the name

given to sacs in which the sperm undergoes most of the stages of

its development, then these sacs in 1. pellucidus are undoubtedly

sperm-sacs, and not testes. As to the true testes, I cannot say

that I have ever seen them, but the following may have some

reference to them.

In certain Enchytreids which are not quite sexually mature,

7. e. when the ovaries and sperm-funnels are small, I have noticed

a peculiar rosette of small cells in the anterior end of each sperm-

sac. This mass of cells, when seen in the living animal, is of a

definite golden-brown colour (Pl. XLVITT. fig. 18). :

91%

316 MR. H. H. STIRRUP ON

Ovaries and Oviducts.—The ovaries are situated in segment 12.

In the mature worm the ova are so large that they almost fill

up the whole of the body-cavity in that region, and are of a

pure white colour. When ripe, they break away from the ovary

and lie apparently quite freely in the body-cavity.

In the lower Oligochetes there are no special ducts for the ova,

e. g. Holosoma simply has a median pore. Beddard (1) says that

“the structure of the oviducts in Enchytreids is such as to

suggest a degenerate condition.” D’Udekem’s opinion was that

the sperm-ducts served as ducts for the ova also. Later it was

suggested that there might simply be pores in the body-wall

through which the ova might escape to the exterior. For some

time these pores could not be found, but Claparéde (4) at last

described a pair of pores behind the male pores in Hnchytreus

vermicularis, and these were afterwards verified by Vejdovsky

and Michaelsen. However, both Vejdovsky and Michaelsen

regarded them as mere pores for the escape of the ova without

any trace of oviducts. Beddard (2) has described the oviduets in |

Pachydrilus (Enchytreide). They consist of a few pear-shaped

cells, not ciliated, which fringe the orifices. He regards them as

degenerate oviducts.

My own observations on #. pellucidus show that there is

certainly a pair of these pores in the body-wall in segment 13.

But, instead of a few non-ciliated cells fringing these openings,

it is quite plain that these two pores communicate with a single,

large, thin-walled sac, which fills up almost the whole of the body-

cavity in this region. Since the mature ova in this Enchytreeid

are large, then either the ova are capable of altering their shape

with ease or else these pores are capable of great distension. The |

former seems to be most likely, as there are no extra muscles

connected with these female pores.

APPENDIX.

Astomatous Ciliate Protozoa in the Gut.

Whilst working at the alimentary canal of this worm, I was

surprised to come across numbers of large ciliates living com-

mensally in the gut. They were usually found in the middle

region, and not at the posterior or anterior ends of the gut.

There were often so many of them present that they almost filled

the cavity of the intestine. Many of these ciliates showed

signs of budding; it was quite a common occurrence to come

across a ciliate with anything from one to six buds at the hinder

end, and the pressure of the cover-slip was only required to make

one or two of the buds break off and swim about independently.

The largest and commonest of these ciliates living in

the gut was Mesnilella fastigiata (3). The length of this

ciliate varies considerably, but an average length is °3 to

‘4mm. It is elongated and cylindrical in shape, with a

AN OLIGOCH ETE WORM. 317

curious swelling at the anterior end on one side. The most

characteristic feature is a strong chitinous-like spicule running

from one end through the body of the ciliate, and ending at about

two-thirds the length of the body in a fine curled thread, almost

like the lash of a whip (Pl. XLVI. fig. 6). The protoplasm is

granular, and there are parallel ridges running along the whole

length of the body, on which rows of cilia are constantly in

motion. The macronucleus is very long, extending the whole

length of the body, and there are numerous vacuoles present.

The micronucleus could not be seen in my own preparations,

and is, I believe, unknown. Fig. 5 shows a longitudinal section

through the gut, in which a number of these ciliates is present.

Less frequently I have found three other ciliates in the gut.

The first is shown in fig.7. It is somewhat similar to Mesnilella,

although not usually so large. It is chiefly distinguished from

Mesnilella by the absence of the spicule. ‘The ciliate shown in

fig. 7 had five buds attached.

The next ciliate was much smaller, and I have never seen any

signs of it budding. Its shape is difficult to describe, and can be

best realized from two drawings illustrating the ciliate turning

round on its short axis (Pl. XLVLII. figs. 8a & 86). It measured

90 « by 30 p. This ciliate was also without a spicule.

The last ciliate I observed was again much smaller than d/esni-

lella and of a peculiar shape. It is best described as hat-shaped

(figs. 9a & 9b). The macronucleus was long and slightly curved.

Vacuoles were present, and were situated in two longitudinal

rows. I have not been able to discover any description of the

three last-mentioned ciliates.

Summary of New and Contirmatory Observations.

(1) New Observations.

(a) The Presence and Structure of the peculiar Black Spots at

the Posterior End of the Brain.—These spots were not arranged

definitely, but were scattered about in the outer cellular portion

of the brain. When the brain was dissected out and examined

under a high power (x 1000), each of the spots was seen to be

composed of a number of small, spherical, oil-like globules.

Although at first I thought these bodies were very similar to

ocelli, I have come to the conclusion that they are probably of

the nature of very minute oil-globules. The reason for this is

that they cannot be seen in any of my preparations or sections

which have been treated with xylol or cedar-wood oil. However,

the true meaning of these black spots in the brain is by no means

clear as yet.

(b) The Structure and Significance of the so-called “ Septal

Glands.” —Hitherto these glands have been regarded as single and

not compound structures. The description given shows that

they contain two definite anatomical components. The meaning

of the connection of the septal glands with the pharynx is not

318 MR. H. H. STIRRUP ON

understood, and makes the question of the origin of the septal

glands much more complicated. The structure of the strands

connecting the septal glands with the pharynx leads one to try

and discover whether these strands are connected in some manner

with the nervous system, but my own observations prove most

conclusively that they end blindly in the substance of the

~ pharyngeal bulb.

The diagram of the septal glands (Pl. XLVIII. fig. 14), recon-

structed from series of transverse and longitudinal sections, is

useful in enabling one to understand the exact shape of these

glands and also their relation with the gut and the solid strands

connecting the glands. Most existing drawings of the septal

glands are sketches taken from the living animal, and, although

these give one a rough idea of their shape, they are never very

satisfactory.

the Haterior by two Pores in Segment 13.—Although in my prepara-

tions ova have not been actually seen in this thin-walled sac,

there can be little doubt but that it is an ovisac. The fact that it

communicates with the female pores on segment 13 proves that

ova must, at some time, be present in the sac in order to escape

to the exterior by the female pores.

(1) Zhe ventral clitellay Glands.—It seems likely that the

function of these glands is to secrete a substance which will

enable two copulating Enchytreids to be held firmly together.

My own observations on two copulating Enchytreids show that

there is certainly something which enables the two worms to

adhere very closely together.

It is difficult to account for the extraordinary muscles which

cover these glands and extend obliquely across to the body-wall.

According to anatomical evidence, the ‘‘ penial bulb” is simply

an enlarged ventral clitellar gland. It is epidermal in origin

like the ventral glands near it, and has a similar but slightly

thicker muscular coat.

These ventral clitellar glands (although it is quite likely that

they are ‘copulation-glands”) must not be confused with the

masses of large cells which almost surround the nerve-cord in

some Enchytreids (segments 13 and 14), and which have also

been called ‘ copulation-glands.”

(e) Astomatous Ciliate Protozoa in the Gut of Enchytreids.

Altogether four different kinds of these ciliates have been found

in the gut of £. pellucidus. The commonest and largest of them

was Mesnilella; this was also the only one which had the

chitinous-like rod or spicule. This ciliate has already been

recorded and described by Cépéde in his extremely useful work

on the Astomatous Infusorians (3), but I have not been able to

find any records or descriptions of the other three ciliates, so that

they may be quite new.

(f) Observations on two Copulating Enchytrceeids.—These obser-

vations on two Enchytreids in copulation are interesting, as one

“AN OLIGOCH ETE WORM. 319

rarely sees such convincing evidence as a transverse section

passing through the male opening of one Enchytreid and the

spermathecal opening of the other. The Enchytreids were

evidently killed and fixed just after the sperms had been actually

transferrea to the spermathecee by means of the sperm-duets.

_ (2) Confirmatory Observations.

After-a fairly exhaustive examination of the nephridia and

celomic corpuscles, | have been able to confirm the excellent

observations of Goodrich (7) in most particulars.

In conclusion, I must thank the Birmingham Natural History

Society for helping me in this work by means of 2 grant from

the Endowment of Research Fund, and I should also like to

express my deep indebtedness to Professor Gamble for his kindly

help and most valuable criticism.

LITERATURE.

(1) BeppArp, F. E.—A Monograph of the Order of Oligocheta.

Oxford, 1895. .

(2) Bepparp, F. E.—-‘‘ Zoological Notes. (1) On some British

Species of Pachydrilus.” Proc. Roy. Phys. Soc. Edinb. x.

pp. 101-106, pl. v.

(3) Crepe, C.—‘“ Recherches sur les Infusoires Astomes.”

Arch. Zool. exp. ser. 5, ii. p. 554, 1909-10.

(4) CrapareDE, E.—‘‘ Etudes Anatomiques sur les Annelides

Beanies Mem. Soc. Phys. Genéve, xvi. pp. 71-164,

pls. 1.-vil.

(5) Frrenp, H.—“ New British Annelids.” Zoologist, 1899.

(6) Frienp, H.—“ Irish Oligochets.” Irish Naturalist, vol. xx1.,

(7) Goopricu, EK. S.—‘ Notes on Oligochetes, with the Descrip-

tion of a new Species.” Quart. Journ. Micros. Sci., n. s.

Ol, SOmbs, IhIO.

(8) MicHartsen, W.—Untersuchungen iiber Hnchytreus mobi

Mich. und andere Enchytreiden. Kiel, 1886.

(9) MicHartsen, W.-—“‘ Synopsis der Enchytreiden,” 1889.

(1G) Micuartsey, W.—“ Oligocheta,” in Das Tierreich. Berlin,

1900.

(11) Sournery, R.—‘ Notes on the Genus Hunchytreus ... .

Irish Naturalist, vol. xv., 1906.

(12) Sournexn, R.—‘<‘ Contributions towards a Monograph of the

British and Irish Oligocheta.” Proc. Roy. Irish Acad.

vol. xxvii. sect. B, no. 8, 1909.

(13) SournerN, R.—‘‘ Oligocheta of Lambay.” Irish Naturalist,

vol. xvi., 1907.

(14) Srepuenson, J.—‘On some littoral Oligocheta of the

Clyde.” Trans. Roy. Soc. Edinb. vol. xlviii. part 1. (no. 2),

1910.

”

320 ON AN OLIGOCH ZTE WORM.

(15) Vasua, J., and Sroxuasa, J.—‘* Die Ruben-Nematoden mit

Anhang tiber die Enchytreiden,” 1896.

(16) Vespovsky, F.—*‘ Monographie der Enchytrziden,” 1879.

(17) SrepHenson, J.—‘‘ Studies on the Oligocheta of the Punjab,”

1909.

