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OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON,

} ,
@ \ , j tou y

HOS) pp e337,

with 49 PuatTEs and 67 TEx1-FIGURES.

£28127

PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK.
LONDON:

MESSRS. LONGMANS, GREEN, AND CO,,
PATERNOSTER ROW.

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.,

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.).........

a2

3

lv

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

© 0

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

37

d1

63

80

85

10.

We

13.

14.

15.

16.

ie

18.

vi

. 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.

to
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

i

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

bo
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

x

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

85

bo

151

152

241

bo

51

xi
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

37

D3

63

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

bo

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

XV
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

OF

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,
23

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.

XX

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.

I

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.

XX

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,
26

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,

»)

me

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,
IB
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,
Ne
salvator, 25.
varius, 5, 7, 10, 16, 24,
34.
Vipera
ammodytes, 149.
russelli, 149, 160.

7

li,

Xanthura
luxuosus, 147.

Zaglossus
nigroaculeatus?, LA0.
Zamenis
gemonensis, 149,
mucosus, 149.
viridiflavus, 153.

i
.
ve
4
t
Bs
he
5! f4

i ocr gia é
Ale WER
OA aula

IN

DEX

OF

~[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.

De

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.,

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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

va
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
¥.

ie
=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.

el

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

1*

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
wa

ee oo
pI se oy ine
Lpiipgy yyy’
Ga ina:
aun hin Wd Gio,
ss s May ZA
< é

Sy

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. :
ie
b 5

Two transverse sections through the strobila of the same species of
Iehthyotenia (Acanthotenia).

ce

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.

od

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

28

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.

3

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

el

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.
a

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
a
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.
ee
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

x

WHIAUO) ais) Ol ish) WAAL SSO eIWASt GNI ESE SLiNshsy WSOUSHAI/ El” YL

‘cut 38 YI] [Sp Useap p

WE Neal SS UGIL Sal

Or
—"

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.
4*

2 MR. G. A. BOULENGER ON NEW AFRICAN FISHES.

OH

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

3

or

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

LS

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 ,,

79
+P

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
5*

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.
© abdomen ..... CeO aia.
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.)
1

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

39
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.

6*

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 &

c

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
7*

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.
M

‘
TT

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.

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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)

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GiLis OF ASTACOPSIS AND PARACHAIRAPS.

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a

GILLS OF ENG4US CUNICULARIUS.

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D, Bo So tess talk Sex

34

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35. ENGAZAUS HEMICIRRATULUS.

Po Ao Ss ISIS), EI, Decay

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ENGAUS CUNICUEARTUS:

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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

Qk

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
\ /
Yi
S
= :
“, =
% SY
“y \
GS &
S
\ 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.
9
0

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

BI

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.

32
29

. Engeus affinis.

33

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.

Fig.

Vig.

Fig.

30.
31.

99
3

39

39

32

39

3”

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.

0

”%

land. Companion to fig. 41.
6. From Glenore, near Hageley, Tasmania: 14.

PuatEe XXIV.

44. Engeus cunicularius, §. From Warragul, Gippsland. X14.

45

39

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,
g*

132 MR. B. F. CUMMINGS ON THE

Text-fig, 25,

Wi iry
gs mIPIRN
a Sy %

EY
ty
Hy
Es
2
=
2
Hy
Hf
z
=
:
H
3
H
4
Hy
H
H
=
FI
H
=|
S

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 :—

5

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.

i

Cc

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
e
S

no

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:
rs

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.
Malay.

S. Africa.

Malay.

N. America.
N. America.

C. America.

S. Europe.

N. America.

S. America.

India.
Gambia.

India.
India.

Madagascar.

N. Europe.

147

TYPE.
Long, thick.

Short, pointed.
Short.

Long and thin.
Long.

Long.

Long.

Long, thick.

Short, thick.
Long, thick.

Long, thick.

Short, thick.

Long, striated.

Long, thin.

Short, thick.

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.
India.
Chili.

E. Africa.
India.
India.
India.
India.
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.
India.

Europe.
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.

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.

50

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

51

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.

52
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,

2

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)
o

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.

4

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,

6

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

7

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.

8

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.

3
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.

iB

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.

2

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.

TERMS FOR THE ADMISSION OF FELLOWS.

Frtitows pay an Admission Fee of £5, and an Annual Contri-
bution of £3, due on the Ist. of January, and payable in advance,
or a Composition of £45 in lieu thereof; the whole payment,
including the Admission Fee, being £50.

No person can become a Frrrow until the Admission Fee and
first Annual Subscription have been paid, or the annual payments
have been compounded for.

Fettows elected in November and December are not liable for
the Subscription for the year in which they are elected.

PRIVILEGES OF FELLOWS.

Fettows have Personal Admission to the Gardens upon signing
their names in the book at the entrance gate, and may introduce
‘Two Companions daily.

The Wire or Huspanp of a Futzow can exercise these privileges
in the absence of the Fellow.