(18) Hewrrr, C. G.—“ On an Enchytreid Worm injurious to the:

Seedlings of the Larch.” Journ. Econ. Biol. vol. iii,,

1908.

EXPLANATION OF THE PLATES.

PratE XLVI.

Fig. 1. Brain of Enchytreus albidus Henle.

2. Brain of Enchytreus pellucidus Friend. cf., central fibrous portion ;

o.cell., outer cellular portion ; conn., esophageal nerve-counectives ; b/.sp.,

black spots on brain.

3. Spermatheca of Hnchytreus albidus, dissected from the worm. ext.op.,

external opening; 7os.gl., rosette of glands; amp., ampulla.

4. Longitudinal sagittal section of #. pellucidus through the first four segments.

ph.b., pharyngeal bulb or ingrowth; br., brain; sal.gl., salivary gland

opening into pharynx; m., muscles attached to pharynx ; sept.gl., septal

glands ; sp., spermatheca; es.conn., esophageal nerve-connective ; mth.,

mouth.

5. Longitudinal section of the gut, to show ciliates in the gut and structure

of gut-wall. cil., ciliates in gut; vac., vacuoles between the cells, some-

times containing a colourless mass.

6. Mesnilella fastigiata, found in the gut of H. pellucidus. mac.n., macro-

nucleus; sp., spicule; vac., vacuole; sw., characteristic swelling at one

end.

7. Ciliate with five buds, found in the gut. mac.m., macronucleus; vac.,

vacuoles,

PratE XLVII.

Fig. 8. Two views (a) and (6) of another ciliate from the gut.

9. Two views (a) and (4) of a fourth ciliate from the gut. mac.n., macro-

nucleus ; vac., vacuoles arranged in two longitudinal rows.

10. Transverse section to show the small blood-vessels connecting dorsal and

ventral vessels with the blood-sinus surrounding the gut. d.v., dorsal

vessel; v.v., ventral vessel; chl.cells, chloragogen-cells surrounding the

gut; neph., nephridium ; v.n.c., ventral nerve-cord.

11. (a) Diagram of segments 8-12, to show peculiar bodies in the nerve-cord.

(6) Nerve-cord, with one of these bodies embedded in it (much higher

power). v.v.c., ventral nerve-cord ; g7.bod., granular bodies in the nerve-

cord.

12, Longitudinal section through nerye-cord and ventral body-wall. ep., epi-

dermis ; ¢.m., circular muscles; 7.m., longitudinal muscles; 7.0., nerve-

branch; v.7.c., ventral nerve-cord; x., nuclei.

13. (a) Nephridium. (4) Nephridial funnel. n,f., nephridial funnel; ezt.e.,

external cilia; 2,c., nephridial canal; 6.c., bunch of cilia; ext.duct., duct

to the exterior; sept., septum; wp.l., upper lip; 1.1., lower lip; int.cil.,

internal bunch of cilia; ext.cél., external independent cilia.

Prate XLVILI,.

Fig. 14. Semi-diagrammatic drawing of the septal glands, obtained by reconstruction

from sections. ph., pharynx; sal.gil., salivary glands; lob. of sep.gl.,

lobe of the septal gland in segment 5; sol.st., solid strands connecting

septal glands with pharynx,

15. (a) Longitudinal sagittal section through segments 4-6, showing septal

glands: sp., spermatheca; ph.m., pharyngeal muscles; d.v., dorsal

vessel ; sept.gl., septal gland in segment 6 ; s.c., solid cord connecting the

three septal glands; sal.gl., salivary glands. (b) More magnified view of

part of the previous section: b/., solid blocks of tissue; s.c., solid cord

connecting the three septal glands.

ON SLEEPING SICKNESS. 321

Fig. 16. Transverse section through the spermatheca, showing communication with

the gut. sp.amp., spermathecal ampulla; comm.g., communication with

gut; d.v., dorsal vessel ; ; sep.gl., septal glands; v.2.c., ventral nerve-cord.

17. Drawing of the ventral clitellar region. lip, lip ov erhanging the groove in

which the male pore is situated.

18. Diagrammatic drawing of segments 10 and 11 of a young mature Enchy-

treid, showing the sperm-sacs. d.v., dorsal vessel; sp.s., sperm-sac ;

v.v., ventral vessel ; ros., rosette of cells in the sperm-sac.

Pruate XLIX.

Fig. 19. Transverse section through two copulating Enchytreids. ov., ovary ;

el.ep., clitellar epithelium ; ; sp.d., sperm-duct ; p.b., penial bulb; p., penis;

op. male opening; sp.gl., ser ae glands ; sep.gl., septal glands ;

amp., ampulla of sper matheca ; glands in the ventr al clitellar region ;

m., muscles attached to these ‘seeing glands and the body-wall; d.v.,

dorsal vessel.

20. Transverse section through segment 12 to show gland-like bodies in the

ventral clitellar region. d.v., dorsal vessel; ov., ovary; cl.ep., clitellar

epithelium ; g., gland-like bodies continuous with the epidermis; ep.,

epidermis ; v..c., ventral nerve-cord ; m., muscles connecting gland-like

bodies obliquely with the body-wall; v.v., ventral vessel; c. in g., ciliates

: in gut.

21. Highly magnified section through the male opening. sp.d., sperm-duct ; p.,

penis-like thickening ; op., male opening; p.b., penial bulb; m., muscles

around the penial bulb; g., ventral clitellar gland.

25. The Relationship of the Big Game of Africa to the spread

of Sleeping Sickness.. By Dr. W. Yorks, Liverpool

School of Tropical Medicine *, With an Appendix con-

taining Remarks by Sir Joan Bianp-Surron, F.R.C.S.,

eS. e (Crose Tete IS WSU, AIS e Jere 18) val

Mincuin, M.A., F.R.S., V.P.Z.S.; The Hon. L. WatrEer

Roruscuiup, D.Sc., F.R.S., F.Z.S.; Sir Henry Srron-

Karr, K.C.M.G., F.Z.S.; and Sir ALFRED SHARPE,

K.C.M.G., LL.D. ; and Reply by Dr. Yorke.

[Received and Read March 18, 1913. |

Although Sleeping Sickness has been recognised as a disease on

the West Coast for nearly two hundred years, human trypano-

somiasis was unknown in Nyasaland and in the greater portion

of Rhodesia until 1908. At the end of that year the first case of

the disease was found in Nyasaland, aud during 1909 and 1910 a

considerable number of cases were discovered amongst the Kuro-

peans and Natives living in Nyasaland and Rhodesia. ‘This state

of affairs was not easy to understand, as the particular tsetse fly,

Glossina palpalis, which is known to transmit Sleeping Sickness

in other parts of Tropical Africa, has not been found in these

countries.

In 1910, it was shown that the parasite causing the disease in

Nyasaland and Rhodesia differed in certain respects from that

* Communicated by Guy CHET WyND, I’.Z.S.

322 DR. W. YORKE ON

causing Sleeping Sickness in other portions of Tropical Africa.

The name Trypanosoma rhodesiense was given to this new

parasite.

Since these discoveries many cases of Sleeping Sickness have

been found in Rhodesia and Nyasaland, and in 1911 the Chartered

Company decided to have the matter thoroughly investigated ; it

was with this object that Dr. Kinghorn and I were sent to North

Eastern Rhodesia.

The first problem that we had to solve was to ascertain the

vector responsible for the spread of Sleeping Sickness in a country

where Glossina palpalis does not exist. Now although Glossina

palpalis has not been found either in Nyasaland or in the

Luangwa Valley of Rhodesia, yet Glossina morsitans, the tsetse

fly which is known to cause ‘“‘ fly” disease in domestic stock, is

present in enormous numbers, and it was soon proved by Dr.

Kinghorn and myself that it is this fly which is responsible for the

spread of human trypanosomiasis. ‘This discovery is one of great

practical significance, for whereas the former fly, Glossina palpalis,

is limited in its distribution to water-courses, and is never found

far from the banks of certain rivers or the lake shores, the latter,

Glossina morsitans, is ubiquitous, its distribution being quite

independent of water. Hence, it is at once obvious that it is

impossible to attempt to deal with Sleeping Sickness in Rhodesia

and Nyasaland by any such simple method as removing the

native population back from the water-courses and lake shores—

a procedure which was attended with such remarkable results in

Uganda. The problem of preventing the spread of Sleeping

Sickness in these countries, which a few years ago were thought

to be in no danger, is one of infinitely greater difficulty than was

that which had to be faced in Uganda.

A large number of wild Glossina morsitans was examined

in the Luangwa Valley in order to ascertain the proportion

capable of infecting man with trypanosomiasis. This information

is Important, as it affords an approximate idea of the potential

danger of the district. We found that 1 in 500 wild Glossina

morsitans was infective in nature. This is an astonishingly

large proportion, and it is at once evident that some host other

than man must be infected with the human trypanosome in order

to account for the large number of naturally infective Giossina

morsitans. With the object of ascertaining what was the chief

vertebrate reservoir of the virus, we examined a large number of

the wild fauna of Africa. In all, we examined 250 wild animals

(including elephant, rhinoceros, hippopotamus, lion, buffalo, 14

different kinds of antelope, caracal, galago, squirrel, genet, hunting

dog, giant rat, and wild rabbit), 256 monkeys, 35 domestic stock,

142 wild rats, and 15 wild mice, making a total of 698. The

results were striking, a large proportion of the antelope being

found to be infected with the parasites which cause Sleeping

Sickness in man and trypanosomiasis in domestic stock, As a

conservative estimate, the percentage of big game infected with

SLEEPING SICKNESS. 323

the trypanosomes of man or domestic stock might at Nawalia in

the Luangwa Valley be placed at 50, and at Ngoa on the Congo

Zambesi watershed at 35.

Taste I.—Percentage of various species of game found infected

with trypanosomes pathogenic to man or domestic

stock at Nawalia, Luangwa Valley.

Animal. Number examined. er eee eee es

IsBuchbucke sie sehen seehe.: 9 66°6

Waterbuck ............ asa 28 | 60°7

etre seetecta- 7 571

Hlartebeest, ii hs8-2f.-n0- 08): 6 | 16°6

OAM cee sece teen setages niu: | 8 125

Wanthogar score ncceentre renter | 9 | 111

PRUUy Saeteeerseseaaceoes tosses 10 10:0

| Mpala iS TINIE SSL sora d 29 | 6:9

| Les al)

Taste [I.—Percentage of various species of game found infected

with trypanosomes pathogenic to man or domestic

stock at Ngoa, Congo Zambesi watershed.

Animal. Number examined. Pate ae ig

| —

SHMEANADUSVIA Shs dspcoa5a0ccaqescs | 2 | 50°0

Waterbuck .. omen ean OH | | 44:4,

Bea Ce i 26°6

WD) tankers eee epee ance cece 9 222

laloantesemernetetck gett. 5 20°0

DUIS eee chee here nicotene 8 12°5

This investigation, therefore, made it perfectly clear that the

main reservoir of the trypanosomes of man and domestic stock

is the big game.

Having ascertained these two essential facts, namely that the

tsetse fly, Glossina morsitams, is the vector by which the disease

is spread, and that the big game is the inexhaustible reservoir of

the virus which causes the disease, we are faced with the problem

324 DR. W. YORKE ON

of what, in the light of this knowledge, can be done to stamp

out Sleeping Sickness or to limit its spread in Nyasaland and

Rhodesia. Obviously the most satisfactory means of prevention

would be the extermination of Glossina morsitans, which conveys

the parasite from one vertebrate host to another. Unfortunately,

however, this is out of the question at the present stage of our

knowledge. The only known method of getting rid of the fly

from a district is by clearing away the bush. In the immediate

vicinity of villages such a procedure is doubtless feasible and

would be attended by valuable results, and natives should be en-

couraged to do everything possible in this direction. The labour

involved in clearing large tracts of country would, however,

be so great that this can be at once set aside as impracticable.

Moreover, it must be remembered that not only would the

country have to be cleared, but it would require to be kept

cleared. KHveryone who has had experience of Tropical Africa is

familiar with the dense shrub growth which springs up in the

site of old garden clearings, two or three years after the natives

have ceased cultivating the land. This shrub growth is exceed-

ingly favourable to Glossina morsitans, so that unless the country

be constantly kept cleared the last state of the district is worse

than the first.

At present but little is known of the bionomics of Glossina

morsitans. The results of investigations carried out up to the

present indicate that this tsetse fly has no particular breeding

places, but that its pupee are deposited in a more or less haphazard

manner in hollow trees and excavations where they are not likely

to be disturbed by game-birds. Regarding the liability of the

fly to disease and of its natural enemies we know nothing. Much

more information is required on this subject, but it seems only

too obvious that the imvestigations will be beset by great diffi-

culties and that the information will only be forthcoming as the

result of much slow and tedious work. In fact, to those familiar

with morsitans country the extermination of the fly must seem

an almost impossible procedure.

In Uganda, where the disease is spread by Glossina palpalis,

the removal of the population a short distance away from the lake

shores and water-courses was followed by most excellent results.

Such a measure, however, is impossible in Nyasaland and Rhodesia,

where the vector Glossina morsitans is practically ubiquitous in

its distribution and not limited to water-courses as is Glossina

palpalis.

In view of the impossibility of exterminating the fly and of the

equal impossibility of removing the population from the fly belts,

we must consider the only way that remains of combating the

disease, that is the advisability of attempting to destroy the

reservoir of the virus. It is obvious that the mere isolation of

infected human beings is futile in view of the fact that the main

reservoir of the virus is the blood of the big game.

And now we come to a subject to which I particularly wish to

SLEEPING SICKNESS. 325

draw your attention—a subject which has excited considerable

controversy already, and one which will, I expect, excite still

more in the near future. I refer to the connection between big

_game on the one hand and trypanosomiasis of man and domestic

stock on the other.

Since the beginning of last year, when Dr. Kinghorn and I

published our paper announcing the fact that a large proportion

of the wild fauna of Africa harboured the trypanosomes of man

and domestic stock, a considerable polemic has arisen over the

question of the advisability of attempting to exterminate the big

game in the vicinity of human habitations.

In discussing this subject, it appears to me that I could not do

better than attempt to answer some of the objections which have

been raised against any prophylactic measure being adopted which

involves interference with the African fauna.

It has been suggested that if the big game be destroyed in

any district, the fly, being deprived of its natural source of food,

might turn its attention solely to man and his flocks and herds.

It appears to me that but little importance should be attached to

this hypothesis. In the first place, cattle do not as a rule live in

the presence of Glossina morsitans. It was suggested that cattle

and other domestic stock might harbour the human trypanosome

for considerable periods without detriment to health. This, how-

ever, is not true in the case of the human trypanosome of Nyasa-

land and Rhodesia, which we proved rapidly killed horses, cattle,

donkeys, goats, and dogs. Moreover, even if the human parasite

did not kill domestic stock, these would still die from the ordinary

cattle trypanosomes such as 7’ pecorwm, T. nanum, and 7. vivax,

with which we found the wild Glossina morsitans to be heavily

infected ; so that it 1s quite obvious that domestic stock cannot

have the same significance as a reservoir of the virus as the

antelope, which are tolerant of the trypanosomes pathogenic for

man and domestic stock. Secondly, the tsetse fly does not invade

the clearings in and around villages to any great extent, and

therefore man 1s only attacked when for any reason he goes forth

into the bush, and it is hard to believe that he would suffer to

any much greater extent in the absence of game. ‘Thirdly, and

this is the most important point: if the game were removed the

reservoir of the virus is destroyed, and therefore in a short time

the fly would tend to become non-infective. The bite of a non-

infective Glossina morsitans hurts nobody. Finally, there is

absolutely no evidence indicating that if the big game in any

particular district were slaughtered, the tsetse fly, unable to obtain

blood from these animals, would attack man and the domestic

animals to a greater extent than at present. It might equally

well be argued that if the food-supply of the fly be removed the

fly would disappear. There 1s, moreover, a considerable amount

of evidence that the tsetse fly spreads with the game. For

example, since the rinderpest swept through Central and South

Africa sixteen or seventeen years ago the big game have increased

326 DR. W. YORKE ON

enormously in numbers, and with this increase in game there has

been a corresponding increase in the number of tsetse fly. At

Nawalia, in the Luangwa Walley, where we were stationed,

Glossina morsitans was present in enormous numbers, and natives

sent out to collect the flies had no difficulty in capturing large

numbers within a short distance of the laboratory. Nawalia is

the site of an old Government station which was closed a few

years ago on account of Sleeping Sickness. The magistrate who

was stationed there in 1905 told me that he only occasionally saw

tsetse flies in this district at that time.

Again, it has been suggested that the big game might be only

one of the reservoirs of the disease, and that the infection might

equally well be conveyed by the small vermin. It must be re-

membered, however, that the small vermin are to a considerable

extent nocturnal in their habits, and although Glossina morsitans

does oceasionally bite at night, especially when the moon is full,

yet nobody who has lived in “ fly” areas can have any doubt but

that this is exceptional, and that for practical purposes the fly feeds

in the daytime only. Dr. Kinghorn and I examined a large

number of small vermin—rats, mice, wild rabbits, ete.—without

finding a single instance of natural infection. Furthermore, it

might be remarked that there is no evidence to show that the

small vermin are tolerant of the human trypanosome as are the

big game. In those which we infected experimentally the disease

ran an acute course and the animals died. If this be the case

with the majority of the small vermin, they cannot have the same

significance as reservou's of the virus as have the big game, which

can probably harbour the parasite for long periods of time with-

out exhibiting signs of disease.

We return, therefore, to the original position. The big game

is the natural reservoir of the infection, and the rélé of the tsetse

fly, Glossina morsitans, is to transfer the virus from the big game

to man and his flocks and herds. At the present state of our

knowledge we are unable to attack successfully the tsetse fly, nor,

unfortunately, is there any prospect of our being able to do so in

the near future. Whether anything would be gained in this

direction by slaughtering the big game is still a moot point: there-

fore I will not consider this side of the question, but advocate

the advisability of attempting to drive back the game from

inhabited regions solely because the game are the reservoir of

the infection.

It has been asserted that the power to slaughter all game

animals in an infected district is unsound in principle, because

the game, when harried, would betake itself to places difficult of

access to man, or scatter in small herds, or in pairs, or singly,

over wide areas, and that should this occur, it is highly probable

that it might be followed by tsetse, thus spreading the danger of

infection to wide areas now free from game and fly.

To such criticism as this it is not difficult to reply. If the

game when harassed betook itself to places difficult of access to

SLEEPING SICKNESS. 327

man, surely this is exactly what is desired. In such places it

would no longer be a menace to civilisation.

Regarding the second alternative, that the hunted game

might scatter into small herds or in pairs and be followed by

the tsetse fly, thus spreading the infection over large districts,

the obvious rejoinder is that should this occur the game must be

destroyed in the new areas, provided these happen to be inhabited

regions. Such a contingency is, however, very unlikely, as it is

very questionable if small herds of hunted game scattering over

wide areas would cause the tsetse fly to migrate with them.

The Colonial Secretary pointed out that “To talk of the

extermination of the wild fauna of a subcontinent was to talk

wild nonsense.” This is perfectly true, but it is no reason why

the game in the vicinity of human habitations should not be

destroyed. It was further stated by the Colonial Secretary that

an attempt made in Nyasaland to get the game in a certain

area killed off was, after twelve months, unsuccessful, though the

natives were encouraged to shoot. This, again, is no argument

against the policy suggested, but merely a confession that the

experiment was not efficiently performed. In this connection it

is of interest to note the remarks of Dr. J. B. Davey at a

recent meeting of the Society of Tropical Medicine. He said :—

‘““T do not think there would be much difficulty in doing this

(destroying the game) if the natives were employed. It is work

they would take up with some avidity. Having lately been a

member of the Commission in Nyasaland, I remember that con-

siderable anxiety was expressed lest we should not be able to get

sufficient material, because during the previous year fifty rifles

had been served out in that area, and the natives had been

encouraged to shoot game. If in that short time fifty rifles could

clear a large area like that, after a year or two they could make

a great impression, As a matter of fact, that permission was

only in foree for about three months, and they made a con-

siderable impression on the amount of game.”

Such experiments as these are unsatisfactory, and do not yield

any definite results. For any reliable information to be obtained,

it is necessary that the work should be done in a scientific

manner, and on a sufficiently large scale in some particular

district. A locality which is fairly well populated and which

contains plenty of tsetse fly and game should be chosen. An

exact census of the population should be made, and the propor-

tion suffering from Sleeping Sickness determined. The same

must be done in the case of the domestic animals, if such exist.

An index of the percentage of infective tsetse fly must be ascer-

tained. This is most important, as it gives one a definite idea of

the potential danger of the district. Finally, the game must be

completely eradicated, and at the same time the percentage

infected with the human and cattle trypanosome determined, and

when once the game has been driven out, it must be kept back

by vigorous action and not allowed to return. After an interval

328 DR. W. YORKE ON

of a couple of years or so the population, domestic stock, and

tsetse fly must again be carefully examined. Then we should be

in a position to decide definitely whether or not driving the fauna

back from the site of human habitations would pay. Such an

experiment as this would take some years to accomplish. That

the big game is the reservoir of the human infection there can

no longer be any doubt, as the work of Kinghorn and myself

has already been confirmed. I submit, therefore, that the time

for temporising 1s past. Sleeping Sickness has already crossed

the Zambesi, and cases have been recorded in Southern Rhodesia.

In my opinion the natives living in fly areas should be allowed

to kill game in their own way, and they might also to an extent

be armed with rifles of some uncommon bore, so that a control

could be kept over the ammunition. Huropeans ought to be

allowed to shoot what they like. Protecting the reservoir of the

trypanosomes causing fatal disease in man and his flocks and

herds by heavy licences, appears to be rather an anomaly.