Until further notice, Frtznows will receive 40 undated Green
Cards, available on any Sunday or week-day up to the end of
February of the year following the year of issue, and 20 White
Cards available on any week-day up to the same date. Twenty
‘of the Green Cards may be exchanged for a book containing two
Orders for each Sunday in the year, and the twenty White Cards
may be exchanged for a similar book of Saturday Orders. Special
children’s tickets will no longer be issued, but the Green and White
Cards will be perforated, and each half will be valid for a Child
under twelve years of age. It is particularly requested that Fellows
will sign every ticket before it goes out of their possession. Unsigned
tickets are not available.

Fetrows are not allowed to pass in friends on their written
order or on presentation of their visiting cards.

Frttows have the privilege of receiving the Society’s ordinary
Publications issued during the year upon payment of an additional
Subscription of One Guinea. This Subscription is due upon the
1st. of January, and must be paid before the day of the Anniversary
Meeting, after which the privilege lapses. Fettows are likewise
entitled to purchase these Publications at 25 per cent. less than
the price charged to the public. A further reduction of 25 per
cent. is also made upon all purchases of Publications issued prior
to 1881, if above the value of F ive Pounds.

Fettows also have the privilege of subscribing to the Annual
Volume of ‘The Zoological Record, which gives a list of the Works
and Publications relating to Zoology in each year, for the sum of
One Pound Ten Shillings. Separate divisions of volumes 39 to
AQ can also be supplied. Full particulars of these publications can
be had on application to the Secretary.

Frettows may obtain a TRANSFERABLE Ivory Tickxr admitting
two persons, available throughout the whole period of F ellowship,
on payment of Ten Pounds in one sum. A second similar ticket
may be obtained on payment of a further sum of Twenty Pounds.

4

Any Fettow who intends to be absent from the United Kingdom
during the space of at least one year, may, upon giving to the
Secretary notice in writing, have his or her name placed upon the
“dormant list,” and will then be called upon to pay an annual
subscription of £1 only during such absence, but after three years
must make a further application to be retained on that list.

Any Fritow, having paid all fees due to the Society, is at liberty
to withdraw his or her name upon giving notice in writing to the
Secretary.

Ladies or Gentlemen wishing to become Fellows of the Society
are requested to communicate with ‘‘ The Secretary.”

P. CHALMERS MITCHELL,

Secretary.
Regent’s Park, London, N.W.,
June, 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

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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.

€
*
e
*%
ee

8
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

5)

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.
7:

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.

39

3)

. 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.

eI
0g

Fig.

Fig.

be)

39

mandible palp.
dorsal view.

Oithona decipiens 2, cephalon, lateral view.

FOODS TB ow po

fs

39

39

mandible palp.
Ist maxilla.
dorsal view.
Ast foot.

2nd foot.

3rd foot.

4th foot.

PratTE XXIX.

99

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
On
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.

33

23

1st antenna.
dorsal view.
8rd foot.
Ist foot.

| Oithona oculata 2, maxillipede.

1
2
3
4.
5. ”
6.
7
8
9

oO 39

oF)

4th foot.

Prarr XXXI.

Paroithona pulla 2, cephalon, lateral view.

Oithona oculata 2, dorsal view.

GOGI ENED BOE ES WS)

39

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.

23

22

marsabitensis, sp. n.
UrGFUESSENSIS, SP. Ni.
percivali, sp. n.
woodhousei, sp. n.

Ennea aberdarensis, sp. n.

39

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.

2?

33
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.

15

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.

bo
Lo
bo

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.

5
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

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PZ.S.1913. Pl. XXX

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Z OO Oe aa
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SNS ecw nema

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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
Arunella subsalsa, gen. et Sp... oe... eee eee eee seer ©2032

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

16

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
YE EOP O OHV HEN D6 © sosenccn cea son sm beaddnosocue cas PARE)
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

x

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

s
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-
form process ...... ; rubescens ©.
. 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.

Lo

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

e

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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Cope pod aig sasesanssce tcctagonc teas oeemerats onsen eee ae neue) SAA
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.

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00
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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.

19

12

Number of

specimens examined.

25

Specimens with equal
number of Radials in right
and left Fins.

3

(two with 15, one with 16).

Greatest difference in

;number of Radials in right

and left Fins.

4

(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.

Gy
is

| 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.

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ANATOMY AND PPOTOZOAL PARASITE OF FNCHYTRZID WORMS.

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ANATOMY AND PROTOZOAL PARASITE OF ENCHYTRAID WORMS.

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ANATOMY OF ENCHYTRAID WORMS.

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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.

ag

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.

(c) The third type of corpuscle is the ameboid corpuscle.
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.

(c) Presence of a thin-walled Ovisac which comnunicates with
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.

tw

(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 ?

b

“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

Dy

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.

10

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.

1]
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.

14

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.

15

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.

18

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

19

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,

yo
22

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

23

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.

24

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,

26

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

27

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,

28

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

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