Finally, I consider that some such decisive experiment as I have

outlined is urgently required, as even under the most favourable

conditions several years must elapse before we should be in a

position to recommend definitely that vigorous steps be taken to

drive back the big game from the neighbourhood of human

settlements on a large scale throughout Tropical Africa.

It may seem an act of vandalism to slaughter the wonderful

fauna of Africa; but surely when it is definitely proved that this

fauna is antagonistic to civilization, then that which stands in

the path of progress must be removed.

AGP ARAB NED 1 axe

Sir Jonn BLAND-SUTTON, F.R.C.S., F.Z.S., did not feel

qualified to express an opinion of value in regard to the drastic

scheme for extirpating the living reservoirs of Sleeping Sickness

proposed by Dr. Yorke. It is not always necessary to destroy

the reservoirs of a disease in order to protect human beings from

infection. For example, when bacteriologists discovered that the

goats of Malta, from which the island derives its milk supply,

were the reservoirs of the Jicrococcus melitensis, the cause of

Malta (or Mediterranean) fever, the Governor was strongly urged

to order the wholesale destruction of the goats. He explained

that such an order would bring about a revolution in the island,

for the Maltese are devoted to their goats. The micrococcus is

conveyed in the milk; it was a simpler plan to banish goat’s

milk from the military and naval dietary. Sailors and soldiers

at Malta are not now allowed to drink goat’s milk; as a con-

sequence, Malta fever has disappeared from the Navy and the

Army.

It is probable that the band of bacteriologists and entomolo-

gists (of which Dr. Yorke is a brillant member) engaged in

SLEEPING SICKNESS. 329

studying Sleeping Sickness will find some prophylactic or remedial

measure less obnoxious than the wholesale destruction of the

great game animals of Africa.

Me. Guy A. K. MARSHALL, F.Z.S., urged that the essential

question for consideration was whether or no the game consti-

tuted a reservoir of Trypanosoma rhodesiense, as opposed to

T. brucei; and from this point of view it was somewhat unfor-

tunate that in giving his percentages of infected game Dr. Yorke

had lumped these two forms together, as this might possibly lead

to misconceptions. He felt very strongly that the unqualified

statement that human trypanosomiasis had “spread” to south

of the Zambesi was not justified in the present state of our

knowledge, and might well cause unnecessary alarm. He agreed

with Mr. Austen that there was no evidence to show that the

disease was really spreading in the countries north of the river,

though its existence there had now been known for nearly five

years. A few years ago, when sporadic cases of Sleeping Sickness

were first discovered in Nigeria, the fear was expressed that an

epidemic similar to that in Uganda would take place. Yet no

epidemic had occurred, nor was such an event probable; for it

was now generally recognised that the disease is there endemic,

the bulk of the population being therefore immune, and these

sporadic cases merely indicated that more or less susceptible

individuals are still being born. The incidence of human try-

panosomiasis in Nyasaland and Rhodesia presents a very striking

sunilarity with that which we find in West Africa, and, in

conjunction with other considerations, strongly suggests that we

are dealing, not with a new disease, but with one which has

already been endemic for some time and has merely escaped

detection, as in West Africa. If this be a sound conclusion, we

must recognise the probability that the immune natives may

themselves be reservoirs of the trypanosome, in which case

Dr. Yorke’s assumption that the removal of the game must

necessarily eradicate the disease may prove entirely fallacious.

Finally, Mr. Marshall pointed out that Dr. Yorke had recorded that

1 in 500 of the wild flies was infected with 7’. rhodesiense, and had

referred to this as an unusually high percentage. But the latter

statement seemed hardly in accord with the results obtained by

other workers, and notably the data for 7’. cazalboni and T. gam-

biense given by Roubaud in a recent paper. Indeed, considering

the high percentage of infected game in the Luangwa Valley, as

estimated by Dr. Yorke, his record of infected flies seemed quite re-

markably low and did not suggest any probability of an epidemic.

In considering any measures for coping with human trypano-

somiasis in Rhodesia and Nyasaland, it is a matter of the very

highest importance, especially from an administrative point of

view, that we should ascertain whether the disease is really a new

one in those countries, or whether it is merely endemic. The

Proc. Zoot. Soc.—1913, No, XXII, 22

33 DR. W. YORKE ON

matter was so inadequately discussed when Dr. Yorke read his

paper to the Society that it seems desirable to consider it here in

somewhat greater detail. In his published papers Dr. Yorke

has throughout assumed the disease to be a new one, and gives

no sivn of having even contemplated the other alternative.

When, at the meeting referred to, I suggested that the disease

is probably endemic, and that in consequence immune natives

may prove to be a reservoir of the trypanosome, he rejected the

suggestion on the grounds (1) that the disease had only recently

been discovered, and it was incredible to suppose that so

distinctive an organism as a trypanosome had been previously

overlooked by medical men; and (2) that the virulence of the

disease was so great in all the cases of infection investigated

(there being no recoveries) that he found it impossible to believe

that human beings could harbour this organism with impunity.

As one who resided in Southern Rhodesia for nearly thirteen

years, I cannot seriously accept the first of these arguments.

The methods of blood-examination which are now matters of

every-day routine for any young doctor trained in our modern

Schools of Tropical Medicine, were certainly not practised in

Rhodesia until quite recently, and the chances of the disease

being correctly diagnosed, at least up to 1906, would have been

extremely remote. With regard to the second contention,

the mere fact that a disease is highly virulent in susceptible

persons is no valid evidence as to the non-existence of immune

individuals. It is now generally admitted that the natives of

West Africa are very largely immune to 7rypanosoma gambiense,

and there seems to be no good reason for assuming that a similar

power of resistance to 7’. rhodesiense cannot exist in Hast Africa.

Indeed, the more virulent the disease, the more rapid will be

the development of a general immunity; and further, the more

deadly the parasite, the less likely are there to be intergrades

between complete immunity and fatal susceptibility in the host.

Tt istrue that a new disease will usually exhibit what is known

as primary virulence; but this virulence is essentially in relation

to population, and not merely in relation to the individual. In

other words, we may reasonably assume a disease to be newly

introduced if there is a very high percentage of cases and a low

percentage of recovery; but if the percentage of cases is small

(as it certainly isin Rhodesian trypanosomiasis), the individual

severity is no proof that the disease is new. ‘To take an example.

The endemicity of yellow fever in West Africa is now hardly

disputed, yet in the epidemics in Accra and Sekondi in 1910 to

1912, out of 23 Europeans attacked no fewer than 22 died.

Another aspect is perhaps worth consideration. There can be

no question that Sleeping Sickness was a new disease in Uganda,

and there was a marked primary virulence in its true sense ; yet

in spite of this, the individual course of the malady, though

eventually fatal, was less severe and much less rapid than in

Rhodesian trypanosomiasis. ee

SLEEPING SICKNESS. 331

The position, then, is as follows. We have in Rhodesia and

Nyasaland a very virulent form of trypanosomiasis, carried by a

Glossina which has a wide local distribution and which is a much

more persistent biter than G. palpalis, and, further, a considerable

percentage of the game is estimated to be infected by J. rho-

desiense. If this be really a new disease, it would seem that the

native population is faced with an epidemic beside which that

in Uganda would be a mere circumstance, and the most drastic

remedies that youthful enthusiasm can devise would appear to be

justified.

On the other hand, we have to consider that the disease has

been known to exist in this area for nearly five years and has

probably been there a good deal longer; for we cari hardly be

asked to believe that by a remarkable coincidence Trypanosoma

rhodesiense chanced to be discovered at the precise moment of

its origin. Yet, in spite of the presence of these factors making

for the rapid development of a severe epidemic during all these

years, nothing of the kind has happened; nor have we any real

reason to suppose that an epidemic is imminent. The evidence,

such as it is, seems to indicate that we are dealing with an endemic

disease, which is not likely to become worse if the present

conditions are maintained. And while further experimental

investigations should certainly be carried on, there is no justifica-

tion for anything in the way of panic legislation.

Proressor EK. A. MINCHIN, M.A., F.R.S., F.LS., V.P.Z.S.,

said that in Dr. Yorke’s paper he had heard nothing to criticise

or controvert, so far as statements of fact were concerned. The

two African trypanosomes deadly to man, namely Trypanosoma

gambiense and T’. rhodesiense, were members of a large group of

trypanosomes which was typified by the well-known 7. brucei,

and might therefore be called the brucei-group. There could be

hardly any doubt, in the present state of knowledge, but that the

brucet-group of trypanosomes was one which was primitively

parasitic upon wild ungulates, and that many species had been

carried secondarily by the agency of biting flies to other verte-

brate hosts, in which they had been able to maintain themselves.

The fact that the trypanosomes of this group were harmless to

the wild ungulates but deadly to other animals showed that the

former were their natural hosts. Thus, 7. gambiense and 7’. rho-

desiense were harmless to antelopes but deadly to man; 7’. brucei

was harmless to wild ungulates but very deadly to domesticated

horses, cattle, or dogs. This conclusion did not apply, however,

to the human trypanosome of Brazil, 7. erwzi, nor to cattle try-

panosomes of the thedleri-type, forms which were quite distinct

from the brucei group.

While agreeing with Dr. Yorke on matters of fact, Prof. Minchin

thought that in the present state of our knowledge the utmost

caution should be exercised in putting into practice administrative

‘measures based upon the data so far established. The intev-

Dies

332 DR. W. YORKE ON

relationship of different organisms in nature was very complex,

and a sudden change brought about in the conditions might have

results altogether different to what was anticipated originally.

Dr. Yorke had argued, it seemed to him, as if the distribution

and occurrence of tsetse-flies were fixed and immutable, and

had denied that there was any danger of their migiating

towards human habitations if the big game, their natural source

of food in the bush, were destroyed. Prof. Minchin found it

difficult to believe, however, that if the tsetses in the bush were

deprived of their food, they would sit down and die of starvation ;

he thought it far more probable that the flies would migrate in

search of food, which they would find in human beings and the

cattle surrounding their habitations. [Dr. Yorke maintained

that the shelter of the bush was necessary for the flies; but such

shelter might be found on cultivated land, especially in the

banana-plantations which often surrounded native huts or

villages. |

Considering the question, therefore, purely from a utilitarian

point of view, Prof. Minchin thought it within the bounds of

possibility, to say the least, that the wholesale destruction of the

big game might lead to a condition of things more dangerous and

disastrous than that existing at present. He urged that such

measures should be undertaken, at first experimentally, on a small

scale and in a restricted area, in order that accurate knowledge

might be obtained of the effects produced by the elimination of

the wild ungulates before destroying them wholesale.

Letter from Tas Hon. L. WALTER ROTHSCHILD,

DSc. ERS, FZS:

Dear Dr. CHALMERS M1TcHELL,

IT am extremely sorry that, as I leave for Monaco on the

18th, I cannot attend the meeting. But I trust you will read

this letter at the meeting. Dr. Yorke suggests the extermination

or partial extermination of the ruminants and large game animals.

I wish to protest against this most emphatically on zoological and

ethical grounds. However, in order to prove to the utilitarians

the absolute uselessness of this proceeding, I should like to point

out that the extermination of the game animals in any large area

would hea task of several years’ duration, and the following would

take place. As year by year the large animals grew scarcer, the

tsetse-flies Glossina palpalis and morsitans, which are the means

of spreading Sleeping Sickness in men and N’gana in animals,

would be driven to bite monkeys, carnivora, rats, mice, and the

numerous small animals of those regions ; these would be infected

and the trypanosomes of the disease would gaily survive. This

would not only mean the continuance of the disease in its present

degree, but would cause a sharp increase of both diseases. The

reason for this increase, to my mind, would be very evident, for

at present, owing to the abundance of large game animals, the

SLEEPING SICKNESS. 333

flies do not on’sight attack EVERY human being or domestic animal

they perceive ; but once let them be reduced to small animals

for their normal supply of blood, it is certain that on the first

appearance of any domestic animal or human being they would

instantly precipitate themselves on it. This would mean that many

more would be bitten than at present, and many more cases of

the disease would occur. Iam sure if the game is exterminated,

the fly will not be; and, on the contrary, the disease will be

increased and not diminished,

Yours sincerely,

Zoological Museu, (Signed) WALTER ROTHSCHILD.

Herts, England.

Mareh 15th, 1913.

Letter from Siz HENRY SETON-KARR, K.C.M.G., F.Z.S.

Dear Dr. CHatmers MrrcHetn,

I am exceedingly sorry I cannot attend the meeting to-

night to hear Dr. Yorke’s paper on the relation of Big Game in

Africa in spreading Sleeping Sickness, and the discussion thereon.

The subject is one of deep interest and importance, to which I

gave considerable attention during my travels in South and East

Africa in 1911. So far as my own information and observation

go, I am entirely opposed to any proposed destruction of Big

Game, or any relaxation of the restrictions on the shooting of

game until the case against them as spreaders of Sleeping Sickness

has been fully and clearly established, which is far from being

the case at present.

The subject was fully discussed in the South-African press a

few years ago by many well-known authorities, and I would

suggest that the letters on this subject be, if possible, obtained,

printed, and circulated,

The main difficulties of the problem, as I understand them,

are :—

1. That there would appear to be other agencies, apart from

Big Game, that spread this dreadful scourge, and that

some of these agencies have yet to be ascertained.

. That inasmuch as there are Sleeping Sickness areas where

Big Game are not plentiful, and Big Game areas where

there is no Sleeping Sickness, the direct connection

between the two has yet to be fully established.

. That other eauses exist for the spread of the disease, such

as, for example, the greater freedom of intercourse

among native tribes under white protection.

4. That the destruction of Big Game in any given areas is

a task of immense difficulty, which, if attempted, might

not have the desired effect. Some game would merely

be driven elsewhere, and the smaller mammals and

game be left, which are also a possible spreading agency

of the disease.

(Se)

334 DR. W. YORKE ON

I feel sure that Dr. Yorke will initiate a most interesting and

valuable discussion, and no doubt throw new light on this im-

portant and mysterious subject. But I earnestly deprecate any

too hasty or premature a conclusion on the matter, and before all

authorities have been fully heard and consulted. If it is conclu-

sively shown that Big Game undoubtedly spread Sleeping

Sickness, then Big Game must go. On the other hand, it would

be an irreparable calamity of the worst kind to permanently

injure or destroy the indigenous wild life of Africa to no purpose,

and this might be the result of premature action.

Yours truly,

47 Chester Square, (Signed) HENRY SETON-KARR.

London, S.W.

March 18th, 1918.

Sir ALFRED SHARPE, K.C.M.G., LL.D., remarked: “‘ The

question appears to resolve itself ito the following heads :—

“1. What varieties of tsetse are proved to be carriers of the

trypanosomes of any form of Sleeping Sickness, and

(which is even more important) what, other carriers are

there ?

“2. Does tsetse really depend entirely on game for its

existence ?

“3. Assuming even that tsetse are the only distributing

agents of all forms of Sleeping Sickness—and that fly

depends solely on game for its existence—is it possible

to destroy every form of life upon which tsetse can

exist ?

“There is a desire on the part of a certain section of the

European population of our African possessions to do away with

all restrictions on the killing of game. This is fomented in some

cases by the local press; and every argument is made use of

which may further their object. In most cases those who carry

on this local agitation are ignorant of the actual conditions

governing the existence of tsetse, and its relation to game and to

Sleeping Sickness.

“1, With regard to my first heading. We know that Glossina

palpalis is considered to be the only distributing agent of

Sleeping Sickness in Uganda. G. morsitans is now credited,

however, with being a distributer in Nyasaland and N.E. Rhodesia.

What we require as regards morsitans is (a) more definite proofs

that it distributes under natural conditions, (6) more knowledge

as to whether there are not other distributing agents in Nyasaland

and Rhodesia. It is remarkable that in these two districts, in

spite of the fact that G'. morsitans exists not only in the low

country but up to a height (in Rhodesia) of 4000 feet above sea-

level, Sleeping Sickness has only been found under the same

conditions as in Uganda, 7. e. in close proximity to the banks of

the large rivers and lakes.

“2. As to the second heading, my experience is opposed to the

SLEEPING SICKNESS. 339

supposition that tsetse depends solely on game for its existence.

There are striking instances in Nyasaland and Rhodesia of game-

less districts full of fly, and flyless districts full of game.

“3. Referring to the third heading—how would it be possible

to annihilate game? It must be remembered that it is not only

what is known as ‘ big game’ that fly feeds on, but all game, big

and small, and almost every form of life in the bush. It would

be necessary to annihilate practically every form of life and to

remove also every native. ‘This would obviously be impossible.

‘“‘ Tn any case it would be a terrible mistake, which could never

afterwards be remedied, to countenance anything in the shape of

an attempt to annihilate game in any part of Africa, unless we

are absolutely certain beforehand that such a course would not only

ensure the extinction of all varieties of tsetse fly, but make sure

that all forms of Sleeping Sickness would disappear. At present

the grounds for such an assumption are totally inadequate.”

Reply by Dr. W. YORKE.

As it is impossible in the limited time at my disposal to reply

in full to the volume of criticism which my paper has evoked,

T must content myself with dealing with a few of the more

important and useful points which have been raised.

Mr. Marshall points out that in giving the percentages of

infected game I grouped together all the trypanosomes pathogenic

to man and domestic stock, and that this might lead to mis-

conception. In our published papers Dr. Kinghorn and I have

separated the parasites one from the other, and full particulars are

given as to number of each antelope infected with each of the

various pathogenic trypanosomes. I might state here that at

Nawalia in the Luangwa Valley 16 per cent., and at Ngoa on the

Congo-Zambesi watershed 3:3 per cent., of the wild animals were

infected with Trypanosoma rhodesiense. The statement that in my

published papers I have throughout assumed the disease to be a

new one in these countries, and not even contemplated the other

alternative, is incorrect. Whether the disease be new or oll

appears to me to be only of secondary importance. What is of

the first importance, however, is whether or not the disease is

spreading—whether more eases are occurring now than, say, eight

or ten years ago. I admit that from the statistics available the

question is difficult to answer; but what evidence there is suggests

strongly that during the past few years sleeping sickness has been

on the increase. Obviously the most reliable information is to

be obtained from an examination of the incidence of the disease

in Europeans. I cannot think it likely that even anterior to 1909

many white people could have succumbed from trypanosomiasis

without the disease having been recognised; yet in spite of the

fact that the main roads to the Congo-Zambesi watershed crossed

the Luangwa Valley, and that these roads were traversed by

336 DR. W. YORKE ON

large numbers of European officials and others, it was not until

towards the end of 1909 that the first white case was discovered.

Since then, although the Luangwa Valley has been closed, quite

a number of Europeans have contracted the disease in North-

Eastern Rhodesia. Moreover, we must remember that some time

before 1909 scientific experts had travelled through the Luangwa

Valley without discovering a single case either native or

European. :

The hypothesis that human beings can harbouw « s parasite

for long periods of time, as do the wild fauna, without exhibiting

signs of disease, is one which I cannot support. Amongst the

cases discovered by us, several presented practically no symptoms,

the only indication of the disease being the presence of trypano-

somes in the blood; yet without exception they were all dead

within six months.

Professor Minchin suggests that if the game in a “fly” area

were destroyed, the “fly” would enter the villages and attack

human beings and the cattle surrounding their habitations.

To this objection I have already referred in my paper. Cattle

are not as a rule found in villages situated in “ fly ” districts, and

the “fly” does not invade villages around which there is a

clearing, even though at the present time they could by so doing

obtain food still more readily than they do in the bush.

Regarding the letter of the Hon. Walter Rothschild, I have

already in my paper discussed the suggestion that after the

extermination of the game the fly would be driven to attack

monkeys, rats, and mice. Even if this occurred, these animals

quickly succumb from trypanosomiasis, and therefore cannot _

nave the same significance as reservoirs of the disease as the big

game which are tolerant of the parasites.

With regard to the first objection raised by Sir Henry Seton-

Karr, I must ask what are the other reservoirs of the virus;

and if there be any, is that a reason why the main reservoir

should not be destroyed 2? Passing to the second point, I submit

that the connection between big game and sleeping sickness has

been fully and amply established. As to the third point,

the greater freedom of intercourse among native tribes under

white protection has undoubtedly played a part in the spread of

the disease. But if civilisation and progress is to continue, this

freedom of intercourse must also continue, and the population

will of necessity run more risk of infection than previously.

Recognition of this fact makes it all the more essential that we

should endeavour to render the “ fly” as non-infective as possible

by destroying the reservoir of the virus. As the last point is

fully discussed in my paper, I need not refer to it again.

In reply to Sir Alfred Sharpe, I must point out that we have

established beyond all doubt that Glossina morsitans does transmit

sleeping sickness under natural conditions. I cannot see how

further knowledge as to whether there are other distributing

agents affects the question, beyond the fact that if this be proved

SLEEPING SICKNESS. 337

to be the case it merely emphasizes the almost hopelessness of

attempting to get rid of the transmitting agents, and demonstrates

clearly that if ‘anything is to be done it must be in the direction

of destroying the reservoir from which Glossina morsitans and

the other (hypothetical) vectors derive their infection.

As I pointed out in my paper, whether the tsetse fly depends

entirely on game for its existence is beside the question. This

subject ~ Iready been most fully discussed on more or less

theoreticat’ grounds, and we now require facts, and not further

discussion. Definite information on this, as on many other

points raised in this discussion, can only be ‘obtained by means of

the limited experiment which I advocate.

In conclusion, may I state my position once more. I have not,

as some of the speakers appear to think, made the wild statement

that the whole of the game in Tropical Africa should be destroyed,

but merely that the restrictions should be removed in “fly”

areas, and that natives and Europeans should be encouraged to

kill game in these areas, especially in the vicinity of human

habitations. Finally, I am strongly of the opinion that some

such limited experiment as I have outlined should be undertaken,

as it is only by this means that the data necessary for our

guidance in the future can be accumulated.

No. 117.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

March 4th, 1913.

Dr, A. Smrra Woopwarp, F.R.S., Vice-President, in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Dr. 8. F. Harmer, M.A., F.R.S., exhibited a hair-ball which

was presented to the British Museum (Natural History) by

Mr. A. Dobrée, in January 1911. Alluding to Mr. E. G.

Boulenger’s exhibit, on February 18th last, of spines of a Mada-

gascar Insectivore in the excrement of a Boa, he pointed out that

this was perhaps another case of the occurrence of spiny hairs in

the alimentary canal of a reptile. Mr. Dobrée’s specimen, which

has been described by Mr. R. Lydekker, F.R.S., in ‘The Field,’

vol. 117, Feb. 25, 1911, p, 383, was found in the Province of

Morondava, West Coast of Madagascar, between Beld on the

Tsiribihina River and Bevilé; and it has the form of a hair-ball,

consisting largely of the hairs of Tenrecs (Centetes). Bearing

Mr. Boulenger’s results in mind, it is possible that the specimen

in question may have come from a Boa; but it seems to be more

probable that it came from the stomach of a Crocodile. The

principal reason for coming to this conclusion is the evidence

supplied by Mr. J. Simao da Costa to the British Museum that

hair-balls of this nature occur in the stomachs of Crocodilians in

N. Brazil, as recorded by Mr. Lydekker in the note above

referred to.

Mr. E. G. Bouuenerr, F.Z.8., Curator of Reptiles, made

remarks on the Lung-Fish, Protopterus wthiopicus, which the

Society had recently received from Mr. C. W. Woodhouse, and

which was the only Dipnoan that had not previously been

exhibited alive in Europe.

* 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.

Dr. P. Coatmers Mircneny, F.R.S., F.Z.S., Secretary of the

Society, exhibited a lantern-slide made from a photograph of a

domestic Donkey (Zquus asinus) taken by Captain C. H. Armitage

in the Gold Coast, and showing a very remarkable pattern of

transversely arranged black stripes on the flank.

Dr. F. E. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the

Society, read a paper on the Anatomy and Systematic Arrange-

ment of the Cestoidea, in which he described a new genus of

Tapeworms, of the family Ichthyoteniide, from the Crossed

Viper (Lachesis alternans).

Dr. W. A. Cunnineton, M.A., F.Z.S., read a paper on the

Branchiura collected by the Third Tanganyika Expedition in

1904-1905. The collection contained over 300 specimens, and

proved that in the case of this group of animals also, Lake

Tanganyika exhibits a number of endemic forms. While two

species of Argulide are known to be widely distributed in the

lakes of Africa, they are associated in Nyasa with a single form

peculiar to that lake, but in Tanganyika with no less than seven

new types. Tanganyika is thus shown to possess not only a con-

siderable number of characteristic species, but a much richer

Branchiuran fauna than the other great African lakes. The

paper was illustrated by lantern-slides made from photcmicro-

graphs of the new species.

Mr. WitiiAM Scuaus, F.Z.8., communicated a paper containing

the descriptions of a lar ge number of new species of Rhopalocera

from Costa Rica. Over 1000 species had been collected, and of

these 04 were found to be new.

A paper was received from Dr. Arraur Wiuury, M.A.,F.BS.,

F.Z.8., containing some notes on Plankton collected across the

mouth of the St. Croix River, opposite to the Biological Station

at St. Andrews, New Brunswick, in July and August 1912.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, March 18th, 1913, at half-past Hight o’clock P.m.,

when the following communications will be made :—

EXHIBITIONS AND Noticzs.

Dr. W. Yorxe.

Remarks on the Relationship of the Big Game of Africa to

the spreading of Sleeping Sickness.

Evita E, BamFrorp.

Variations in the Skeleton of the Pectoral Fins of Polypterus.

H. H. Stirrup, B.Sc.

a a eae eer ee nent

A Descriptive Study of an Oligochete Worm of the Family

Enchytreida.,

The following papers have been received :—~

C. Tate Rucan, M.A., F.Z.S.

A Collection of Fishes made by Professor Francisco Fuentes

at Haster Island.

A Revision of the Fishes of the Genus Kuhlia,

S. F. Harmer, M.A., Sc.D, F.B.S., F.Z.8.

The Polyzoa of Waterworks,

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZooioaicaL Society or Lonpon,

Rucent’s Park, Lonpon, N.W.

March \1th, 1913.

No. 118.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

March 18th, 1913.

EK. G. B. Meapr-Watvo, Esq., Vice-President,

in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

The Secrerary read a Report on the Additions made to the

Society’s Menagerie during the month of February, 1913.

Miss Epiru E. Bamrorp presented a paper, communicated by

Dr. H. Gavow, F.R.S., F.Z.S., entitled “ Variations in the

Skeleton of the Pectoral Fins of Polypterus.” An examination

had been made of the ample material brought back by Budgett

from his West African Expeditions, in order to account for the

discrepancies which occur in the descriptions of the fins of

Polypterus as given by different investigators, These discre-

pancies were found to be due to the very numerous variations in

the fins and to the previous scarcity of material,

A description is given of the variations found in the radials,

mesopterygium, propterygium, metapterygium, and the distal

cartilages, and their bearing on the different descriptions and the

theories of other investigators is indicated.

Mr. H. H. Srirrvp, D.Sc., contributed a paper, communicated

by Prof. F. W. Gamsue, F.R.S,, F.Z.S., containing a descriptive

study of an Oligochete Worm of the family Enchytreide. A

number of new and interesting observations were recorded therein,

* 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.

including an account of the structure and significance of the

so-called ‘septal glands,” which had been found to contain two

definite anatomical components. An appendix to this paper con-

tained an account of some Astomatous ciliate Protozoa, which the

author had discovered living commensally in the gut of the

worm which formed the basis of his paper. One of these ciliates,

the largest and commonest, had been identified as Mesnilella

fastigiata, but no description had been found of three others

discovered along with it, which would probably prove to be new

species.

Dr. W. Yorxs, of the Liverpool School of Tropical Medicine,

read a paper, communicated by Mr. Guy Currwynp, F.Z.8., on

“The Relationship of the Big Game of Africa to the Spread of

Sleeping Sickness.” The author stated that sleeping sickness in

Nyasaland and Rhodesia is due to a different parasite from that

causing the disease in other parts of Tropical Africa. In these

countries the disease is transmitted by Glossina morsitans and

not by Glossina palpalis. As Glossina morsitans is ubiquitous,

and not limited in its distribution to water-courses, this fact has

an important bearing on the measures that can be recommended

with a view to prophylaxis.

The chief reservoir of the virus is the big game, the tsetse-fly

transmitting the trypanosome from the big game to man and

domestic stock,

In the consideration of prophylactic measures, the impossibility

of exterminating Glossina morsitans was mentioned, and the

removal of population from fly-areas was equally impracticable.

The advisability of attempting to destroy the game in the

inhabited portions of fly-areas was then discussed, and the

necessity of a thoroughly scientific experiment being undertaken,

so that definite information of the results of exterminating the

game in a_ particular district may be forthcoming, was

recommended.

In the discussion which followed several members and visitors

took part, including Prof. E. A. Minchin, F.R.S., Sir Alfred

Sharpe, Mr. E. E. Austen, Mr. F. C. Selous, Mr, Guy Marshall,

Sir John Bland Sutton, and Dr. 8S. F. Harmer, F.R.S., and letters

were read from the Hon. L. Walter Rothschild, D.8c., F.R.S.,

and Sir Henry Seton-Karr, C.M.G.

The general opinion of the Meeting was not in favour of

Dr. Yorke’s views.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, April 8th, 1913, at half-past Hight o’clock P.M.,

when the following communications will be made :—

Exuisitions AND Noricus.

SS SE ES

C. Tats Reaan, M.A., F.Z.S.

1. A Collection of Fishes made by Professor Francisco

Fuentes at Haster Island.

9. A Revision of the Fishes of the Genus Kwuhlia.

The Affinities of Canis antarcticus.

Major G. EH. H. Barrerr-Hamitron, ¥.Z.8., and M. A. C, Hinton.

On a ae of Mammals oa ine ese ides, Slane

The following papers have been received :—

S. F. Harmer, M.A., Se.D., F.R.S., F.Z.S.

The Polyzoa of Waterworks.

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

The Marine Fauna of British East Africa and Zanzibar,

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

in the Years 1901-1902. Bryozoa—Cheilostomata.

Communications intended for the Scientific Meetings should

be addressed to

P, CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL SocteETy oF LonDon,

Recent’s Park, Lonpox, N.W.

March 25th, 1913.

No. 119.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

April 8th, 1913.

Sir Jonn Rose Braprorp, K.C.M.G., M.D., D.Sc., F.B.S.,

Vice-President, in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The SecRETARY read a Report on the Additions made to the

Society’s Menagerie during the month of March, 1913.

Mr. H. J. Ewes, F.R.S., F.Z.8., exhibited the head of an

Asiatic Wapiti remarkable for the outward extension of the bez

tines.

Mr. C. Tare Recan, M.A., F.Z.8., read a paper on Fishes

from Easter Island collected by Professor F. Fuentes. The

collection included examples of 10 littoral species, 4 widely

distributed in the tropical Indo-Pacific and 6 new to science; of

the latter 2 were related to tropical forms and the rest to species

described from New South Wales or from Norfolk Island.

Mr. Recan also read a paper entitled ‘‘A Revision of the

Fishes of the Genus Kuhlia”; 12 species were recognized,

including 3 described as new to science.

Mr. R. I. Pococg, F.R.S., F.Z.S., Curator of Mammals, read

a paper on “The Affinities of Canis antarcticus,” in which he

* 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 Sta Shillings per annum, payable in advance.

showed :—(1) that C. antarcticus and C. latrans are not closely

related, as have been claimed; (2) that the affinities of

C. antarcticus lie with certain South-American species of Canide ;

and (3) that C.latrans must be affiliated with the wolves and

large jackals of the northern hemisphere. These conclusions

were based mainly upon cranial and dental characters, and the

points were illustrated by a series of lantern-slides of the skulls

of several species of Canide.

A paper by Major G. E. H. Barrerr-Haminroy, F.Z.8., and

Mr. Marri A. C. Hinton on a collection of Mammals from the

Inner Hebrides, was read by Mr. Hinton. This collection had

been made during an expedition organized and managed by

Mr. W. R. Ogilvie-Grant. Three new forms were discovered :

of these one (Sorex grantii) is regarded by the authors as an

insular development of S. araneus, whilst they are inclined to

think that the other two (Zvotomys alstont and Microtus agrestis

macgillivrat) are slightly modified survivals from the Pleistocene

period.

SoOREX GRANTH, sp. n.

Habitat. Islay.

Size rather large (average: head and body 77°3, tail 36, hind

foot 12°6 mm.; condylo-basal length of skull 18-2 to 19°3 mm.);

colour pattern, the dusky upper side in strong contrast to light

flanks ; dentition, posterior unicuspid above frequently suppressed,

the four remaining unicuspids being then enlarged in compen-

sation.

EvoroMYs ALSTONI, sp. n.

Habitat. Mull.

Size slightly larger than in LZ. glareolus, approaching that of

norvegicus; tail shorter than in latter, about as in former; ear

and colour nearly as in glareolws; hind foot as in norvegicus and

skomerensis (dimensions of type: head and body 108, tail 44, hind

foot 18, ear 11 mm.). Skull large as in norvegicus (condylo-basal

length 24:1-25°3 mm.), jugal heavy, brain-case broad and smooth,

convex above in profile instead of flattened, rostrum and bull

like those of norvegicus; posterior cheek-tooth above with the

third inner fold and fourth inner salient angle well developed.

MICROTUS AGRESTIS MACGILLIVRAI, subsp. n.

Habitat. Islay.

Similar to J/. a. exsul in size and general proportions ; coat

much thinner, the amount of yellowish wash on underside much

less, so that slaty bases of hairs take part in general coloration,

the upper side slightly less brightly coloured ; dental characters

conspicuously of the exsul type; skull (the comparison being

between skulls of equal age) with deeper rostrum, lighter jugal,

less sloping occiput, and less reduced interparietal, the anterior

portion of temporal muscle not ascending so high upon the

anterior shoulders of the brain-case. Broadly speaking, the

external and cranial characters of macgillivrai are those of

immature exsul, which here persist with very little modification

throughout the life of the individual.

The authors propose to divide the Western European members

of the agrestis group into two species, viz,: (1) IL agrestis,

with the subspecies agrestis, macgillivrat, exsul, levernedtiit, and

neglectus ; (2) M, hirtus, with the subspecies hirtus (Southern

Britain) and bailloni (France). J. agrestis is the older inhabitant

of the region, and the subspecies macgillivrai is regarded as its

most primitive known form; J/. hirtws is a new immigrant,

which, in the southern, lowland, or non-ingsular districts, has

replaced J. agresias.

In conclusion, the authors argue that the evidence shows that

Islay, and perhaps Jura, was separated from the old Hebridean

land-area as well as from the mainland of Scotland earlier than

were the other islands. Secondly, they think it likely that the

severance of the Hebridean district transpired before that of

the Orkneys. Lastly, they consider that the evidence of the

mammals supports the suggestion of a former direct land-con-

nection between Western Norway and the Hebrides, put forward

by Stejneger.

Mr. R. Lyprexxsr, F.R.S., F.Z.8., described, under the name

of Bubalis ceama selbornei, subsp. n., a mounted male hartebeest

from the Transvaal, in the British Museum, which differs from

the typical 6. caama by the yellowish-fawn body-colour, the

admixture of tawny hairs in the dark markings, the small extent

of the face-blaze, which is interrupted by a large tawny interval

in the region of the eyes, and by the dark markings on the front

of the lower part of the fore legs forming only a cap at the knee

and a small patch on the shank,

The next Meeting of the Society for Scientific Business will bo

held on Tuesday, April 22nd, 1913, at half-past Eight o’clock P.m.,

when the following communications will be made :—

EXHIBITIONS AND NorIcEs.

S. F. Harwer, M.A, Se.D., E.RS., F.Z8.

The Polyzoa of Waterworks,

A. W. Warmers, F.LS., F.GS.

The Marine Fauna of British East Africa and Zanzibar,

from Collections made by Cyril Crossland, M.A., B.Sc., F.Z.8.,

in the Years 1901-1902. Bryozoa—Cheilostomata.

Major J. Srevenson-Haminton, C.M.Z.S.

Notes on Albinism in the Common Reedbuck (Cervicapra

arundinum), and on the Habits and Geographical Distribution

of Sharpe’s Steenbuck (Raphiceros sharpet).

The following papers have been received :—

FrAnK EK. BeppArp, D.Sc., M.A., F.R.S., F.Z.S.

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.—X. On Two Species of Tapeworms from

Genetia dongolana.

J. A. MiILne.

Pacific Salmon: an Attempt to evolve something of their

_ History from an Examination of their Scales.

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZooLocicaL Socinry or Lonpon,

Recent’s Park, Lonpon, N.W.

April 15th, 1913,

No. 120.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

April 22nd, 1913.

EK. T. Newton, Esq., F.R.S., in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. R. H. Burne, M.A., F.Z.S., exhibited two pairs of

deformed antlers of an Axis Deer (Cervus azxis), lately presented

to the Museum of the Royal College of Surgeons by Capt. Stanley

S. Flower, F.Z.S. The deer was born in the Giza Zoological Gardens

in 1899, the deformed antlers being shed in 1905 and 1906. The

second pair showed a common malformation, 2. e. duplicity of the

brow tine, but the first pair suggested rather an injury during

growth than a congenital malformation. At a similar point in

each antler the beam was abruptly bent inwards upon itself at a

very acute angle. The apical tines also were stunted, probably

owing to an interference with the blood-supply brought about by

the sharp bend in the beam.

Mr. E. G. Bounzencsr, F.Z.S., Curator of Reptiles, exhibited a

number of living specimens of the Leaf-Insect (Phylliwm cruri-

foliwm), presented to the Society by Dr. Alfred Russell, which

had been reared from eggs laid in captivity, and which showed

various stages of development.

Dr. S. F. Harmer, F.R.S., F.Z.S., read a paper on “The

Polyzoa of Waterworks.” He gave some account of the serious

trouble which had been caused by the occurrence of a rich and

* 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,

varied fauna in the pipes of certain foreign waterworks, notably

at Hamburg and Rotterdam. As was first shown by Kraepelin,

the Polyzoa play a prominent part in the activity of the pipe-

fauna, by feeding on Diatoms and other microscopic organisms,

and serving in their turn as the food of other animals which prey

on one another, The nutritive matter rendered available by the

presence of enormous numbers of Polyzoais thus in large measure

responsible for the existence of other constituents of the fauna,

which may include even fishes, such as the Hel and the Stickle-

back, The organic material supplied by the disintegration of the

Polyzoa and other animals is believed to be important for the

nutrition of Iron-Bacteria, which are well known to cause the

most serious trouble in waterworks.

He then gave some account of five cases, which had recently

come under his own observation, of the occurrence of Polyzoa in

English waterworks in sufficient numbers to give rise to very

serious inconvenience. In one or two of these cases the advice

given by Kraepelin, in his paper on the Hamburg pipe-fauna, was

being followed, by the introduction of a system of filtration, the

principal object of which is to remove the microscopic organisms

on which the Polyzoa, and ultimately the whole assemblage of

animals in the pipes, depend for their nutriment.

The Polyzoa found in the five systems considered were referred

to four species, and some account was given of the synonymy and

distinguishing features of these. One of the species found was

Paludicella articulata Ehrb., and the specimens were remarkable

for the profusion with which hibernacula or winter-buds were

being produced. The examination of the hibernacula resulted in

the discovery of some evidence with regard to the mode of

development of these structures—a subject on which no observa-

tions appear to have been hitherto published. The evidence

recorded may have some bearing on the question whether there

is any homology between the hibernacula of Paludicella and the

statoblasts of Phylactolemata.

A paper, communicated by Mr. Cyrim CrossLanp, was received

from Mr. A. W. Waters, F.L.S., F.G.8., entitled ‘The Marine

Fauna of British East Africa and Zanzibar, from Collections

made by Cyril Crossland, M.A., B.Sc., F.Z.8., in the Years

1901-1902. Bryozoa—Cheilostomata.”

In the collection dealt with from the neighbourhood of Zanzi-

bar there are 76 species or varieties of Cheilostomatous Bryozoa,

almost all being from 10 fathoms or under, so that for a shallow-

water collection it is a very large one.

The nature of the articulation is traced in the Scrupocellaride

from the simplest to the more differentiated forms, and Cateni-

cellidee with long nodes with many ovicells are described. ‘The

ovicells at the end of erect Aetea are again described, The author

says that our position at present is a very difficult one, as it is

uncertain how much of recently proposed alteration of classification

will be accepted. He considers that, while part of Levinsen’s

classification will no doubt be accepted, there will be modification

in others, and he believes that in the future classification will be

more based upon anatomical structures. The position and nature

of the ovaria give some characters of value, but these characters

require careful study to see how far they can be made available.

The fact is again emphasized that characters in one group or

family are almost useless in the next, and all attempts at fixing

certain characters as of primary importance and others of

secondary lead to no result.

Major J. Srevenson-Haminton, C.M.Z.S., Game Warden of

the Transvaal, contributed a short paper recording the occurrence

of albino examples of the Reed-buck (Cervicapra arundinum) in the

Sabi Reserve, and containing some interesting notes on the habits

and distribution of Sharpe’s Steenbuck (Laphiceros sharpei),

which resembles the Grysbok much more closely than it resembles

the common Steenbuck in mode of life, and ranges from Nyasa-

land to the Transvaal, but gradually dies out to the south-east of

that country.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, May 6th, 1913, at half-past Hight o’clock P.m.,

when the following communications will be made :—

EXHIBITIONS AND Novices.

Frank E. Bepparp, M.A., D.Sc., F.R.S., F.Z.8.

Contributions to the Anatomy and Systematic Arrangement

of the Cestoidea.—X. On Two Species of Tapeworms from

Genetia dongolana.

J. A. MILne.

Pacific Salmon: an Attempt to evolve something of their

History from an Examination of their Scales.

KATHLEEN HADDON.

Note on Peripatoides woodwardii Bouvier.

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

Field-Observations on the Enemies of Butterflies in Ceylon.

The following papers have been received :—

Dr. R. W. Suureupt, C.M.Z.S.

On the Patella in the Phalacrocoracide.

Dr. R. Broom, C.M.Z.8.

Observations on the South African Rhynchocephaloid Reptile

Luparkeria and allied Genera.

P. Coatmers Mircnett, M.A., D.Sc., LL.D., F.R.S., F.Z.S8.

Observations on the Anatomy of the Shoe-bill (Baleniceps

7€0).

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society or Lonpon,

REGENT’s Parx, Lonpon, N.W.

April 29th, 1913.

No. 121.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON,*

May 6th, 1913.

Dr. Henry Woopwarp, F.R.S., Vice-President, in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. E. G. Boutznesr, F.Z.S., Curator of Reptiles, exhibited a

living melanistic specimen of the Green Lizard (Lacerta viridis)

recently received from Dalmatia.

A new Species of Golden Mole.

Dr. R. Broom, C.M.Z.S., exhibited an example of a new species

of Golden Mole from near Springs, Transvaal, and described it as

follows :—‘ Though closely allied to: the Natal species, Bematiscus

villosus, it differs mm having a much softer fur of a dark reddish-

brown colour, while the skull differs principally m having a tem-

poral bulla almost half as large again asin B. villosus. For the

new form the name Bematiscus transvaalensis is proposed. The

following skull-measurements are given for comparison with those

of B. villosus :—

Length. Breadth. Dental Series.

“ B. villosus, Smith’s type ......... 33 20°5 13:5 mm.

£. villosus, Dobson’s specimen... 34 22 13) mm,

DUC WSCAULEMSISIMER ee he teak on 34:9 23°5 14-5 mm.”

Dr. Broom also exhibited an adult female of the large 8. African

Lizard, Zonurus giganteus, with two newly-born young, and

three adult male specimens of the allied species Pseudocordylus

microlepidotus.

* 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 Sia Shillings per annum, payable in advance,

Dr. F. E. Bepparp, M.A., F.R.S., Prosector to the Society,

read a paper on “ The Anatomy and Systematic Arrangement of

the Cestoidea.” This paper, the tenth of the series, contained an

account of two species of ‘lapeworms found in a Dongolan Genet,

both of which were described as new, one being made the type of

a new genus.

Mr. J. A. Minye read a paper, communicated by the Secretary,

on “Pacific Salmon: an Attempt to evolve something of their

History from an Examination of their Scales,” and illustrated his

remarks with a large series of lantern-slides. Reasoning from

the similarity of their appearance to the scales of the other

Salmonide, he pointed out that all the migratory species except

Onchorhynchus kita remain for at least a year in fresh water

before proceeding to the sea—in the Fraser River district, at any

yate. He also showed the scale of a Quinnat, and pointed out

that it was hardly possible to avoid the conclusion that that fish

had already spawned once before it was captured. The ages, and

lengths of the different fishes at varying ages, were also dealt

with in the paper.

A paper, communicated by Prof. J. Sranuey Ganpiner, M.A.,

F.R.S., F.Z.S., was received from Miss Karsuren HaAppon, con-

taining some notes on Peripatoides woodwardu Bouvier.

This paper was based on material collected in Western

Australia, consisting of twenty specimens, male and female,

ranging in size from 17 to 46 mm., thus considerably exceeding

in length those described by Prof. Bouvier. Various types of

coloration are exemplified, some being blue-green with small

yellow spots, while others have the yellow pigment increased so as

to give a tawny appearance to the animal ; a dark variety of this

latter type also occurs.

Peripatoides woodwardii is characterised by the possession of

sixteen pairs of legs, and coxal glands are present in the male in

all except the 4th, 5th, and 15th pairs. The female has a re-

ceptaculum seminis on each oviduct shortly after its emergence

from the ovary.

Mr. J. C. F. Fryer, M.A., gave a brief account of some field-

observations on the enemies of butterflies in Ceylon. During a

residence of nearly two years in this island the more common

insectivorous birds were watched almost daily. Excluding the

depredations of the Wood-Swallow (Artamus fuscus Vieill.), thirty

attacks on butterflies were noted, of which thirteen were success-

ful. In five of the successes the butterfly was flying badly. Two

attacks were made on insects supposed to be relatively distasteful,

and three on the mimetic females of Papilio polytes Linn. The

W ood-Swallow lived almost entirely on butterflies and confined

itself, with few exceptions, to members of the Hupleinz and

Danaine, insects usually considered distasteful. Asilid Diptera

killed butterflies indiscriminately. It was concluded :—

i. That in Ceylon, with the exception of the Wood-Swallow,

birds are not formidable enemies to butterflies ;

il. That owing to the propensity of the Wood-Swallow for

members of the genera Danais and Huplea, a resemblance

to them would be not a safeguard but a danger.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, May 20th, 1913, at half-past Eight o’clock p.m.,

when the following communications will be made :—

EXHIBITIONS AND Novicxs.

The Rev. T. R. R. Sressine, M.A., F.BS., F.Z.8.

Notice of some important works on Zoological Nomenclature

now in progress,

Dr. R. Broom, C.M.Z.S.

Observations on the South African Rhynchocephaloid Reptile

Luparkeria and allied Genera.

E. G. Bouencsr, F.Z.S.

Experiments on the Metamorphosis of the Axolotl (Ambly-

stoma tigrinwm) conducted in the Society’s Gardens.

G. EK. Buuren,

On some cases of Blindness in Marine Fishes.

Dr. R. W. SHUFELDT, C.M.Z.S.

On the Patella in the Phalacrocoracide.

The following papers will be taken at the Scientific Meeting on

June 3rd, 1913, which closes the session :—

Sir Arruur H, Cuurca, K.C.V.O., M.A., D.Sc., F.R.S., F.S.A.

Notes on Turacin and T'uracin-bearers,

Surgeon JosepH C. Taompson, U.S.N.

Contributions to the Anatomy of the Ophidia.

Prof. T. Wingate Topp, M.B., F.R.C.S.

Observations on Osteomalacia in the Zoological Collections of

Manchester and Cleveland.

T. H. Wiruers, F.G.S.

Some Miocene Cirripedes of the Genera Hexelasma and

Scalpellum from New Zealand.

P. Caatmers Mitcuett, M.A., D.Sc., LL.D., F.RB.S., F.Z.S8.

Observations on the Anatomy of the Shoe-bill (Baleniceps

rex).

Communications intended for the Scientific Meetings should

be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZoouLoeicaL Society or Lonpon,

Recent’s Park, Lonpon, N.W.

May 13th, 1913.

Papers (continued).

Page

19. The Dwarf Buffalo of Southern Nigeria; with a Revision of the Dwarf Buffaloes of

Western Africa. By R, Lyprxker, F.R.S., F.Z.S. (Text-figs, 42-44.) ......- Satay ofa 2

20. Contributions. to the Anatomy and Systematic Arrangement of the Cestoidea.—IX. On

a New Genus of Ichthyoteniids. By Frank EK. Bepparp, M.A., D.Sc., F.R.S., F.Z.S8.,

Prosector to the Society. . (Wext-figs, 49-53.). 22.00 . 2c. seeecerssscecesscorsvers 243

21. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A.

Cunnington, 1904-1905.—Report on the Branchiura, By Wiuuiam A, Cunnrneron,

VEC token fass cbse Ne leg OLIV cor are eralpiniate os evel ais» Gjelo'e «telalnioie tia vise se serate 262

22, Notes on Plankton collected across the mouth of the St. Croix River opposite to the

Biological Station at St. Andrews, New Brunswick, in July and August 1912. By

Arruor Witury, M.A., D.Sc., F.R.S., F.Z.S., McGill University, Montreal. (Text-

ele edt OES ine at ee reek erie ti colon nae sree ate ailele seas ols «Sea eww 'are S Cake wieie e slahern 283

23, Variations in the Skeleton of the Pectoral Fins of Polypterus. By Epiru EH. Bawrorp,

Newnham College, Cambridge. (Text-figs. 56-6U.) ......sc cece secs cece eres cereee 202

24. A Descriptive Study of an Oligochexte Worm of the Family Enchytreidx; with an

Appendix on certain Commensal Protozoa. By H. H. Stirevp, B.Sc. (Birm.), Lecturer

in Agricultural Biology, East Anglian Institute of Agriculture, Chelmsford.

(Pls. XLVI.-XLIX. and Text-figs, 61-67.) 012... see ee eee eee cece eee e eee eee ees 300

25. The Relationship of the Big Game of Africa to the Spread of Sleeping Sickness. By

Dr. W. Yorks, Liverpool School of Tropical Medicine. With an Appendix containing

Remarks by Sir Jonn Buanp-Surton, F.R.C.S., F.Z.8.; Guy A. K. Marsnatt, F.Z.8. ;

Prof. HE, A. Mincnty, M.A., F.R.S., V.P.Z.S8.; The Hon. L. Water Roruscurnp, D.Sc.,

F.R.S., F.Z.S.; Sir Henry Seron-Karr, K.C.M.G., F.Z.S.; and Sir Aurrep SHArPs,

K.C.M.G., LL.D. ; and Reply by Dr. Yorne............+- i shale oretty staste a a) errata 321

Titlepage ......00.6 £50 Bib NSS GEOR ISea eS eecinn hip Hin USO IBOEN nao CO Cop Sarre OOn OC i

List of Cannel ATI CUOIMCENSHity toca crores o sieiiiate: Sie lore el otie ave eaiscalasemteis Savesane einen ataleee reas ii

Been Oe ee ibs Scag oe ood eh ae re ayes can eon Gunes iii

Alphabetical List of Contributors ..... SEA Ach anne cinerea: subse cupierarat OE ayer ix

New Generic Terms JB oS OA ES EN EES OEEIORRSD HOU OGLy Ueno ricror ROSES CA See OR Oc xvii |

Index of Scientific Names ...... ess ee cere cece ee tence A TROT RTS Ts Dae XIX

Mile xno Mla St ra GlOMaleae cas san sie eco ale lesciel a) cvaiinleiaielis smater eles ayeleleueiel's ses -eleleaelesorere rele alee cece XKVIL

— Deer

LIST OF PLATES.

1913, Parr II. (pp. 158-337).

Plate Pags

KXVI. Hehinocardwum cordatwm 0.0... c..000 cece eseceres sous 169

XXVII. 1-8. Oithona vivida. 9-12. O. fallaw .... 0+ ....ee cc aeee \

XXVIII. 1-3. Oithona fallax, 4-11. O. decipiens......++,.20-.. |

XXiX. 1-9. Paroithona pulia. 10-14. Oithona simpler ........ r 181

XXX. 1,2. Otthona simplex. 3-7. O. attenuata. 8,9. O. oculata |

XXXI. 1. Paroithona pulla. 2-9. Oithona oculata ........++.- )

XXXII. New Streptaxis and Ennea from Equatorial Africa ......

XXXIII.

XXXIV. | New Hnnea from Equatorial Africa.......... stave esterane eve 194

XXXV.

DORN. SCy1acOps CAPCNSIS: ve nie «leis alesels) oeloi wie emiete eicVaes sale sieielaietee

XXXVII. Scymnognathus hgriceps s0..8 vate Coke Ma enhen ae } 225

KA VIL, Diaptomus pusills o. «css «2c ela e einla nloiriswis a clalels jell els

RANT | Araneta SUbscilstesg: sh witeen meet DOVE BROUBMaC Boon OC [a

XL.

XLI. 1,2. Argulus incisus. 3-5. A. rubropunctatus. 6-8. A.)

PPETSONATUS aa uccrsesncrnceeccescevtresecrcerence

XLIT. 9. Argulus ewiguus. 10, 11. A. angusticeps. 12-14. A.

striatus. 15,16. A. rubescens .. ..s..eceees

XLII. 17,18. Argulus incisus. 19,20. A. lbropunetntie: “21, 22. + 262

A, €X1GUUS......- ae ts wieee sa sicise

XLIV. 23-25. Aigutae nena "96, 27. Oe ‘inaisticons See

XLY. 28-80. Argulus striatus. 31,32. A. rubescens ..........)

XLVI. |

XLvit J Anatomy and Protozoal Parasite of Enchytrzid Worms ..

300

Se \a Anatomy of Enchytreid Worms .......+...... Neer srioto c

XLIX.

NOTICE.

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,

so that the complete reference is now P. Z. 8. 1913, p.... The Distribution

is as follows:— ‘

Part I. issued in March.

paid Eee June,

3 Le September,

rae Oh ae ntes December,

‘ Proceedings,’ 1918, Part I. (pp. 1-152), were published on

March 7th, 1913.

The Abstracts of the ‘ Proceedings,’ Nos. 117—121, are

contained in wis eer